1

BUD DORMANCY

D ormancy can be defined as a developmental process involving a temporary

suspension of visible growth of any plant structure containing a meristem. It
is a survival mechanism assuring a seasonal synchronization of growth, but is also
contributing to the control of plant architecture. Dormancy may be imposed by
certain external, internal or genetic factors. In order to overcome or survive against
adverse environmental conditions plants undergo a period of dormancy with suitable
modifications. Lower plants avoid these unfavourable conditions through the
production of endosperms, zygospores, auxospores, akinetes, etc. The vascular
plants produce dormant structures to overcome such unfavourable conditions.
Among the many structures which exhibit dormancy, seeds and buds are important.
Perennial plants like shrubs and trees have to go through different seasons in a
year. Because of extreme variations in the temperature, especially cold conditions
during the growth and even survival of plants become difficult. In order to survive
against such adverse conditions the growth regions like apical buds, axillary buds
and underground structures like rhizomes, tubers, etc. undergo a period of dormancy
i.e suspended growth. Leaves wither during the season not due to change in the
temperature but due to change in the photoperiodic conditions.There is temporary
suspension of growth in meristematic cells also and these are protected by thick,
hairy and waxy bud scales which provide thermal insulation to the meristematic
zones and prevent them from cold or frost injury.

DEFINITIONS
Various researchers have defined bud dormancy in different ways as:
● Dormancy is the state of reduced activity of development.
● Any rest period or reversible interruption of the phenotypic development of
any organ.
2 Fruit Tree Physiology

● A period of markedly reduced growth rate with few or in some cases no cell
division in the terminal or lateral meristem.
● A condition in which a tissue is pre disposed to elongate does not do so
(Dorenbos, 1953).
● A temporary suspension of visible growth of any plant structure containing
meristem (Lang, 1987).
● When the mitotic activity in the cells of buds is zero - a condition occurring
from December through February (Owens and Molder, 1973).
● The inability to initiate growth from meristems (and other organs and cells
with the capacity to resume growth) under favourable conditions.
Ecodormancy state is not covered by this definition (Rohde and Bhalerao,
2007).

CLASSIFICATION
Lang et al. (1987) classified dormancy into following categories :
Paradormancy (Ectodormancy) : The inhibition of growth by factors outside the
bud (distal organs).
Endodormancy : The inhibition of growth by factors internal to bud; and
Ecodormancy : The inhibition of growth by unfavourable environmental conditions.
Paradormancy : This phenomenon is regulated by the factors which are
within the plant but are external to the dormant structure e.g apical dominance or
photoperiod in certain cases. It is synonymous with correlated dormancy and
summer dormancy. Basipetal transported auxin has been observed as the primary
signal that regulates paradormancy by many workers.
Endodormancy : The physiological changes internal to the bud that prevent
untimely growth during seasonal transitions, when environmental conditions often
fluctuate between those permissive or inhibitory to growth lead to endodormancy.
It helps to protect the vegetative buds by suspending the growth of meristems
until return of suitable environmental conditions. Induction and breaking of
endodormancy is regulated by light and temperature. Light plays a dominant role
in woody plants. In deciduous plants, terminal buds produce leaf primodium
forming scales instead of leaf buds due to short-day conditions (Okuba, 2000).
Ecodormancy : External environmental factors such as cold or drought stress
that prevent bud growth impose ecodormancy. These environmentally unfavourable
growing conditions are generally required for the breaking of endodormancy, but
at the same time imposing ecodormancy.
 Bud Dormancy 3

(Horvath et al; 2003)

Fig. : Diagram of signals and typical seasons corresponding to the different types
of dormancy. Lang et al; 1987 categorized dormancy into paradormancy,
endo dormancy and ecodormancy. This figure illustrates the cycle of three
types of dormancy and the general signals and seasons that are associated
with dormancy of perennial woody plants (shrubs and trees).

Hypothetical dormancy classification system

Organisation level Term for event

I. Base condition: The temporary suspension of visible Dormancy

growth of any plant structure containing a meristem.
II. Initial regulation by (process):
- Physiological factors within the dormant structure. Endodormancy
- Physiological factors within the plant, outside Paradormancy
the dormant structure.
- Environmental stresses evoking non specific Ecodormancy
responses.
III. Controlling factors (inputs) :

[Table Contd...
4 Fruit Tree Physiology

Contd. Table]

Organisation level Term for event

A. Endodormancy
Chilling Cryogenic endodormancy
Photoperiod Photoperiodic endodormancy
Light quality Spectral endodormancy
Endogenous rhythms Rhythmic endodormancy
B. Paradormancy
Apical meristem Apical paradormancy
Subtending leaves, budscales or cotyledons Laminar paradormancy
Seedcoat Testa paradormancy
Periderm Peridermal paradormancy
Chilling Cryogenic paradormancy
Photoperiod Photoperiodic paradormancy
Light quality Spectral paradormancy
C. Ecodormancy
Temperature Thermal ecodormancy
Water Hydrational ecodormancy
Nutrient Nutritional ecodormancy
Level of CO2, O2 Atmospheric ecodormancy

(Lang et al; 1987)

A new concept of dormancy was put forth by Rhode and Bhalerao (2007).
According to them dormancy is the ability to cease meristem activity and to
establish a dormant state in which the meristem is rendered insensitive to growth
promoting signals for some time before it is released and can resume growth. They
identified different stages of growth in plants which are as under (Table and Fig.):
Table : The growth–dormancy status and the corresponding meristem stages.

Stage Description

I Cessation of cell division

II Establishment of dormancy
III Maintenance of dormancy
IV Release from dormancy state
V Resumption of cell division

Locus and Site of Dormancy Perception

The locus of dormancy is the apical meristems of shoots. Buds act as the sites
of perception for inducing dormancy, although leaves perceive changes in
photoperiodic effects. Dormancy is induced in buds only after the falling of
leaves and subjecting them to long photoperiodic treatment or interrupting the
long dark periods by red light leads to breaking of dormancy.
 Bud Dormancy 5

(Rhode and Bhalerao; 2007)

Fig. : Transitions in seasonal growth–dormancy cycling in woody perennials. The
inner circles depict the growth–dormancy status and the corresponding
meristem stages: I- cessation of cell division; II- establishment of dormancy;
III- maintenance of dormancy; IV- release from dormancy; and V- resumption
of cell division. Gap between stage IV and V denotes the phase where growth
does not occur because of purely environmental restraints

Physiological Mechanism of Bud Dormancy

It is generally recognised that dormancy is induced by the onset of low temperatures
and shorter day-length associated with late autumn. The metabolic rate of the
plant decreases to such an extent, that the residual metabolism is just sufficient
for the plant to survive. Growth ceases for the period of dormancy. During
dormancy, the level of endogenous plant growth inhibitors is maintained at a high
level in the dormant organs. It is still unclear whether the increase in the
concentration of such substances is a pre requisite for the induction and/ or the
continuation of the dormant stage. Dormancy involves a temporary suspension of
visible growth of plant structures containing a meristem. Dormancy is an important
mechanism ensuring a seasonal synchronization of growth and also contributes to
the control of plant architecture. Establishment of the dormant condition in time
before the unfavourable season requires sensing and processing of a regular and
reliable environmental seasonal signal. In temperate trees and shrubs a short-day
6 Fruit Tree Physiology

control of growth cessation and dormancy induction has been widely demonstrated.
Physiological studies, including recent studies of the effects of monochromatic
red, far-red and blue light on growth and bud formation, have pointed to an
important role of the phytochrome system as the day length sensor might suggest
also a role of one or more blue light receptors. Studies of woody plants with
changed expression of phytochromes might suggest that perception of photoperiod
is related to levels of phytochromes. Also, recent studies have shown that steady
state mRNA levels of different phytochromes in woody plants are affected by day
length. The phytochrome system apparently interacts with biosynthesis of plant
hormones, particularly gibberellin. Also, the involvement of abscisic acid in control
of dormancy related processes has been shown. Dormancy induction is associated
with reduced rates of cell division, and it appears that these hormones act, at least
partly, through their interaction with the mechanisms of cell cycle regulation. In
some species, like apple and pear, no effect of photoperiod on growth cessation
and dormancy induction has been found, whereas low temperature is highly
effective. However, it is a paradox that the same temperature regime that induces
dormancy in these species also controls its release. Moreover, in a number of
species in which growth cessation occurs under short photoperiod, a low night
temperature can bring about growth cessation and bud set even under long days.
Bud set under a non inductive photoperiod has been shown even in day length
insensitive plants of phytochrome A overexpressing Populus when exposed to a
low night temperature. These and other studies indicate that phytochrome action
is affected by temperature, and is suggestive of the existence of a photoperiod
independent pathway in addition to a photoperiodic control resulting in growth
cessation and bud set.
After the onset of quiescence (latter part of summer), there is build- up of
carbohydrates reserves and other compounds through photosynthesis. Besides
there is synthesis of high molecular weight, acid-insoluble phosphorus compounds.
As dormancy progresses, this ceases as acid soluble phosphorus compounds
(apparently for storage) are manufactured. Phosphorus uptake and, correspondingly,
phosphorus metabolism cease almost entirely during the deepest part of rest. The
DNA-RNA-protein system has been working actively in autumn and by the time
rest passes to imposed dormancy the metabolic activity is completed through
DNA transcription. RNA and protein synthesis occurs during dormancy and rRNA
being the primary nucleic acid produced. However, Yeh Feng et a1. (1974) noted
that later in dormancy (January to March) changes in the nucleus occur and more
protein and RNA are produced. Transcription may be complete by end of rest but
mRNA is not able to enter the cytoplasm because of a double membrane around
the nucleus and the absence of nucleopores. Other changes associated with the
onset of dormancy include a decrease in amino acids, perhaps due to active
 Bud Dormancy 7

protein synthesis and increase in the protein content of the ribosomal and
mitochondrial fractions. There is rapid hydor1ysis of polysaccharides and the
appearance of large quantities of oligosaccharides also during dormancy. Smash
(1954) reported that the protop1ast in the cell contract and assume a convex shape. They
are withdrawn from the cell wall, rupturing the plasmodesmata, and become
covered with a lipoid layer.
During dormancy there is increase in nuclear dimensions and protoplasmic
content, increased metabolic activity of cell wall formation and lipid droplets
(comprise the source for membranes of new organelles formed later) are at a
maximum in early December (Yeh et al; 1974). Many changes seen at the inception
of dormancy are reversed as the dormancy is coming to a close. An increase in
amino acid content occurs due to rapid breakdown of the protein, nucleopores
reappear, allowing mRNA to enter the cytoplasm, carbohydrates are markedly
decreased especially starch, organic acid content increases just prior to break
(El-Masy and Walker, 1969). Loss of nitrogenous material and phosphorus uptake
is resumed. The respiratory quotient decreases from about 3 (indicating anaerobiosis)
to between 1 and 1.5 (Bachelard and Wightman, 1973). The metabolic activity
changes from catabolism to anabolism about two weeks prior to renewed growth.
Before this occurs, the carpel size of the golgi bodies increases. There is increase in
number of mitochondria indicating that more energy is needed for cell reactions.
Smooth endoplasmic reticulum (ER) is transformed into rough ER in preparation
for synthesis of new proteins. The protoplasts swell and reestablish plasmodesmatic
connections (Samish, 1954). There is increase in total sugars especially mono-
sachhrides (glucose and fructose) about two weeks after the completion of rest.
Various growth inhibiting compounds such as abscissin synthesis is stimulated
by the onset of short day or long dark photoperiods in winter. Greater amounts
of ABA have been found in dormant buds than in actively growing ones. ABA
inhibits the synthesis of proteins, RNA and other metabolic processes, imposing
dormancy in meristematic tissues. Short days and dormancy reduce the frequency
and pore size of plasmodesmata in apical bud cells. This disrupts the intercellular
communication and leads to cessation of growth. In addition more Ca2+ gets
precipitated in cytosol during dormancy and restricts the intercellular
communication thereby leading to dormancy under these conditions. Besides the
symplastic pathways (plasmodesmatal connections) are shut down in the apical
meristem during dormancy induction in response to short days. This blockage is
brought about by formation of 1,3-β-D- glucan, produced by activation of
1,3-β-D- glucan synthase complex which prevents functioning of apical meristem.
Low temperature (chilling) restores the symplastic organization by enhancing the
production of1,3-β-D- glucanases and their delivery into the vicinity of
8 Fruit Tree Physiology

plasmodesmata where glucans are digested by glucanases. This results in restoration

of symplastic connections and allows cells in the apical meristem to exchange
cell metabolites and acquire growing capacity (Rinne et al; 2001)
Phytochromes are present in plastid membranes and other surface membranes
and also play an important role in imposing dormancy. Due to photoperiodic
changes phytochromes induce synthesis of ABA in plastids and also fascilitate its
release. It is then translocated to buds where it leads to temporary suspension of
growth and inhibits metabolic activity. Dormin (Abscissin II) has also been found
to be involved in dormancy. In some plants such as leafy Spurge, a compound
probably sugar has been found to induce dormancy.

Molecular Mechanisms of Hormone Action in Bud Dormancy

Since various plant hormones influence the onset and maintenance of dormancy
in various plant buds, there has been considerable effort to understand how these
hormones act to control dormancy. Much work has been focused on cloning
genes that are differentially expressed in response to hormonal or environmental
signals known to affect dormancy. Studies have shown that auxin increases cell
growth and division by interacting with the degradation pathways controlling the
level of cyclins and other key components of the cell division machinery. However,
the process by which auxin inhibits growth of distal buds remains unknown.
Genes like Rmsl1 and Rmsl2 have been identified, whose products are transported
through the plant and they interact with auxin to increase apical dominance
(Beveridge et al; 2000). Further, it is now known that ABA likely acts through
the cyclin-dependant kinase inhibitor Ick1 to prevent cell division and to maintain
dormancy in affected tissues. The induction of Ick1 by ABA may also explain the
inhibition of bud growth caused by low temperatures. Little is known about the
action of GA, but it is clear that GA can promote cell cycle activity in some way
(Sauter et al; 1995). GA has been shown to be sufficient for induction of
endoreduplication of DNA in Arabidopsis and for induction of S-phase, but not
M-phase, of the cell cycle. It is possible that antagonism between GA and ABA
signaling results in GA inhibiting Ick1 production and allows cell cycle progression
through the S-phase. Recent work has indicated that overexpression of cytokinin
biosynthetic genes increased cytokinin levels and reduced correlative inhibition
in axillary buds. Increased cytokinin levels can induce Knat1, a gene involved in
meristem growth and development. Cytokinins also seem to play a role in the
induction of cyclin D transcripts, which are required for cell division (Gaudin
et al; 2000). Phytochrome responsive genes have provided significant insights
into the mechanisms by which phytochrome controls growth and development.
Expression of GA biosynthesis genes and interactions of GA with ABA and sugar
 Bud Dormancy 9

signaling indicate that phytochrome control of dormancy probably acts through

GA and ABA (Neff et al; 2000). With this information, the method by which
environmental stimuli such as light and temperature may control dormancy
induction and termination through specific plant hormones is becoming more
evident. Light and temperature clearly influence production of sugar and alter
phytochrome signaling. Both of these signals in turn affect ABA and GA
biosynthesis. Certainly ABA, and probably GA, influences adventitious bud growth
through the action of Ick1 and possibly other cell cycle inhibitors. In addition to
environmental signals, the action of auxin and cytokinins produced by growing
meristems also clearly affects cell cycle activity. Additional studies have also
indicated a substantial amount of cross-talk between the signaling pathways of
auxin, cytokinin, GA, ABA, and sugar/ light (Xin et al; 1998).
Expression of several dormancy responsive genes like genes coding for
KNOTTED-like homeodomain proteins in the buds of deciduous trees and other
plants has been characterized. Certain members of this family, along with members
of the CLAVATA group and WUSCHEL, play a crucial role in the proliferation of
undifferentiated cells of shoot meristems and are required for maintenance of growth.
During the induction and release of endodormancy some proteins are relocated
or modified in addition to expression of genes. Along with changes in gene
expression, there are epigenetic changes associated with endodormancy induction
and release. Changes in DNA methylation have been observed during the induction
and breaking of endodormancy. Increased DNA methylation is associated with
induction while demethylation leads to release of dormancy.
10 Fruit Tree Physiology

Mitotic activity in buds: In dormant buds the mitotic activity in the cells of
buds is zero a condition occurring from December through February.

SIGNIFICANCE OF DORMANCY
The main function of dormancy is to ensure that the plant is able to survive
periods of adverse environmental conditions, such as low temperature and reduced
wate ravailability. Plant cells normally contain a large amount of water which is
liable to freeze at low temperature and can damage the protoplasm e.g tropical
plants but those of temperate and arctic regions are frost resistant. In these frost
resistant plants the growth is arrested and the whole plant, including the apical
meristem, is relatively frost resistant because shoot apices cease active growth
and remained enclosed in bud scales to form winter resting buds which become
dormant and are frost resistant than actively growing buds. It is observed that
frost resistance of dormant tissue is due to certain protoplasmic characters, and
is not primarily due to the presence of the bud scales, the protective function of
which is probably concerned with reduction of water loss.
 Bud Dormancy 11

There is also reduced water loss due to bud scales and hence prevention from
drying from drought during winter conditions. There is also reduced surface area
for evapotranspiration due to falling of leaves and thereby prevents the plants
from water stress.

FACTORS AFFECTING BUD DORMANCY

1. External/ environmental : Temperature, photoperiod, restricted O2 supply.
2. Internal (Hormonal) : ABA, gibberellins, auxins, cytokinins and ethylene.
3. Nutritional : Nutrient diversion, Ca2+.
4. Genetic factors
1. External/ environmental factors:
Temperature : Terminal buds of apple shoots rapidly enter dormancy in
autumn and then start to exit dormancy initially slowly but more rapidly in
winter before spring bud burst. The buds have the ability to enter into and
exit from dormancy even at warm temperatures (15oC) but cold temperatures
are generally considered as the environmental cues responsible for the initial
induction (in autumn) and subsequent alleviation (in winter) of the chilling
requirement during bud dormancy. Apple and walnut buds take longer time
to enter into and exit from dormancy at warm temperature. The shoots in
Granny Smith apple reach maximum dormancy after only 100 Utah chill
units (CU) in a cold area while those from a warmer area reach maximum
dormancy after 600 CU. To satisfying the rest requirement chilling
temperatures are not effective until a significant amount of leaf fall has
occurs in the fall. The most efficient continuous temperature for satisfying
the rest is 8oC. Temperature above 8oC is less effective and all chilling
effectiveness is loss with temperatures above 12oC. Chilling effeciency of
low temperature is enhanced when moderate temperatures (i.e.13-15oC) are
interspersed, for 8 hours in a 24 hour cycle, with chilling temperatures.
However high temperature (i.e. 19oC or high), interspersed in the same manner,
will result in negation and the degree of negation depends upon the level and
length of high temperature exposure period. However, short periods (2 to 4
hours) of 20oC in a diurnal cycle can enhance the chilling effect of the low
temperatures. The degree of exposure to winter chilling that an apple receives
can influence the rate of starch conversion in the root system. The conversion
of starch to sugar under chilling is an important process for freeze protections
and bud burst in deciduous fruit plants. In persimmon, high root and shoot
temperatures may delay the starch conversion, causing a delay in bud break,
in comparison to lower temperatures. Rate of starch conversion is influenced
in apple root system by the degree of exposure to winter chilling. This
conversion of starch to sugar is a mechanism that provides freeze protection
and prevents bud burst in deciduous fruit plants.
12 Fruit Tree Physiology

Photoperiodism : Photoperiodism plays an important role in regulation of bud

differentiation and flowering. The buds of woody species respond more to the
photoperiod than to cold temperatures. In general long day conditions promote
vegetative growth and short-days bring about cessation of growth. However,
some species of Pyrus, Malus and Prunus are relatively insensitive to day length
changes. In many woody plants, dormancy is controlled by photoperiod or
temperature variation. The response of woody plants to photoperiodic cycles
appear to depend upon length of the dark period and if a long dark period is
interrupted by a short period of light, the effect of dark is nullified and dormancy
is delayed. At the time of maximum growth activity, usually only a short time
after the completion of the rest period, the buds are able to break, leaves to
expand, and shoot internodes to grow in a wide range of photoperiods.
When the long daily dark period in diurnal cycles are interrupted by a short
light period, the plants show a growth behavior similar to that of plants kept
under long day conditions leading to cessation of growth and formation of
terminal buds e.g in Actinidia chinensis, 8 h days at higher temperatures (25oC)
induced bud dormancy and stem elongation and leaf production were halved
after about 90 days whereas growth continued in 16 h days. Buds of woody
plants respond more to photoperiod than cold temperatures and under short
day conditions, the level of inhibitors increases leading to cessation of growth.
Short day (SD) conditions commonly results in full endodormancy in many
deciduous perennials e.g both cold and SD are required for dormancy induction
in grape (Vitis sps.) (Wake and Fennell, 2000). However, sometimes temperature
alone can also induce dormancy in some species. Short day responses require
the involvement of phytochrome A (PHYA). It has been observed that short
pulses of red light during the dark cycle completely inhibit bud dormancy,
whereas far red light has opposite effect (Howe et al; 1996). The following
Fig. shows the interaction of signals impacting dormancy, flowering and growth.

(Chao et al; 2007)

Fig. : Interaction of signals impacting dormancy, flowering, and growth.
 Bud Dormancy 13

Effects of environmental factors such as long term (LT) and short term
(ST) cold and long day (LD) and short day (SD) length on the expression of
proteins such as Phytochrome A (PhyA), Vernalization Insensitive3 (VIN3),
Constans (CO), Flowering Locus T (FT), Dormancy Affecting Madsbox (DAM),
Flowering Locus C (FLC), and on plant hormone abscisic acid (ABA) are shown.
These proteins and hormones in turn impact growth, flowering and dormancy.
Arrows indicate stimulation of the signal, gene action, or developmental process,
and bars indicate inhibition. Cold (ST) and SD both inhibit PhyA action.
Inhibition of PhyA in turn down regulates CO then FT thus inhibiting flowering
and growth and stimulating dormancy. Likewise, cold (LT) induces VIN3 and
blocks expression of DAM/FLC and thus stimulates FT, flowering, growth,
and breaks dormancy
Oxygen (O2) supply : Secondary dormancy in buds is induced by high
temperature and limited oxygen supply. Under such conditions glycolysis is
faster but oxidative breakdown of pyruvate through Krebs cycle is restricted
(because of covering structures i.e bud scales and external leaf primodia in
buds) and some other intermediates are formed and the oxidation of pyruvate
and acetyle CoA takes some other direction. This alternate path is not coupled
with growth processes but with those which increase resistance to unfavourable
external environmental conditions and thereby causes bud dormancy.
2. Internal or hormonal factors :
Role of growth hormones : Plant growth regulators play an important role
during dormancy. Various phases of dormancy like induction, maintenance,
trigger and release are regulated by hormonal and environmental signals.
Endogenous plant growth substances are directly involved in the dormancy
process. The various phases are shown in Fig.

Fig. : Changes in levels of PGRs during different phases of dormancy (Olsen, 2003)
14 Fruit Tree Physiology

Induction : There is decrease in promoter and an increase in inhibitors

activity. Environmental factors such as temperature and light may affect or
trigger this phase.
Maintenance : In this phase, general metabolism and endogenous promoters
are very low while levels of inhibitors are very high. Environmental changes
may shorten this period but will not have an immediate effect on ending of
this phase. Buds become susceptible to frost injury due to premature sprouting.
Trigger : There is dramatic decrease in inhibitors and a sharp rise in promoters.
It is especially sensitive to environmental cues or signals.
Bud break : There is high amount of promoters and low inhibitors leading
to increase in enzymatic activity. Depending on genus, species or other factors,
the length of each phase varies.
(i) Abscissic acid (ABA) : Accumulation of ABA in buds is the primary
factor during the dormancy induction and is associated with the rest of
buds. During dormancy ABA level increases and reaches maximum during
the deepest dormancy stage, when water levels in the bud is low. As soon
as water levels in the buds increase at the end of dormancy and at
budbreak stage the ABA levels decrease rapidly. The conversion of the
apical meristem into a dormant bud is mediated by ABA. The newly
developing leaves growing above the meristem become converted into
stiff bud scales that wrap the mersistem closely and will protect it from
mechanical damage and drying out during the winter. In the metabolic
processes of the bud abscissic acid plays a more active role.
(ii) Gibberellins : Gibberellins are closely related to chilling based dormancy
phenomena in dormant buds. GA3 concentration in buds increases as
they expand in the spring. The increase in concentration is more rapid
after the chilling requirement has met e.g a marked decrease in the level
of gibberellins and a rise in the activity of the inhibitors during deep
dormancy (Nov.- Dec.) have been observed in the flower buds of the
cherry cultivar Napoleon. The endogenous gibberellins level rise sharply
after breaking deep dormancy (Jan.) and enforced dormancy (March-
April). Chilling temperature may affect gibberllins through an effect on
membrane permeability. Inhibitor studies support the theory that chilling
temperature activates the gibberellins mechanism. Ramirz and Cadens-
chavet (1989) reported that endogenous gibberellins content in fruit buds
of apple cv. Red Delicious varied during dormancy. GA3, GA4, GA4+7
are effective in overcoming bud dormancy in many fruit crops but more
satisfactory results are obtained only when application is made directly
to the bud. Knowledge on GA’s role in regulating the induction and
 Bud Dormancy 15

release of bud dormancy is rudimentary, despite extensive research on

plant responses to GA (Razem et al; 2006). In general, GA is involved
in two major regulatory roles during dormancy: i) SD-induced growth
cessation in trees and ii) vegetative bud growth following dormancy
release.
(iii) Auxins : They play a limited role in the release and control of bud
dormancy but are essential for active growth. There is decline in activity
of free auxins and appearance of inhibitors during transition towards
dormancy and the activity of free inhibitors is maximum and no auxins
are found in the buds of apple during deep dormancy. High amounts of
auxin like substances and low amounts of β inhibitors are present during
the early stages of dormancy (Jan.) in pear cvs. LeConte and Bartlett. In
LeConte trees, the deep dormancy period starts when the promoters
disappear completely and inhibitors are present in high amount in early
Feb. and in Bartlett deep dormancy starts in late Jan. After deep dormancy,
the auxin like substances increases while β inhibitors decreased gradually.
Auxin like substances increase to maximum level, while the β inhibitors
decrease and reach minimum at bud break.
(iv) Cytokinins : Role of cytokinins in dormancy is still unclear, however,
they play a role in bud development and shoot growth that occurs after
release from dormancy. Metabolic processes that are involved with growth
such as DNA-RNA and protein synthesis, an increase in energy
metabolism and a decrease in metabolic paths are stimulated by cytokinins.
Synthetic cytokinins such as TZD and 6-BA stimulate bud break in apple
indicating the role of cytokinins in releasing bud dormancy. Relative
ratio of a specific plant growth regulator is an important factor to determine
its role in any metabolic process. During bud break the relative ratio
between cytokinins and auxins are important, as cytokinins reduce or
soften the inhibitory effect of auxins. Cytokinins are not the cause but
play a supporting role during bud break of fruit trees and accelerate the
development of buds that have been partially released from dormancy.
(v) Ethylene : The breaking hormone (necrohormones) may be ethylene. It
is well documented that the production of ethylene is universal to all
stresses. Exogenous applications of ethylene break dormancy in many
plant tissues. Wang et al. (1986) showed that ACC precursor from
dormancy to active state in Prunus avium and Prunus serrulata in Radiant
Crab apple for delaying flowering and increasing winter bud hardiness
when used at rates upto 200 ppm. But in grape and apple, ethylene does
not ensure budbreak and the ethylene releasing product, ethephon, even
delays the budbreak. Ethepon also delays the bud break of green summer
16 Fruit Tree Physiology

buds and resultant shoot growth. It is thus unlikely that ethylene plays
a role in bud in the control of bud break. In peach ethephon has been
found to prolong bud dormancy by increasing chilling requirements,
however, the mechanism of action is not clearly understood.
3. Nutritional factors :
(i) Nutrient diversion hypothesis (Sachs, 1973, Sachs and Hackett, 1983):
According to this theory the environmental signals modify source/ sink
relationships within the plant in such a way that the shoot apex receives
a better supply of assimilates under favourable conditions while under
noninductive conditions the relationship is altered in such a way that
plant cease their growth to avoid the unfavourable conditions. Sucrose is
the main storage reserve that is transported to the buds during dormancy
release and is mainly stored in stems and roots. Thus release of dormancy
is associated with changes in sucrose, mineral nutrients and thereby
enhanced metabolic activity. The following Fig. shows the various events
during onset and release of bud dormancy:

(ii) Calcium : During active growth Ca2+ is present in vacuoles, cell wall and
intercellular space. As PP and ATP decrease, Ca2+ increases in cytosol,
nucleus. Various ultrastructural changes are observed such as thickening
of cell wall, decrease in plasmodesmata connections, symport transport
stops, limited cell to cell communication which may lead to dormancy.
Wang et al. (2008) observed that short sunlight and low temperature were
the main factors inducing growth cessation, dormancy, and freezing
resistance development and the inducement effect of short sunlight was
closely related to the actions of Ca2+, because Ca2+ played a messenger
role in the signal transduction of short sunlight. By supplementing light,
it was found that with the decrease of temperature, the Ca2+ had an
increasing influx from the vacuole, intercellular space, and cell wall to
the cytosol and nuclei. At the same time, plant growth slowed down and
finally ceased, dormancy started then, and freezing resistance developed,
which indicated that as a messenger in the signal transduction of natural
 Bud Dormancy 17

low temperature, Ca2+ plays an important role in the inducement of

growth cessation, dormancy, and freezing resistance development by
natural low temperature.
4. Genetic factors : The genetic control of the chilling requirement for breaking
the dormancy varies greatly within and among species. Although vernalization
is controlled by a single gene in some species, but multigenic control is the
rule. Mutants have been found in Arabidopsis in which dormancy is controlled
by a single gene. Existence of these mutants suggests that similar cases may
exist in other species also.

Status of Water in Buds

The water is present in the bound state in dormant buds which gives resistance
against frost. However, when the chilling unit requirement is satisfied, a part of
water disasociates from other macro molecules and becomes free e.g conversion
from bound to free water occurred at the end of chilling unit accumulation (600
and 4000 CUs) in low chilling Anna and high chilling Northern Spy apples,
respectively. In cold climates, maximum hardiness (implying maximum bound
water) may occur after satisfying the chilling unit requirement (i.e ecodormancy).

Release of Bud Dormancy

The release of bud dormancy is a sequential process, with each phase characterized
by different metabolic and hormonal effects. Current evidences indicate that control
of bud dormancy is very complex and involves interactive effects of growth
hormones and other compounds. Changes in cell cycle specific gene expression
occurs during release of axillary buds of pea (Pisum sativum) potato, adventitious
buds of leafy spurge (Euphorbia esula) (Horvath et al; 2002) and Jerusalem
artichoke (Freeman et al; 2003) from dormancy. Dormancy breaking results in up
regulation of genes that act at G1-S phase transition such as histones (Horvath
et al; 2002). In addition, several post translational modifications to key enzymes
involved in cell cycle regulation occur shortly after dormancy break in several
plant systems. The attaractive model of Saure (1988) suggests that release of bud
dormancy may occur in following phases
Phase I : Removal of the primary cause of dormancy, largely involving the
enzymes that are not most active at chilling temperatures. These
changes cause transition from endodormancy to ecodormancy.
Phase II : This is a catabolic phase and is characterized by increased metabolic
activity, increased GA activity which promotes synthesis and/ or
activation of hydrolytic enzymes (e.g amylase, protease and
18 Fruit Tree Physiology

ribonuclease) and development of a translocation system, causing

mobilization of reserve carbohydrates essential for growth. During
this phase, bud swelling indicates transition from endodormancy to
ecodormancy.
Phase III : In this phase there is additional increase in auxin, GA and cytokinin
activity, mobilization of reserve carbohydrates, mitosis and complete
release from dormancy. It ends with bud break and rapid shoot growth.
There are different approaches to examine mechanisms of dormancy release
in plants:
1. Regulation within the apical meristem due to changes in the cell to cell
communication and plasmodesmatal connections (Rinne et al; 2001) or in
the cell cycle (MacDonald, 2000).
2. Regulation of water where initial reports based on supercooling examined
the vascular connections into the bud (Sakai, 1979). More recently, the
sequence and regulation of water uptake into the bud (De Fay et al; 2000)
and water status/ availability (Faust et al; 1997) during dormancy has been
addressed.
3. Molecular events involved in the perception and transduction of dormancy
breaking signals during chemical induced dormancy release in a Vitis model
were used by Or et al. (2000, 2002).
4. The approach of Champagnat (1989) and others is based on a mechanism of
dormancy induction (and release) via a metabolic or communication block,
or a permeability barrier between the bud and adjacent tissues.
5. Gevaudant et al. (2001) studied both the buds and the underlying tissue in
peach and found greater accumulation of PPA (Prunus persica H+-ATPase)
transcripts in the underlying bud tissues compared to the buds themselves at
the beginning of the dormancy period (October). This group attributed
increased sucrose absorption in tissues underlying the bud during October to
a stimulated H+/ sucrose cotransport driven by PPA genes and suggested this
to have a role in paradormancy. It is hypothesized that in the evolution from
paradormancy to endodormancy there is involvement of chilling induced
specific decrease in certain PPA isoforms in tissues underlying the buds in
November and December could be involved.
Molecular mechanisms of bud dormancy release : To identify genes and
gene products that mediate signal transduction of a dormancy release signal or
depression of meristematic activity, Or et al. (2000) while examining release of
bud dormancy in grape by hydrogen cyanamide (HC) application compared RNA
population from HC treated and control buds. They identified transcripts for a
 Bud Dormancy 19

sucrose non fermenting (SNF) like proteins and suggested that this kinase may
be involved in perception of stress signal induced by HC in grape buds. They
hypothesized that disruption of respiratory metabolism caused by H2O2 (generated
by HC induced oxidative stress) and shuts off catalase (free radical scavenger)
gene expression soon after HC application. These observations indicate that bud
dormancy release coincides with upregulation of antioxidant system. It is
noteworthy that cold acclimation of temperate fruits (chill induced response) is
often accompanied by an upregulation of antioxidant machinery required for
protection against freezing stress. Leida et al. (2010) found that genes like DAM4,
5 and 6 coding for MADS-box transcription factors are related to growth cessation
(dormancy) and terminal bud formation in the evergrowing mutant of peach. The
main classes of genes relevant for the bud outgrowth controlling mechanism are
as under:
Functional categories of genes expressed during bud release, identified from
c-DNA microarray experiments in different plant species
1. Stress/ response/ defense / detoxification
2. Sugar metabolism
3. Hormones-induced genes
4. Cell cycle and DNA processing
5. Energy generation
6. Transcriptional factor and signal transduction
In particular in all the researches carried out it appears evident the activation
of stress-response/ defense related genes as well as genes controlling sugar
metabolism, cell cycle and affected by hormones. It is interesting to compare the
gene list of differentially expressed genes because it is possible to determine
several recurrent themes emerging as well as genes which are not described in
other species (Mazzitelli et al; 2007).

Methods to Overcome Bud Dormancy

1. Chemical agents : Various chemicals have been reported to overcome rest
or other forms of dormancy in temperate woody perennials. To date, the
following groups of chemicals have been reported to break dormancy in
plants. Mineral oils, nitrogen containing compounds, uncoupling agents,
sulphur containing compounds, salts, acids, toxic compounds, growth
regulators, cyanamide (Hydrogen cynamide, calcium cynamide) e.g calcium
cyanamide in the presence of CO2 and by acidification produces free hydrogen
cyanamide. Calcium cyanamide undergoes a partial hydrolysis to hydrogen
20 Fruit Tree Physiology

cyanamide and cyanamide ions are the active form of chemical. Exogenous
applications of ethylene help to break dormancy in many plant tissues. An
inhibitor of ethylene biosynthesis, S-(E)]-2-amino-4-(2-aminoetoxy)-3-
butanoic acid (AVG) delays bud break. Sublethal stress may overcome rest
by increasing membrane permeability. It is speculated that an increase in
membrane permeability may be the cause of breaking rest. Increased
production of ethylene due to sublethal stresses may be due to the release or
activation of the ethylene forming enzymes (EFE) that is reported to be
associated with membrane. Various chemicals used for breaking the bud
dormancy are: GA, HCN, KNO3, IBA, NAA, BAP, ascorbic acid etc. used
at different concentrations in different fruit crops.
Dormex (special formulation of hydrogen cyanamide) leads to high and
very uniform bud break has proved to be a particularly effective agent for
breaking dormancy, particularly in woody plants such as grapes. Since the
period of dormancy is accompanied with high levels of natural growth
inhibitors and hence, cessation of growth. The main natural plant growth
inhibitor believed to be involved with dormancy is the hormone- abscisic
acid. Studies have been initiated to investigate the relationship between abscisic
acid levels in grape buds following Dormex application and release from
dormancy. It was found that there was no correlation between abscisic acid
levels in the plant and Dormex induced release from dormancy. The enzyme
catalase catalyses the breakdown of hydrogen peroxide to water and oxygen.
It possesses a particularly important role in the plant, as many enzyme
reactions produce hydrogen peroxide as a by-product of metabolism, which,
if not detoxified, could have serious deleterious effects on the plant. It has
been known for many years that hydrogen cyanamide inhibits the action of
catalase Youngman and Trostberg (2008). However, since the application of
Dormex at the recommended rates does not lead to phytotoxicity, it must be
assumed that the plant possesses an alternative mechanism to detoxify
hydrogen peroxide. It is conceivable that the activation of this second pathway
is related in some way to the mechanism of dormancy breaking by Dormex.
Dormex inhibits the action of catalase, the plant subsequently detoxifies
hydrogen peroxide via a sequence of reactions which are eventually coupled
to the oxidative pentose phosphate pathway. It could be shown that the
presence of Dormex stimulates the reaction between hydrogen peroxide and
ascorbate, which in turn leads to an increase in the rate of turnover of the
pentose phosphate pathway. The consequences of this are two fold: by
stimulating the oxidation of ascorbate by hydrogen peroxide, Dormex ensures
that although it inhibits the catalase dependent breakdown of hydrogen
peroxide, another pathway is stimulated, thus allowing its detoxification.
 Bud Dormancy 21

Due to the concomitant activity increase in the pentose phosphate pathway,

a range of essential substances such as lipids, RNA, DNA and pentose sugars
are produced at higher rates. These compounds are fundamental for new
growth, as would be required during the release of buds from dormancy.
These studies demonstrated that Dormex exerts an effect on the plant whereby
the synthesis of compounds essential for new growth is markedly stimulated.
Nevertheless, it is likely that compounds exerting an effect on a
physiological process such as dormancy also influence the hormone status of
the plant. Investigations have shown that Dormex affects the cytokinin
physiology via certain mediator compounds. It could also be demonstrated
that these mediators per se led to a further stimulation of the pentose phosphate
pathway.
2. Environmental factors :
(i) Chilling/ stratification : The rest period is overcome by exposure to low
chilling temperatures in temperate perennial woody plants. The speed of
bud break and the vigour of the elongating shoot are related to the degree
of chilling experienced by plants. The effective chilling temperatures and
the duration of the chilling period required to satisfy the rest requirement
is dependent on the genetic make up of the plant and possibly on the
environment of the preceding season. According to Westwood (1978)
chilling requirement is an adaptive feature and depends on the plants
native area of origin. For example relatively short chilling periods are
required by plants indigenous to mild climates with long frost free seasons
and those native to high latitudes or elevations with continuously long,
cold periods. While long chilling requirement to ensure the maintenance
of cold hardiness through the freezing period are required by plants native
to moderate climates, characterized by fluctuating warm and cold periods
during the winter. The effective temperature range for chilling satisfaction
appears to vary with the plants. Generally low temperature promotes rest
release and in many woody plants temperatures between 0°C and 7.2oC
for 2 to 3 months break the rest.
High temperature results in negation of chilling. The temperature
range for induction of secondary dormancy decreases progressively during
the post dormancy period. Moderate temperatures applied during the early
part of dormancy can cause chilling negation. In peach, moderate
temperature during November and December counteract chilling. Short
periods of high temperature during a daily cycle can negate chilling.
There is a greater chilling negation if the high temperatures occur every
2 days rather than 4 days.
22 Fruit Tree Physiology

(ii) Photoperiod : Although photoperiod is the key agent leading to dormancy

but its effect can be overcome by temperature, light intensity during
photoperiod, concentration of nutrients in soil medium, soil moisture etc.
Low light intensities produce dormant buds at temperatures of 10-230C.
Cool temperature rather than a short photoperiod causes floral induction
whereas warm temperature rather than long photoperiod inhibits flowering.
3. Calcium signaling : By exogenous application of HCN in grapes several
Ca- signaling related genes with remarkable differences in expression level
were observed between HC-treated and control buds. This indicates that
Ca2+ signaling is involved in grape bud dormancy release. Ca- dependent
protein phosphorylation studies have revealed that in HCN treated buds the
Ca- dependent histone phophorylation was higher upto 70 per cent which
indicates the role of calcium signaling in bud dormancy release (Pang, 2007).

CONCLUSION
Dormancy is a complex process that results from the interaction of several factors
like environmental, hormonal, nutritional and genetic ones, along with their
corresponding signalling pathways. The environmental controls of bud dormancy
are mediated by hormones and a balance between growth inhibitors and promoters
is important in regulating both entry and exit from the dormant phase in buds.
Molecular mechanisms suggest that dormancy is under the control of multigenes
in various species. Several chemicals like Dormex (hydrogen cyanamide),
polyamines etc. are used to overcome dormancy.
Proper understanding of the mechanisms of dormancy induction, maintenance
and release can be achieved through a multidisciplinary approach involving,
horticulturists, physiologists, biochemists, and molecular geneticists at different
levels. Molecular approaches like microarray technique, gene mapping studies
etc. will provide information about gene groups that control bud dormancy related
traits in different fruit crops in future.

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release in tree buds: changes in degree of dormancy, respiratory capacity, and major
cell constituents in over wintering vegetative buds of populus balsamifera. Can J
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Beveridge CA, Symons GM and Turnball CGN (2000) Auxin inhibition of decapitation
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RMS1 and RMS2. Plant Physiol 123: 689-698.
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Champagnat P (1989) Rest and activity in vegetative buds of trees. Ann Sci For
46 : 9–26.
Chao WS, Foley ME, Horvath DP and Anderson JW (2007) Signals regulating dormancy
in vegetative buds. Inter J Plant Dev Biol 1(1): 49- 56.
De Fay, Vacher EV and Humbert F (2000) Water related phenomenon in winter buds
and twigs of Picea abis until bud burst: A biological, histological and NMR studies.
Annl Bot 86: 1097-1107.
Doorbens J(1953) Review of the literature on dormancy in buds of woody plants.
Medelingen van de Landbouwhoge School to Wageningen / Netherland 53: 1-24.
El-Mansy HI and Walker DR (1969) Seasonal fluctuations of amino acids, organic
acids, and simple sugars in Elberta peach and Chinese apricot flower buds during
and after rest. J Amer Soc Hort Sci 94: 184.
Faust M, Erez A, Rowland LJ ,Wang SY and Norman HA (1997) Bud dormancy in
perennial fruit trees: physiological basis for dormancy induction, maintenance and
release. Hort Sci 22: 371-377.
Freeman D (2003) Isolation, characterization and expression of cyclin and cyclin-
dependent kinase genes in Jerushalem artichoke (Helianthus tuberosus L). J Exp
Bot 54: 303-308.
Gaudin V, Lunness PA, Forbert PR, Towers M, Khamlichi C, Murray JAH, Coen E and
Doonan JH (2000) The expression of D-cyclin genes defines distinct developemental
zones in snapdragon apical meristems and is locally regulated by the cycloidea
gene. Plant Physiol 12: 1137-1148.
Gevaudant F, Patel G and Guilliot A (2001) Differential expression of four members of
the H+ ATPase gene family during dormancy of vegetative buds of peach trees.
Planta 212: 616-626.
Horvat DP, Chao WS and Anderson JV (2002) Molecular analyses of signals controlling
dormancy and growth in underground adventitious buds of leafy spurge. Plant
Physiol 128: 1439-1446.
Horvath DP, Anderson JV, Chao WS and Foley ME (2003) Knowing when to grow:
signals regulating bud dormancy. Trends in Plant Sci 8:11-15.
Howe GT, Gardner G, Hackett WP and Furnier GR (1996) Phytochrome control of
short-day-induced bud set in black cottonwood. Physiol Plant 97: 95-103.
Lang GA (1987) Dormancy: A new terminology. Hort Sci 22(5):817-820.
Leida C, Terol J, Marti G, Gerardo MA, Mara L, Badenes L and Rios G (2010)
Identification of genes associated with bud dormancy release in Prunus persica by
suppression subtractive hybridization. Tree Physiol 30 (5): 655- 666.
24 Fruit Tree Physiology

MacDonald JE (2000) The development basis of bud dormancy in one year old Picea
and Pseudostuga seedling. In: JD Viemont and J Crabbe (eds). Dormancy in Plants.
CABI, New York.
Mazzitelli L, Hancock RD, Haupt S, Walker PG, Pont SDA, McNicol J, Cardle L,
Morris J, Viola R, Brennan R, Hedley PE and Taylor MA (2007) Co-ordinated gene
expression during phases of dormancy release in raspberry (Rubus idaeus L.) buds.
J Exp Biol 58 (5): 1035–1045.
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Genes Dev 14: 257-271.
Okuba H(2000) Growth cycle and dormancy in plants. In: JD Viemont and J Crabbe
(eds.) Dormancy in Plants–From Whole Plant Behavior to Cellular Control, CABI,
Pp. 1–22.
Olsen JE(2003) Molecular and physiological mechanism of bud dormancy regulation.
Acta Horticulturae 618: 437-453.
Or E, Belausov EPI and Ben Tal Y (2000) Changes in endogenous ABA levels in
relation to dormancy cycles in grapevine grown in a hot climate. J Hort Sci &
Biotech 75(2): 190-194.
Or E, Vilozny I, Eyal Y and Ogrodvitch A (2002) Dormancy in grape buds: isolation
and characterization of cDNA and analysis of its expression following chemical
induction of bud dormancy release. Plant Sci 162: 121-130.
Owens JL and Molder M (1973) Astudy of DNA and mitotic activity in the vegetative
apex of Douglas fir during the annual growyh cycle. Can J Bot 51: 1395-1409.
Pang (2007) Involvement of calcium signalling in dormancy release of grape buds. J
Exp Bot 58(12): 3249-3262.
Ramirz H and Cardenas-Chavez LF (1989) The effect of temperature and photoperiod
on gibberellin activity in apple buds during dormancy. Acta Horticiculturae 239:
257–259.
Razem FA, Baron K and Hill RD (2006) Turning on gibberellin and abscisic acid
signaling. Curr Opinion Plant Biol 9: 454- 459.
Rinne PLH (2001) The shoot apical meristem restores its simplistic organization during
chilling-induced release from dormancy. Plant J 26: 249–264.
Rohde A and Bhalerao RP (2007) Plant dormancy in the perennial context. Trends Plant
Sci 12: 217-223.
Sachs RM (1977) Nutrient diversion: a hypothesis to explain the chemical control of
flowering. Hort Sci 12: 220-222.
Sachs RM and Hackett WP (1983) Source sink relationships and flowering. In Beltsville
Symposia in Agricultura Research. Pp. 263- 272.
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Sakai A (1966) Seasonal variation in the amounts of polyhydric alcohol and sugars in
fruit trees. J Hort Sci 30: 49-52.
Samish RM (1954) Dormancy in woody plants. Ann Rev Plant Physiol 5:183.
Saure MC (1985) Dormancy release in deciduous fruit trees. Hort Rev 7: 239-300.
Sauter M, Mekhedov SL and Kende H (1995) Gibberellin promotes histone H1 kinase
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of three Vitis genotypes to short photoperiod. Physiologia Plantarum 109: 203-210.
Wang HB, Wang XD, Cheng CG, Wang BL, Li M, Gao DS and Liu FZ (2008) Natural
inducing factors of peach bud dormancy and roles of Ca2+ in the dormancy induction.
NCBI 19(11): 2333-2338.
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a plant bioregulator, thidiazuron. Phytochemistry 25(2): 311-317.
Westwood NH (1978) Temperate zone Pomology. WH Freeman San Francisco.
Xin X, Lammeren AAM, Vermeer E and Vreugdenhil D (1998) The role of Gibberellin,
ABA and sucrose in the regulation of potato tuber formation Invitro. Plant Physiol
117: 575-584.
Yeh F, Campbell WF and Walker DR (1974) Ultrastructural changes in peach flower
buds during rest. J Amer Soc Hort 99: 427.
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Further Suggested Readings

Crabbe and Barnola (1996) A new conceptual approach to bud dormancy in woody
plants. In: Plant Dormancy G A Lang (Edt.), CAB International, Wallingford, Oxon,
UK. Pp. 83-113.
Pallardy SG and Kozlowski TT (2008) Physiology of woody plants, Academic Press
London, Pp. 42-67
 2

CHILLING REQUIREMENT
IN FRUIT CROPS

C ertain fruit species require low temperature exposure (chilling) for normal
and abundant bud break to occur in the spring. The classic definition of
chilling requirement is the number of hours the temperature is below 45 degrees
F and above 32 degrees F. More recently, it has been suggested that hours above
65 degrees F be subtracted from the previous calculation. While it sounds feasible
at first, the National Weather Service does not routinely calculate and publish
these statistics. Chilling temperatures are not effective in satisfying the rest
requirement until a significant amount of leaf fall has occurred in the fall. In
certain years, insufficient chilling has been recognised as an economic problem
in the production of several temperate zone fruit species, particularly those
temperate fruits which are grown in sub temperate climates. Symptoms of
insufficient chilling are many but generally can be recognised as a delay in flower
and vegetative bud break, bud break spread over an extended period of time, or
in severe cases a total lack of bud break. It is not uncommon, in insufficiently
chilled trees, to find large fruit and flowers on the same shoot as well as sparse
bud break levels.

IMPORTANCE OF CHILLING
1. Delayed foliation : A classic symptom of insufficient chilling is delayed
foliation. A tree may have a small tuft of leaves near the tips of the stems
and be devoid of leaves for 12 to 20 inches below the tips. Lower buds will
break eventually but full foliation is significantly delayed, fruit set is reduced,
and the tree is weakened. Furthermore, heavy suckering from lower parts of
the tree causes management problems, and normal development of next year’s
fruit buds can be impaired.
 Chilling Requirement in Fruit Crops 27

2. Reduced fruit set and buttoning : Flowering, in response to insufficient

chilling, often follows the pattern seen with leaf development. Bloom is
delayed, extended, and due to abnormalities in pistil and pollen development,
fruit set is reduced. In many peach cultivars, flowers drop before or around
shuck split, but in others such as ‘Jersey Queen’ and ‘Harvester’, buttons
form. Buttons result from flowers which apparently have set but never develop
into full-size fruit. The fruit remains small and misshappen as they ripen. If
they are cut these fruit open, the seed is dead. Because buttoning is not
apparent early in the season, growers can not thin off the abnormal fruit and
the developing buttons serve as a food source and over wintering site for
insects and diseases.
3. Reduced fruit quality : The effects of insufficient chilling on fruit quality
are probably the least discussed but appear to be very common. The effects
on leaf growth and fruit set are dramatic but the effects of insufficient chill
on fruit quality are subtle, and can occur when other symptoms do not.
Insufficient chilling will cause many cultivars to have an enlarged tip and
reduced firmness. Furthermore, fruit ground coloration may be greener than
usual, possibly due to the fruit losing firmness before the ground color can
fully change from green to yellow. The extent of these quality problems
depends on the cultivar and the degree of chilling deficiency.

Length of Chilling Requirement in Fruit Species

There are some observable symptoms in fruit trees that have chilling hour’s
incompatible with the local climate. Fruit trees with lower chilling requirement
than necessary, frequently experience crop losses due to early bloom and late
spring frost damages the crop. In these situations, the chilling requirement exceeds.
Conversel, planting varieties with higher chilling requirements than required can
result in uneven bloom and/ or delayed foliation. In these cases, the chilling
requirement is not met. Chilling requirements are based on averages and calculated
averages are made up from extremes and each chilling requirement has some
latitude. In general, for elevations above 6,000 ft, varieties with chilling
requirements above 1,000 hours, for elevations between 4,000 and 6,000 ft, varieties
with chilling requirements between 750 and 1,000 hours and for elevations between
2,500 and 4,000 ft varieties with chilling requirements between 500 and 750
hours should be used. There is some flexibility depending on orchard site/ micro
climate.
Beginning sometime in late summer or early fall, many deciduous fruit buds
gradually proceed into a state of inhibition that culminates into a deeply inhibited
state from which buds will emerge following the accumulation of a certain amount
28 Fruit Tree Physiology

of low temperatures. The amount of low temperature required to release buds

from this inhibited state will vary with species, cultivar within a species, as well
as bud type. For example, the peach cultivar Florida Prince has a chilling
requirement of aproximately 50 hours (at 80C) while Mayflower requires over
1200 hours of low temperature for release from the resting state (Savage et al;
1963). Also, in peach, vegetative buds located on the shoot apex (i.e. terminal
buds) have the shortest chilling requirement while lateral vegetative buds have
the longest. Generally, flower buds are intermediate in chilling requirement
although in rare cases the flower bud chilling requirement may exceed that of the
lateral vegetative buds (Scalabrelli and Couvillon, 1986; Savage et al; 1963).
Because of this characteristic, apple and other pome fruit crops are less affected
than Prunus by insufficient chilling since pome flower buds are produced terminally
along with a vegetative bud, both which have a short chilling requirement. In
Prunus, the flower buds are always borne laterally. For example, apple yields in
middle Georgia were unaffected by the low chilling winter of 1973-74, whereas
peach yields were reduced over 60 per cent (Kirman, 1975).
There is also data available which show that effective accumulation begins
only after significant leaf fall has occurred in the fall. Chandler (1960) reported
that shoots which hold their leaves late into the autmn seem to require more
chilling hours to complete rest. Although, the winter in southern California was
colder in 1958 than that of 1957, fruit trees bloomed later in the spring of 1959
than the spring of 1958 due to the late leaf fall that occurred in the fall of 1958.
Walser et al . (1981) reported that time of leaf abscission in the fall was related
to duration and intensity of rest in Gleason Elberta peach leaf terminal buds.
Also, Reeder and Bowen (1977) reported bloom delay in peaches to which nitrogen
application were made the fall preceding spring bloom. Trees to which fall nitrogen
applications were applied held their leaves later into the year than the trees which
received no nitrogen. Walser et al. (1981) as well as others have reported bloom
delay on trees which receive fall application of gibberlic acid. The gibberlic acid
application resulted in leaf retention well into the fall. Thus, it seems that chilling
temperatures received while a significant portion of the leaves remain on a tree
are not effective or as effective as chilling temperatures received once significant
leaf fall has occurred.

Type of fruit Aproximate hours at 450F Equivalent time if continuously

needed to break dermancy exposed to 450F

Almond 200-300 8-13 days

Apple 1200-1500 7-9 weeks
Apricot 700-1000 4-6 weeks
[Table Contd...
 Chilling Requirement in Fruit Crops 29

Contd. Table]

Type of fruit Aproximate hours at 450F Equivalent time if continuously

needed to break dermancy exposed to 450F

Cherry, sour 1200 7 weeks

Cherry, sweet 1100-1300 6-8 weeks
Chestnut 300-400 2-3 weeks
Fig A few hours
Filbert (Hazelnut) 1500 9 weeks
Kiwifruit 600-850 3.5-5 weeks
Olive 200-300 8-13 days
Peach/ Nectarine 650-850 4-5 weeks
Pear 1200-1500 7-9 weeks
Pecan 400-500 3-4 weeks
Persimmon 100 4 days
Pistachio 1000 6 weeks
Plum, American 3500 5 months
Plum, European 800-1100 5-6 weeks
Plum, Japanese 700-1000 4-6 weeks
Pomegranate 200-300 8-13 weeks
Quince 300-400 2-3 weeks
Walnut, Persian 700 (Payne)-1500 (Franquette) 4-9 weeks

Table : Apple varieties of different maturity requiring different chill hours.

High chill Medium chill Low chill

Early maturity Gala Akane, Dorsett Golden (250), Anna (300),

Gravenstein Ein Scheimer (400), 60-39 (400),
88-20 (375), Tropic Mac (300),
Tropic Sweet (300)
Medium maturity Delicious, Golden
Delicious, Jonagold
Late maturity Fuji, Braeburn Granny Smith
Sundowner
Pink Lady
30 Fruit Tree Physiology

Table : Pear varieties with certain species ancestry and different chill requirements
in Australia.

High chill Medium chill Low chill

Pyrus communis Comice, Dr Jules Guyot Packham’s Flordahome, Hood,

Williams, Beurré Bosc Josephine Moonglow, Magness
Conference
Pyrus pyrifolia Nijisseiki, Shinsui, Kosui, Hosui – –
Dan Bae
Pyrus x
bretschneideri – – TsuLi, YaLi

Root stocks : The chilling requirement of a root stock can have an effect on
the chilling requirement of the variety. Most of the clonal apple root stocks used
commercially were selected and bred in much colder climates, so how they work
in warm areas is not been fully tested. Granny Smith seedlings are not so high
chill may be used but the tree vigour tends to be too vigorous. Pear root stocks
have more choice as Williams’ on their own roots require over 1100 hours,
P. calleryana alone requires 400 hours and Williams on P. calleryana require an
intermediate number of hours (Westwood and Chestnut, 1964). Where a root
stock is high chill (e.g MM1111-1100 hours, Young, 1992), the effect on a low
chill scion variety may be less than the other way about as with pears, as the buds
of the scion have had enough chill.
Table : Cumulative chilling hours (Jill Campbell, 2005).

Country name Station name Hours < 45°F Hours >32° and <45°F

Alameda Oakland Foothills 505 495

Pleasanton 922 858
Union City 672 646
Butte Durham 941 798
Colusa Colusa 829 727
Contra Costa Brentwood 791 741
Concord 896 837
Moraga 1127 998
El Dorado Camino 1404 1301
Fresno Firebaugh/ Telles 808 742
Five Points South West 889 808
Five Points/ WSFS USDA 1071 867
Fresno State 872 828
Orange Cove 1029 977
Panoche 705 664
[Table Contd...
 Chilling Requirement in Fruit Crops 31

Contd. Table]

Country name Station name Hours < 45°F Hours >32° and <45°F

Parlier 854 800

Westlands 878 781
Glenn Orland 935 810
Imperial Calipatria/ Mulberry 607 590
Meloland 303 303
Palo Verde II 755 666
Salton Sea East 301 301
Salton Sea West 16 16
Seeley 374 362
Inyo Westmorland North 294 294
Bishop 2077 917
Owens Lake North 1831 1386
Owens Lake South 1905 1114
Kern Arvin-Edison 824 789
Belridge 942 890
Blackwells Corner 950 840
Famoso 1051 1000
Shafter/ USDA 1039 957
Kings Kettleman 611 590
Stratford 944 876
Lassen Buntingville 2506 914
Los Angeles Glendale 341 339
Long Beach 323 322
Monrovia 213 213
Palmdale 1606 1136
Pomona 373 373
Santa Clrita 73 73
Santa Monica 59 59
Madera Madera 1039 924
Marin Black Point 773 717
Point San Pedro 815 788
Mendocino Hopland FS 916 851
Sanel Valley 1056 830
Merced Kesterson 1067 934
Los Banos 1000 907
Merced 1545 1084
Modoc Alturas 2410 1252
Monterey Arroyo Seco 733 683
Castroville 552 524
King City-Oasis Rd. 864 810
Pajaro 413 395
[Table Contd...
32 Fruit Tree Physiology

Contd. Table]

Country name Station name Hours < 45°F Hours >32° and <45°F

Salinas North 832 805

Salinas South 1072 1044
Napa Carneros 751 630
Oakville 854 760
Orange Irvine 95 95
Placer Auburn 935 876
Riverside Blythe NE 550 537
Cathedral City 237 231
Indio 2 427 291
Oasis 123 123
Ripley 657 638
Temecula 193 192
Temecula East II 867 845
U.C. Riverside 239 239
Winchester 882 859
Sacra Mento Fair Oaks 690 651
Twitchell Island 935 873
San Benito San Benito 732 694
San Juan Valley 799 755
San Bernardino Barstow NE 1129 985
Big Bear Lake 2426 761
Lake Arrowhead 2257 1187
Victorville 1359 1101
San Diego Escondido SPV 617 597
La Jolla 3 3
Miramar 124 124
Otay Lake 106 106
San Diego II 28 28
San Joaquin Lodi West 824 734
Manteca 717 653
Tracy 772 722
San Luis Obispo Atascadero 1176 966
Nipomo 284 283
San Luis Obispo 338 337
San Luis Obispo West 509 501
Santa Barbara Cuyama 1444 1205
Goleta Foothills 27 27
Santa Barbara 282 281
Santa Ynez 1060 948
Sisquoc 711 694
Santa Clara Gilroy 165 165

[Table Contd...
 Chilling Requirement in Fruit Crops 33

Contd. Table]

Country name Station name Hours < 45°F Hours >32° and <45°F

Santa Cruz De Laveaga 601 578

Green Valley Road 394 382
Shasta McArthur 2472 1463
Siskiyou Tulelake FS 2504 1316
Solano Dixon 896 872
Hastings Tract East 181 181
Suisun Valley 761 703
Winters 919 819
Sonoma Bennett Valley 1327 1186
Petaluma East 904 812
Santa Rosa 962 813
Windsor 950 801
Stanislaus Denair 169 169
Modesto 1048 920
Oakdale 947 893
Patterson 788 762
Sutter Nicolaus 895 798
Tehama Gerber 950 844
Tulare Alpaugh 1428 1185
Delano 1058 994
Lindcove 814 777
Porterville 1253 1124
Ventura Camarillo 377 377
Oxnard 82 82
Santa Paula 424 424
Yolo Bryte 712 679
Davis 959 870
Esparto 514 488

Table : Varieties of stone fruits and chill units required (Steven and Swan, 2006)

Variety Chill units required

Snow Queen Nectarine 650

Snow Zee Nectarine 900
Fairlane Nectarine 700
Early O Henry Peach 900
Maycrest Peach 600
Florda Gold Peach 300
Santa Rosa Plum 150
Fuji Apple 200-300
Royal Gala Apple 300-400
34 Fruit Tree Physiology

Progression into and out of Resting State

Deciduous fruit buds gradually progress into rest untill the most intense state of
inhibition is reached. At this point, little can be done to force buds from rest.
During chilling, resting buds gradually progress out of the resting state untill bud
break occurs. The transition from endodormancy (deep rest) to bud break is not
abrupt, but a gradual occurrence (Couvillon and Erez, 1985). Wolak and Couvillon
(1976) treated non-resting peach buds (i.e. 50% of the buds broke within 2 weeks
at forcing temperatures, indicating that sufficient chilling was received to allow
for normal bud break) with thiourea which resulted in bud break following the
accumulation of 2,000 less growing degree hours (GDH) than the control. However,
since buds continued to respond to thiourea application, some degree of inhibition
remained, thus the buds were under the rest influence even though they were
sufficiently chilled to allow for normal bud break without insufficient chilling
symptoms. Buds out of endodormancy (deep rest) may force following some
degree of chilling, provided a sufficient number of GDH are accumulated
(Richardson et al; 1974, 1975). The number of GDH required for bud break
depends upon the amount of chilling received, i.e the longer the chilling period
the fewer the GDH required for bud break. Peach and pecan buds have been
observed to break in cold storage with the accumulation of few GDH (Sparks,
1993).

Movement of the Rest Influence

The rest influence is localised in the buds and can move through graft unions
(Chandler, 1960., Chandler, 1957). The growth of fully chilled scions of Beverly
Hills apple grafted to either fully chilled or unchilled apple root stock was affected
by root stock chilling. Although, fully chilled scions grafted to chilled root stocks
grew normally, those grafted to unchilled root stocks broke and produced
significantly less growth, indicating movement of the rest influence into the
chilled scions. Thus, the rest influence seems to be localized in the buds and can
move through stems and across graft unions influencing bud break in the scion.
This has economic implications and shows the need for low chilling root stocks
in deciduous fruit producing areas that experience warm winters.

Effective Chilling Temperatures

Chilling temperatures of 80C have been shown to be the most effective, in a
continuous chilling cycle, in satisfying the rest requirement. Temperatures above
or below this level provide chilling, but are less effective than 80C (Erez and
Couvillon, 1987). For instance, 120C was 46 per cent as effective as 80C in
 Chilling Requirement in Fruit Crops 35

satisfying the rest requirement and 00C was only 33 per cent as effective. However,
when chilling temperatures are interspersed with moderate temperatures (130C to
150C for 8 hours with 16 hours of chilling temperatures) during rest, there is an
enhancement of chilling response. Moderately effective chilling temperatures of
00C became as effective as 80C in satisfying the chilling requirement, when
chilled in a cycle of 8 hours of 150C and 16 hours of 00C. The most effective
moderate temperature applied in a diurnal cycle was 130C. The cyclic temperatures
were most effective when applied during the later 1/3rd of the chilling cycle, the
portion representing emergence form deep rest and the transition to the non-
resting state (Erez and Couvillon,1987). These data relating to temperature and
rest allowed for the development of a scheme for peach chilling response to
temperature based on a similar scheme proposed by Purvis and Gregory (1952)
for the vernalization of rye. The proposed scheme has two steps which explain
the effect of temperature on bud rest. The first step involves the conversion from
the unchilled to the chilled state by chilling temperatures. This stage can be
reversed by high temperatures. The second stage is not reversible and involves
the conversion, by moderate temperatures, of the unstable intermediate formed in
step 1 to a stable material, which, when accumulated to a certain level, will result
in rest completion (Erez and Couvillon, 1987).

Table : List of fruit species based on effective chilling requirement units (Luedeling
et al; 2009)

Low chilling requirement Intermediate chilling High chilling requirement

requirement

Date Pomegranate (100) Walnut (400)

Rose Peach (200) Apple (400)
Lime Apricot (350) Pear (600)
Sweet Lime Almond (400) Plum (700)
Bitter Orange Grape (100)
Orange Fig (100)
Osbeck
Lemon
Prickly Pear
Citron, Guava, Mango, Sapodilla
Pigeon Pea, Banana, Papaya
36 Fruit Tree Physiology

Chilling Negation by High Temepratures

High temeperatures (i.e. 19, 20, 240C and higher), applied at least 8 hours in a
diurnal cycle with chilling temperatures, result in chilling negation. The degree
of chilling negation depends upon the length of cycle in which the high temperature
is applied as well as the level and duration of high temperature. Applied chilling
was completely negated when 8 hours of 240C was provided in a 24 hour cycle
with 40C, but as the length of the chilling period increased, before exposure to
the high temperature, the negating effect of high temperature decreased (Erez
et al; 1979b). Bud break did not occur in the trees exposed to a 1 day cycle
(8 hours 240C and 16 hours at 40C), but a significantly greater percentage of buds
broke on trees exposed to a 3 day cycle (48 hours at 40C and 24 hours at 240C).
This was suggested a chilling fixation effect in the 3 day cycle. Bud break in the
3, 6 (108 hours at 40C and 36 hours at 240C), and 9 day (162 hours at 40C and
54 hours at 240C) cycles did not differ from that of the continuous chilling
control which received the same number of chilling hours. Thus chilling
accumulated only during the 20 to 40 hours prior to the onset of the high
temperatures was susceptible to high temperature negation on the other hand,
temperature of 200C intersperssed for 2 or 4 hours in a diurnal cycle with 40C
resulted in 15 and 17 per cent, respectively, enhancement of the chilling effect.
However, longer periods of 200C resulted in a 19 per cent (6 hours of 200C) and
70 per cent (8 hours of 200C) reduction in bud break over the continuous chilling
control. Two, 4, 6 and 8 hours of 240C interspersed in a diurnal cycle with 40C
resulted in chilling negation of 14, 45, 82 per cent and total negation, respectively,
of the applied chilling. The chilling negation effect of high temperature is greater
when occuring during the later ¼ of the chilling period than earlier (Couvillon
and Erez, 1985). A modulation factor was developed for level and duration of
high temperature in a daily cycle. Temperature of 15, 18, 19, 20 and 210C
interspersed for 8 hours in a daily cycle with 40C for 16 hours resulted in
modulation factors of 1.9, 0, -0.8, -1.5 and -1.9, respectively (Couvillon and Erez,
1985). Chilling reversion by high temperature, follows the temperature effect on
chilling described in the scheme (Erez and Couvillon, 1987) modelled after that
developed by Purvis and Gregory (1952) for the vernalization phenomenon.

Heat Accumulation, Stress and Rest

It is clear that release of buds from rest is a function of both chilling and heat.
Smith (1907) reported that unchilled peach buds from high chilling peach cultivars
grown in southern California, broke in August. He accredited this phenomenon
to the accumulation of heat. Waring (1956) reported that immersing resting twigs
in a water bath at 450C (113 F) had a rest breaking influence. However, in this
 Chilling Requirement in Fruit Crops 37

study, the reduction in oxygen levels which occurred due to immersion of the
buds in water could have had a rest breaking effect, thus it is difficult to accredit
the bud release response obtained by Waring to heat alone (Erez and Keys, 1980).
Chandler (1960) who reported that temperatures of 450C will release buds from
rest presented much more definitive data than did Waring and accredits late
summer and early autmn bloom of fruit species to rest release by exceptionally
high summer temperatures. In his study, Chandler exposed unchilled trees of
Beverly Hills apple to ambient temperatures and heated others for 6 hours at
450C only 13.6 per cent of the unheated buds broke while 78.3 per cent of those
heated broke and grew. This was verified in a southern California apple orchard
where temperatures during the summer of 1955 resulted in the release from rest
of insufficiently chilled buds on apple trees which bloomed on September 28th
(Chandler, 1960). Sparks (1993) developed a model to describe bud break in
pecan. He found that bud break varied inversely with chilling and pecan bud
break could occur without chilling, provided sufficient heat occurred. Thus, sparks
suggested that long chilling species/ cultivars actually have a long heat requirement
which can be modified by chilling. This reasoning becomes more plausible in
light of data (Key et al; 1982) which show that many plants respond to heat stress
by synthesising a new set of proteins known as heat shock proteins. These proteins
are formed, in soyabean, during exposure to 39-410C, can be detected within
3-10 minutes after exposure to high temperature, and are accompanied by rapid
synthesis of mRNAs for these proteins. Of course these data related to proteins
produced, in soyabean, to protect plants form high temperature injury, but it is
conceivable that other unknown proteins may be produced in plants exposed to
high temperatures which may override the rest influence. Chandler (1960) reported
bud release form rest following exposure of plants to 6 days at 450C, a temperature
level above that reported for heat shock protein production in soyabean
(Key et al; 1982). This may be an area worthy of attention and could add to our
knowledge relating to rest release in plants.
It has been found in many fruits crops that the stress helps to release buds
from the rest. It seems that many types of stress (i.e. heat, drought, toxicities, cold
damage, etc.) will result in a lowering or replacement of chilling requirement
resulting in early bloom during the subsequent year. In some cases, rest release
due to induced stress is used for economic advantage in fruit production.
Replacement of chilling by stress (induced by defoliation) allows for the
commercial production of long chilling apples in areas that experience high winter
temperatures. The long chilling apple cultivar Rome Beauty is grown in Java by
leaf removal following fruit harvest which induces flowering for the next crop
(Janick, 1974). Trees are continuously fruited year after year. In temperate areas,
trees treated in this manner generally enter deep rest following a few defoliation
38 Fruit Tree Physiology

cycles. Probably the high temperature experienced in tropical Java play a role in
rest release as well as defoliation of the mature trees. Although these stress
related treatments allow for bud break in Rome Beauty apple without chilling,
other factors such as the use of low chilling root stocks could also be involved.

Pre requisites for construction of a model

Preparation of a fruit tree model, or another crop simulation model, cannot be
undertaken unless some background knowledge of the system is available. It
might be said that in order to develop useful and robust models one must have:
● Reasonable conceptual understanding of the physiological processes involved
● Imaginative, quantitative thinking
● An extensive agricultural/ biological database

MODELS FOR FRUIT TREES

Models are extensively used in research, in the social sciences (e.g economics)
as well a the physical, chemical and biological sciences. As a preliminary definition
one could think of a model as an attempt to describe a certain process or system
through the use of a simplified representation, preferably a quantative mathematical
expression, that focuses on a relatively few key variables that control the process
or system.
Currently available agricultural crop models may be found in two areas:
A. Climate based models : These were originally designed to correlate and
predict a specific biological phenomenon with climate data, mostly temperature
data. Models using cumulative heat or chilling units to predict spring growth
and phenology, to predict fruit ripening dates or to predict the satisfaction of
chilling requirement in perennial dormancy and their more sophisticated
descendants belong to this category (Ben Mechlia and Carroll, 1989., Bustan
et al; 1996., Erez et al; 1990., Fuchigami and Nee, 1987., Hall and McPherson,
1997; Reuther, 1973; Salinger et al; 1993). Modelling of vegetation and crop
water use is driven by climatic variables (Allen et al; 1998). More recently
these simplified models have often become sub-models of more comprehensive
plant or crop models discussed below.
B. Ecological systems : Models have been and are still major tools in the hands
of ecologists in their efforts to identify and quantify the state variables which
affect the ecosystem and the interactions of its components. Agri-systems
have many features in common with ecosystems but they are much simplified,
heavily managed, often limited in interactions amongst species and as such
 Chilling Requirement in Fruit Crops 39

are outside the scope of ecologists. However, a recent broadening of views

of the interactions of agri-systems with the surrounding environment has led
to more common ground (Room et al; 1996).
Many fruit and nut trees require cold temperatures during the winter to
overcome their seasonal dormancy (Erez, 2000). Most fruit and nut species that
evolved in temperate or cool sub tropical climates have such chilling requirements
that need to be fulfilled each winter to achieve homogeneous and simultaneous
flowering and regular crop yields. In order to select appropriate fruit and nut
species and cultivars for the climate of a given site, researchers have developed
chilling models, which convert temperature records into a metric of coldness
(Bennett, 1949., Richardson et al; 1974 and Erez et al; 1990). Using the same
chilling model to quantify both a cultivar’s chilling requirement and the amount
of winter chill available at a given location enables growers to predict whether
the cultivar will perform well under the specific temperature conditions of their
sites. Chilling models also constitute tools to understand and manage the inter
annual variation in the time, at which tree crops complete their dormancy. Many
growers use estimated winter chill to determine the time line for certain
management measures, such as the spraying of rest-breaking chemicals, or to
predict their yield potential for the season.
What remains unclear is how well different chilling models predict winter
chill, when temperature conditions deviate from historic patterns. Since
overwhelming scientific evidence suggests that the global climate is warming
(IPCC, 2007), most growing regions might experience substantial temperature
increases in the near future, with likely consequences for available winter chill.
In California, the strongest warming trend has been identified for daily minimum
temperatures during the winter months (La Dochy et al; 2007), indicating that
winter chill might be strongly affected by climate change. Fruit and nut orchards
might thus be among those agricultural systems in California, and probably in
other parts of the world, that are most vulnerable to the environmental stress
invoked by climate change. While most annual crops are primarily affected by
temperature changes during the spring and summer months, the sustainability of
an orchard depends on the prevalence of a certain temperature regime throughout
the entire year. Failure to meet the climatic requirements of tree crops can have
devastating consequences for growers, since in order to be economically viable,
orchards must remain productive over several decades to pay off the investments
needed for orchard establishment. It is thus crucial for the sustainability of an
orchard operation to accurately estimate the effect of changing climatic conditions
on the biology of the cropping systems. This is especially important during the
dormant season, since insufficient chilling can severely compromise fruit and nut
yields.
40 Fruit Tree Physiology

The implications of climate change for winter chill have occasionally been
investigated (Baldocchi and Wong, 2008), but no studies have compared the
effects of temperature increases on winter chill, when quantified with different
chilling models.

Common Chill Accumulation Models

450F and under model: 1 hour <= 450 F =1.0 chill unit
32-450F model: 1 hour between 32 and 450 F =1.0 chill unit
Utah model :1 hour below 34 degrees F =0.0 chill unit
1 hour 35-36 degrees F = 0.5 chill units
1 hour 37-48 degrees F = 1.0 chill units
1 hour 49-54 degrees F = 0.5 chill units
1 hour 55-60 degrees F = 0.0 chill units
1 hour 61-65 degrees F = – 0.5 chill units
1 hour >65 degrees F = – 1.0
Chilling hours model : The Chilling hours model (sometimes referred to as
Weinberger Model; Bennett, 1949 and Weinberger, 1950), as originally proposed,
simply calculates the number of hours, when the temperature (T) is below 7.2 °C
(450 F, sometimes converted to 7 or 7.22°C). It soon became apparent that freezing
temperatures did not contribute to winter chill accumulation, leading to the
exclusion of such temperatures. At a given time t during the dormancy period (in
hours after a fixed starting time at the beginning of the dormancy season), the
number of accumulated chilling hours (CH) is thus given as:

Utah model : (Richardson et al; 1974): The second model tested was the
Utah model which is similar in concept to the chilling hours model, but assigns
different weights to different ranges of temperatures. This approach reflects research
showing that chilling efficiency varied with temperature, including negative chilling
accumulation by high temperatures. The exact definition of the temperature steps
used in this model has been modified to suit the climate of different regions and
but in California only the original version is currently used:
 Chilling Requirement in Fruit Crops 41

Scientists noticed that under controlled condition’s temperatures of 6°C attributed

most towards the completion of tree rest than any other temperature. So, one hour at
6°C was given the value of 1 chill unit (CU). They also found that lower and higher
temperatures could counter-act those effects. Scientists at Utah State University in
Logan developed the following model to help calculate chill units received.
● 1 hour below 1.4°C = 0.0 chill unit (CU)
● 1 hour between 1.5-2.4°C = 0.5 chill units (CU)
● 1 hour between 2.5-9.1°C = 1.0 chill units (CU)
● 1 hour between 9.2-12.4°C = 0.5 chill units (CU)
● 1 hour between12.5-15.9°C = 0.0 chill units (CU)
● 1 hour between 16-18°C = - 0.5 chill units (CU)
● 1 hour over 18°C = - 1.0 chill units (CU) chill unit
Positive Utah model : In addition to the models typically used in California,
one modified version of the Utah model is worth considering, because it has been
shown to perform well in regions with a warmer climate than present-day
California, such as South Africa. In this Positive Utah model, the negative
contributions of warm temperatures to accumulated chilling were removed from
the original equation of the Utah model:
t
Utah ti TU, with TU
i 1
T 1.4 C :0
1.4 C T 2.4 C : 0.5
2.4 C T 9.1 C :1
9.1 C T 12.4 C : 0.5
T 12.4 C :0
42 Fruit Tree Physiology

Dynamic model (Fishman et al; 1987) : The Dynamic Model is currently

used in Israel and South Africa. It has also been adopted by cherry growers in
California, found to be superior to currently used models in Spain (Ruiz et al;
2007) and suggested for general use in Chile (Perez et al; 2008). The Dynamic
model is currently the only model that explains experimental evidence from
controlled temperature studies in Israel. The main findings from these trials were
that moderate temperatures enhanced previous chilling, and that only recently
accumulated chilling was subject to negation. The Dynamic model postulates that
winter chill accumulates in a two-step process. Initially, cold temperatures lead
to the formation of an intermediate product. Once a certain quantity of this
intermediate has accumulated, it can be transformed into a so-called chill portion
by a process requiring relatively warm temperatures. The equations used to calculate
chill portions are more complex than the other models. While they are difficult
to derive from the original publications, we extracted them from a spreadsheet
used by plant physiologists to calculate chill portions.

The experimentally derived constants slp, tetmlt, a0, a1, e0 and e1 were set
to 1.6, 277, 139500, 2.567 × 1018, 12888.8 and 4153.5, respectively, according to
standard practice in horticultural applications. TK is the measured hourly
temperature in Kelvin, while t denotes the time during the season (in hours), with
t0 being the starting point of chilling accumulation.
 Chilling Requirement in Fruit Crops 43

Other Utah models : In the early 1970s, researchers at Utah State University
began to develop a series of models to predict chill unit accumulation necessary
for the completion of rest and the growing degree hours (GDH) necessary for
spring flower bud development. The models subsequently developed for field
prediction utilises the most commonly available temperature data, maximum and
minimum temperatures. These models relate instrument shelter temperature to
actual bud temperatures.
(a) The Utah chill unit model : It quantifies temperatures necessary to complete
the rest cycle of deciduous fruit trees (Richardson et al; 1974). One chill unit
is defined as one hour at optimum temperature needed to meet the chilling
requirements. According to this model, temperatures below 00c (as measured
in an instrument shelter) do not contribute to accumulated chill units and
temperatures above 14.50c reduce accumulated chill units.
(b) The ASYMCUR growing degree hours medel : The chill unit and growing
degree hour models are usually used in combination, once the chill unit
accumulation indicates that rest has been completed for a cultivar, the model
begins accumulating growing degree hours, as defined by the ASYMCUR
fruit- tree model. In general, two equations (1 and 2 below) describe the
asymmetric curvilinear model. Equation 1 defines the curve between the
base and optimum temperatures and equation 2 defines the curve between
the optimum and critical temperatures. The cardinal temperatures for all
deciduous fruit trees investigated to date are: TB=4 0c, TU= 25 0c and TC=
36 0c, where TB is the base temperature below which little or no growth or
development will occur, TU is the optimum temperature at which the maximum
rate of growth and/or development will occur, and TC is the critical
temperature above which little or no growth will occur.
GDHC = TU–TB)/2) (1 + cos ( π + π(TH–TB)/(TU–TB))) ......(i)
GDHC = TU–TB)/2) (1 + cos ( π/2 + π2 (TH–Tu)/(Tc–Tu))) ......(ii)
where :
GDHC = accumulation of growing degree hours for an hour at temperature
= to TH
(c) Maping potential production areas : To determine whether the climate of a
specific site could support deciduous fruit production of a particular cultivar,
normal temperature data for the sites or for the nearest site with similar
climate can be used in the model. If winter hardners, incomplete chilling,
insufficient growing season or late spring freeze problems were likely, other
cultivars or cultural practices would be needed.
44 Fruit Tree Physiology

Mean temperature model : The mean temperature model uses mean winter
(December and/or January) monthly temperatures to estimate accumulated chilling
units. Researchers in Georgia and Florida independently developed a relationship
between the mean monthly temperature of their coldest month(s) and total chill
unit accumulation. Combining data from both studies (Fig.), the Stone Fruit
Breeding Program at Texas A&M University developed a method to estimate
chill accumulation which has demonstrated to be accurate for estimating chill
accumulation in Texas from the lower Rio Grande Valley up to the Red River, and
should work well throughout the south eastern United States.

Fig.: Total chill unit accumulation based on mean winter temperatures.

Chilling accumulation, determined with this model, has been tested and
compared to peach tree behavior at Stephenville, Fredricksburg, College Station,
Yoakum, and Weslaco, TX. The coldest month or months are used for the
calculation. In low chill regions (regions where average January temperature is
59-630F) where January represents the dormancy season, January mean temperature
is most accurate for estimation. In high chill regions (regions where average
January temperature is below 480F) a mean December-January temperature is
recommended. In medium chill regions (regions where average January temperature
is 48-580F) January mean temperature has been best for calculating chill
accumulation except in years when mean temperatures between December and
January differed by more than 6 degrees F. In this case, the December-January
mean was more accurate.
 Chilling Requirement in Fruit Crops 45

CONCLUSION
Rest and dormancy have long been of interest to pomologists and are well
researched areas. Although the cause of rest has been elusive and much progress
has been made in describing the response of resting buds to the temperature
experienced in nature. Possibly, in the near future, new technology such as
molecular biological techniques will allow for elucidation of the rest phenomena.
A comparison of genetic make up of plants of the same species with or without
a cold/ heat requirement could lead to identification of the gene(s) responsible for
rest in these plants. Of course it would also be interesting to determine if proteins,
capable of overriding the rest requirement, are produced in resting buds in response
to high temperature exposure.
The four chilling models tested predicted different rates of change in winter
chill in California. While all models showed a decline, winter chill loss predicted
by the currently common models, the Chilling hours model and the Utah model,
was much greater than losses predicted by the Positive Utah model and the
Dynamic model. While available information is insufficient for deciding which
model is most suitable for describing imminent changes in winter chill, research
on chilling models needs to be intensified, in order to prepare fruit and nut
growers in the world and other for the agro climatic changes that are occurring
globally at a faster rate due to climate change.

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growing regions of California. Climatic change 87: 515-516
Ben MN and Carroll JJ (1989) Agroclimatic modeling for simulation of phenology,
yield and quality of crop production. II. Citrus model implementation and
vertification. Int J Biometeorol 33: 52-65.
Bennett JP (1949) Temperature and bud rest period. Calif Agri 3: 9- 12.
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48 Fruit Tree Physiology

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 3

HEAT UNITS AND

THERMOPERIODISM

I n the absence of extreme conditions such as unseasonal drought or disease,

plants grow in a cumulative stepwise manner which is strongly influenced by
the ambient temperature. Growing degree days take aspects of local weather into
account and allow gardeners to predict (or, in greenhouses, even to control) the
plants pace towards maturity.
Unless stressed by other environmental factors like moisture, the development
rate from emergence to maturity for many plants depends upon the daily air
temperature. Because many developmental events of plants and insects depend on
the accumulation of specific quantities of heat, it is possible to predict when these
events should occur during a growing season regardless of differences in
temperatures from year to year. Growing degrees (GDs) is defined as the number
of temperature degrees above a certain threshold base temperature, which varies
among crop species. The base temperature is that temperature below which plant
growth is zero.
Temperature plays an important role in growth and development of plants.
The temperature dependent growth and development is often described as heat
units or growing degree days (GDDs). Growing degree days (GDD) also called
growing degree units (GDUs), are a heuristic tool in phenology. GDD are a
measure of heat accumulation used by horticulturists, gardeners, and farmers to
predict the date that a flower will bloom or a crop reach maturity. The transition
from one developemental stage to another requires energy which is provided by
heat and the amount of heat required for this transition remains constant from
year to year, but depending on weather conditions, the amount can vary. Each
plant has a minimum base temperature or threshold below which development
50 Fruit Tree Physiology

does not occur. The difference between the average of the daily maximum and
minimum temperatures and a certain critical basal or developmental threshold
temperature is used to calculate the daily heat units. The threshold temperature
varies among species. The daily heat unit values can be summed for the season
to derive cumulative seasonal heat units. The following equation is used to calculate
heat units.

(Balogh and Pfeiffer, 1998)

or (DD) = Average daily temp. - Base Temp. = (max. + min.) / 2 - Base temp.
GDDs are typically measured from the winter low temperature. Any
temperature below Tbase is set to Tbase before calculating the average. Likewise,
the maximum temperature is usually capped at 30°C because most plants and
insects do not grow any faster above that temperature. However, some warm
temperate and tropical plants do have significant requirements for days above
30°C to mature fruit or seeds. For example, a day with a high of 23°C and a low
of 12°C (and a base of 10°C) would contribute 7.5 GDDs.

A day with a high of 13°C and a low of 10°C (and a base of 10°C) would
contribute 1.5 GDDs.
13 10
10 1.5
2
Fruits are classified into tropical, sub tropical, or temperate based on climatic
requirements and have varying amounts of climatic adaptations. They can grow
only in their specific areas/ zones e.g mango and pineapple can grow in tropical
while citrus in sub tropical climate and apple grows best in temperate climate.
However, some fruit crops are day neutral such as strawberry and some apple
cultivars and can grow in low chill sub tropical climates while others in high chill
temperate climates. Some fruits have more specific adaptation limitations than
others as seen by their specialized areas of production. Climatic adaptation precedes
breeding for commercial fruit qualities. Without climatic adaptation, the breeder
may not be able to make hybrids and cannot adequately fruit and evaluate them.
Once the breeder finds climatic adaptation for tree growth and fruiting, selection
for more specific climatic requirements is possible, and primary attention can
then be given to fruit characteristics necessary for making the crop economically
viable. Photoperiod, light intensity, chilling and heat units, soil type, and
 Heat Units and Thermoperiodism 51

temperature and water tolerance are adaptation factors for tree growth, flower bud
formation, flowering, and fruit growth and maturation (Sherman and Beckman,
2003). The degree days or heat summation can be used to predict the harvest
dates of fruits and vegetables e.g accumulated heat units in apple for optimum
maturity are computed by calculating the time in relation to temperature above
a certain base temperature i.e 500C. Hence a day with an average temperature of
680F (200C) would provide 18 degree days F (10 degree days C) of heat units.
A day with an average temperature of 410F (50C) would provide zero degree days
of heat units.
Table: Heat unit requirement in terms of degree days of some fruits (Verma and
Joshi, 2006).

Fruit Cultivar Base temperature Degree days

Apple Red Delicious 500F 1659-1705

Pear Bartlet 450F 2240-2296
Grape Thompson Seedless 500F 1600-2000
Bhokri 3284
Gulabi 3508
Banglore Blue 3562
Mango Bangnapally 180F 1426 ± 45
Ber Gola 450F 1980-2236
Kaithali 2236-2568
Umran 2516-2920
Litchi Shahi 150F 813
Banana 9.80F 1930

Sindh province in Pakistan has considerable potential for diversifying into

new horticultural crops where 100-550 chill units (<7.2°C) and 3700-4500 heat
units (>12.8°) per year are available. Nine agro-climatic zones were identified to
be suitable for these new crops. Low chill crops like peaches, plums, apples,
pears, persimmons, pomegranates, grapes, figs, zizyphus, almonds and pecans
were tested for their suitability at a site receiving 300 chill units. It was found that
peaches, followed by pomegranates and grapes, could be grown successfully
while apple production was erratic (Panhwar et al; 1995).
Models like chill unit and flower bud model are used to find the suitability of
crops to be grown in a particular region. Chill unit model calculates the completion
of rest (endodormancy) from the accumulation of temperature dependent chill unit
in temperate fruits while flower bud phenology model is a growing degree-hour
model based on the heat units required for buds to develop to their different stages
following rest completion. The models are accurate for temperate continental
52 Fruit Tree Physiology

climates, but not for sub tropical climates, where the models can conflict. Models
must be continuously developed and refined to give better fits with observed data
because the factors like photoperiod dependency and cultivar effects vary.

Uses of Heat Units or Growing Degree Days (GDDs)

1. Help in predicting the suitability of a site for a particular fruit crop.
2. To eliminate areas that can not grow a particular fruit crop.
3. Help in selection of variety for a particular locality.
4. To predict harvest date.
5. To predict pest outbreak during growing season.
6. To estimate the growth stages of crops.
7. Predict best timing of fertilizer or pesticide application.
8. To estimate the heat stress on crops.
9. To plan spacing of planting dates to produce separate harvest dates.
Crop specific indices that employ separate equations for the influence of the
daily minimum (night time) and the maximum (day time) temperatures on growth
are called crop heat units (CHUs).
Table : Degree day accumulations for selected Ohio sites (http.//ipm.osu.edu/fruit/
index.html).

Actual DD accumulations Forecasted degree day accumulations

Location Base 43° F Base 50° F Base 43° F Normal Base 50° F Normal

Akron 2591 1767 2817 2579 1944 1736

Canton
Cincinnati 3030 2109 3285 3265 2315 2293
Cleveland 2593 1780 2745 2523 1896 1677
Columbus 3156 2226 3368 2859 1954 1964
Dayton 2905 2040 3153 2927 2238 2032
Elyria 2729 1917 2961 2657 2100 1814
Fremont 2475 1698 2735 2579 1909 1763
Mansfield 2509 1690 2745 2555 1876 1719
Norwalk 2611 1807 2845 2414 1992 1699
Toledo 2707 1894 2905 2511 2050 1699
Wooster 2669 1838 2909 2442 2030 1614
Youngstown 2390 1598 2608 2383 1767 1571
 Heat Units and Thermoperiodism 53

Phenology

Range of degree day accumulations

Coming events Base 43° F Base 50° F

Codling moth 2nd flight peak 1587-3103 1061-2212

Apple maggot flight peak 2033-2688 1387-1804
Obliquebanded leafroller 2nd flight begins 2124-3040 1412-2076
Oriental fruit moth 3rd flight begins 2172-2956 1553-2013
Peach tree borer flight subsiding 2230-3255 1497-2309
Red banded leaf roller 3rd flight begins 2389-3113 1722-2209
Spotted tentiform leaf miner 3rd flight peak 2415-3142 1728-2231
San Jose scale 2nd flight subsides 2494-3257 1662-2303
Red banded leaf roller 3rd flight peak 2514-3225 1818-2625
Oblique banded leaf roller 2nd flight peak 2634-3267 1789-2231

HEAT UNITS IN FRUIT CROPS

Apple : Development of apple bud is regulated by temperature for both the
satisfaction of the chilling requirement to break dormancy and the accumulation
of growth units to resume growth. Regulation of temperature in apple orchards
can be achieved by water management to suppress bud development, protect
against ice formation, and ameliorate occurrence of sub freezing temperatures.
Table : Growth stages and growth units for Red Delicious apples (http.// ipm.edu/
fruit / index. html).

Stage Growth units oF Growth units oC

1 Silver tip 3710 2061

2 Green tip 4580 2544
3 Half-inch green 5580 3100
4 Tight cluster 7090 3939
5 First pink 8740 4856
6 Full pink 9740 5394
7 First bloom 11110 6172
8 Full bloom 12480 6933
9 Post bloom
10 Harvest 98000 54440

Effect of daily temperature on cell multiplication and fruit growth leads to

increased cell number in reproductive bud in the areas with most chilling units,
54 Fruit Tree Physiology

while the lowest number in areas with lowest chilling in Royal Gala apples.
Increased fruit development and improved fruit size is achieved when higher
temperatures occur about 30-40 days from full flowering (Greybe et al; 1998).
Grapes : Heat units are calculated by multiplying the mean daily temperature
over and above 100C by the number of days. Certain amount of heat units is
required for different growth stages and for fruit ripening it varies from 1600-
3500 degree days. Fewer heat units are required by early ripening cultivars and
those adapted to cool, short season locations than cultivars that ripen later and are
adapted to warmer locations. Average heat units received by selected Idaho
communities’ are shown in the following table (Barney et al; 1995)

City Heat units City Heat units

Ashton 1,300 Moscow 1,650

Black foot 2,000 Mountain Home 2,700
Boise 2,650 Payette 2,900
Burley 2,200 Pocatello 2,100
Coeur d’Alene 1,600 Rexburg 1,700
Idaho Falls 1,800 Salmon 1,900
Kellogg 1,800 Sandpoint 1,500
Lewiston 2,700 Stanley 500
Malad 1,900 Twin Falls 2,000
McCall 950

Seedless grapes

Cultivar Color Cold hardiness degrees F Heat units Ripens

Canadice Red -15 to -25 1500-2500 3-4

Concord Seedless Bluish black -15 to -25 2000-2500 5
Himrod White 0 to -15 1500-2500 3-4
Interlaken Seedless White +5 to -5 1500-2500 3
Reliance Red -15 to -25 1500-2500 3-4

North American grapes

Cultivar Color Cold hardiness degrees F Heat units Ripens

Campbell’s Early Red -15 to -25 1500-2500 3-4

Catawba Red -10 to -20 2500-3000 5
Concord Bluish black -15 to -25 2000-2500 5
Delaware Red 0 to -10 2000-2500 5
Niagara White -5 to -15 2000-2500 5
Steuben Bluish black -10 to -20 2500-3000 5
 Heat Units and Thermoperiodism 55

Hybrid grapes

Cultivar Color Cold hardiness Heat units Ripens

Aurore White -5 to -15 2000-2500 3

Chancellor Bluish black 0 to -10 2000-2500 4
Chelois Bluish black +5 to -5 2000-2500 4
De Chaunac Bluish black 0 to -10 2000-2500 4
Foch (Marechal Foch) Bluish black -5 to -15 2000-2500 3-4
Rosette Bluish black -5 to -15 2000-2500 5
Seibel Pink +5 to -5 2000-2500 3
Verdelet White to yellow +5 to -5 2000-2500 2-4

European grapes

Cultivar Color Cold hardiness Heat units Ripens

Cabernet Sauvignon Purplish black +10 to 0 2000-3000 5

Chardonnay White +5 to -5 2000-3000 3
Gewürztraminer Pinkish red +10 to 0 2000-3000 3
Pinot Noir Blue +10 to 0 2000-2500 3-4
Sylvaner White +5 to -5 2000-3000 3-4
White Riesling White +5 to -5 2000-3000 5

Date palm : The mean temperature between the period of flowering and
ripening of the fruit should be above 210C rising to 270C or higher for at least
one month. For successful fruit maturation, nearly 3000 heat units are required
and are available in most of the north-western districts of India during end February
(time of flowering) to July, hence these areas are suitable for date palm cultivation
(Ashwani, 2010). Keena (2003) found that dates need 2000 heat units from
flowering (August- September) to fruit maturity (February- April). The average
daily maximum temperatures in leading date growing countries ranges from 27
to 32°C and dates can withstand temperatures as high as 50°C (Burt, 2008).
Swingle (1904) considered 18°C as base temperature (since the flowering process
does not commence below 18°C) and added the daily temperature maxima for a
period of 184 days. He obtained the value of heat units for several date plantations
as shown in the table.
56 Fruit Tree Physiology

Table : Value of heat units at various date growing areas in Algeria and Iraq (Zaid
and Wet, 2008).

Plantation/Country Heat units (°C) Plantation/Country Heat units (°C)

Laghouat/Algeria 2,327 Touggourt/Algeria 3,666

Biskra/Algeria 3,049 El Golea/Algeria 3,990
Ayata/Algeria 3,295 Baghdad/Iraq 3,898

Table : Heat units at various date growing countries.

Plantations/Country Heat units (°C) Plantations/Country Heat units (°C)

Elche/Spain 792 Nema/Mauritania 1,903

Laghouat/Algeria 990 Basra/Iraq 1,872
El-Kantara/Algeria 1,170 Kidal/Mali 1,841
Touggourt/Algeria 1,854 Kayes/Mali 1,980
Bechar/Algeria 1,620 Agadez/Niger 1,827
Atar/Mauritania 1,860 Bilma/Niger 1,860
Kankossa/Mauritania 1,836 Largeau/Chad 1,860

Heat unit values can help in three various ways in the cultivation of date palm as:
1. To find the suitability of a site for growing a productive date palm.
2. To eliminate areas that cannot grow date palm, and
3. To help in variety selection.
Table : Accumulated heat units for date production (Burt, 2008).

Sum of heat units(°C) from Comments

flowering to harvesting

Less than 2000 No varieties ripen

2000 to 2250 Early varieties ripen
2250 to 2750 Many varieties will ripen
2750 to 3250 All varieties ripen, Deglet Noor of variable quality
Greater than 3250 All varieties ripen, Deglet Noor of top quality

Mango : Heat units work on the basis that the fruit needs a certain amount
of degrees temperature to enable it to mature. The use of heat sums allows
growers to make fairly accurate decisions on when to start dry matter testing the
fruit and aids in predicting harvest dates. A combination of heat units, dry matter
levels and internal colour will give the grower the best indications of when
harvest should commence (Greg Owens, 2004). Total heat unit requirements for
commercial mango varieties Alphanso, Kesar and Ratna have been calculated by
Burondkar et al. (2000) who found that Alphanso requires minimum duration
 Heat Units and Thermoperiodism 57

(111-93 days) and heat units (718-701 DDs), Kesar (818- 98 days; 779-799 DDs)
and Ratna (127-122 days; 840-869 DDs) for proper maturation.
Citrus : The growth rate of plants depends on the amount of heat they receive.
For each species there is an optimum temperature range for growth, if no other
factors (such as water) are limiting. For citrus, the optimum temperature range for
growth is commonly considered to be 13–35°C. The hours of heat within this range
are referred to as heat units or growing degree days (GDD). Heat units can be used
in citrus to assess the suitability of a region for growing citrus, for estimating
the length of phenological (growth) stages and for predicting fruit maturity times.
It is also equally important to determine the incidence of very cold temperatures
(< –2°C) and frosts when assessing an area for growing citrus. There are several
methods used to calculate heat units. The most common method is relatively straight
forward. The heat accumulated each day is determined by adding together the
maximum and minimum temperatures and dividing the total by two to obtain a
daily average. The crop threshold for citrus of 13 is then subtracted from this average.
The final value then represents the daily heat units useful for crop growth.
Daily Heat Units = [(maximum temperature + minimum temperature) / 2] – 13.
Results are then added to determine the accumulated weekly, monthly or
yearly heat units. When calculating daily heat units all results below zero (negative
results) are not used. Additionally all maximum temperatures above 35°C
(≤ 35.1) are changed to 35. An example of how to calculate heat units using both
winter and summer temperatures is contained in table.
Table : Heat unit calculations for both winter and summer temperatures (Hardy and
Tahir, 2007).

Date Max temp Recalculated Min Average Daily heat Accumulated

max temp temp temp (max units (DHU) heat units
(for temps + min / 2) (average (negative
above 35°C) temp – base values for
temperature DHU are not
of 13) used)

Example winter temperatures

1 11 11 4 7.5 -5.5 0
2 14 14 4 9 -4 0
3 14 14 3 8.5 -4.5 0
4 10 10 -2 4 -9 0
5 12 12 -1 5.5 -7.5 0
6 13 13 2 7.5 -5.5 0
7 15 15 4 9.5 -3.5 0
8 15 15 5 10 -3 0
9 16 16 6 11 -2 0
10 14 14 5 9.5 -3.5 0
[Table Contd...
58 Fruit Tree Physiology

Contd. Table]

Date Max temp Recalculated Min Average Daily heat Accumulated

max temp temp temp (max units (DHU) heat units
(for temps + min / 2) (average (negative
above 35°C) temp – base values for
temperature DHU are not
of 13) used)

Example summer temperatures

1 34 34 20 27 14 14
2 38 35 21 28 15 29
3 38 35 22 28.5 15.5 44.5
4 40 35 22 28.5 15.5 60
5 37 35 20 27.5 14.5 74.5
6 36 35 20 27.5 14.5 89
7 35 35 19 27 14 103
8 35 35 20 27.5 14.5 117.5
9 35 35 21 28 15 132.5
10 36 35 22 28.5 15.5 148

Very high heat units of low land tropics (e.g 5000-6000 units during fruit
growth) leads to faster growth but produces poor quality fruits because of the
increased respiration and therefore net carbohydrate accumulation is less. Low
heat units delay growth and result in higher acids and lower sugar content. For
oranges, root, shoot and fruit growth and development slows considerably below
13°C (Bevington and Castle, 1986). This value is then treated as the crop threshold
and temperatures below 13°C are discounted when calculating heat units. In
Valencia and Navel oranges relationship between quality and temperature sums
(effective heat units-EHUs) for the periods the fruit were held on tree were tested
and relationship of juice acid with EHUs was stronger than the brix acid relationship
with EHUs while a linear reduction in per cent acidity with EHUs was evident
(Hotton and Landsberg, 2000). In grapefruits, water (irrigation) requirements can
be determined through heat unit based crop coefficients (Martin et al; 1997). In
Satsuma orange mulching with black polyethylene increased maximum canopy
temperatures by up to 1.70C in spring and early summer, but in winter the canopy
temperatures were lower than for unmulched trees. Soil temperatures were up to
60C higher than those without mulch. In reflective mulch treatments the
accumulation of heat units from flowering until harvest was highest while black
polyethylene had lowest values. However, bud break was advanced and vegetative
growth enhanced to a greater extent under black polyethylene than under reflective
foil (Richardson et al; 1993).
 Heat Units and Thermoperiodism 59

Raspberry : Climatic factors (solar radiation, daylength, water supply, growing

degree days (GDD), cummulative heat units (CHU), soil and air temperatures)
were monitored to determine which factors could account for the variability in
yield of 3 primocane fruiting red raspberry cultivars. It was observed that soil
temperature and water supply had a major influence, while daylength, solar
radiation and above ground temperature (i.e air or CHU) had a lesser influence
on these components. Soil temperature had the largest influence during April and
May, while water supply was equally influential at all times during the season.
Air temperature and solar radiation had their largest influence during the period
of flower initiation and development (i.e. June and July), while day length was
most influential from June to October. Berry count, weight and yield had the
highest frequency of associations among the climatic factors, indicating the
complexity of the association between these yield components and climate. Total
number of nodes/ cane, length of the fruiting section/ cane and the harvest period
showed the fewer number of associations. Autumn Bliss was less sensitive to
climatic variation than Heritage or Redwing. The anomalous behaviour of Redwing
was usually related to air or soil temperatures (Prive et al; 1993). Effects of
temperature on time of harvest and cane development were investigated in cultivars
Heritage and Redwing in Minnesota, New York, Michigan, Ontario and British
Columbia. The harvesting of Heritage before Redwing began at British Columbia
and New York while in the remaining locations, harvest of heritage began after
Redwing. Heat unit accumulation and latitude data were not sufficient to explain
the differences in harvest date between the different cultivars and locations. In
Minnesota, parameters of cane height, and cumulative yield were collected. Cane
height was correlated with accumulated heat units for both cultivars. However,
the difference in yield over years for each cultivar was not due to differences in
accumulation of heat units. Changing the pattern of heat unit accumulation may
influence the cane height but the timing of harvest may not necessarily be achieved
by altering the accumulation of heat units (Hoover et al; 1989).
Peach : The time of bud development is crucial for the potential persistence
of the reproductive organs. Vegetative buds need heat units to break them while
floral buds respond differently to higher temperature in peach and nectarine.
Competition for water and nutrients among the floral parts affect the floral and
fruit drop. Cool climate favours quicker flower development but in warm conditions
leaf production may advance to the extent that flowering will occur concomitantly
with vigorous vegetative growth (Erez et al; 2000). Three peach cvs. San Pedro,
Y9/106, Rubidoux, at different stages of flowering and fruiting were observed
and chilling units were recorded according to the total hours
< = 7.20C degree days and were 275, 294, 323 for San Pedro, Y9/106, and
Rubidoux, respectively from bud swelling. According to the total hours < =100C
degree days the heat units were 632, 697, 837 (El et al; 2002).
60 Fruit Tree Physiology

Table : Average fruit development, degree days and growing degree hours for
peach cultivars (Marra, 2002).

Cultivar Fruit development Degree days (DD) Growing degree hours (GDH)
period (FDP-days)

Maycrest 72 6393 227791

Mayglo 90 8037 285057
E LADY 123 14372 445920
Fantasia 137 16917 496254
O’Henry 150 19774 568934

Apricot : The heat requirements for flowering as determined by Ruiz et al.

(2007) in 10 apricot cultivars ranged between 4078 and 5879 growing degree
hours (GDH). The cold units (CU) and heat units (HU) required for the flowering
of 13 apricot cultivars grown in Abaran (Spain) were observed by Andres and
Duran (1999). The end of the dormancy period was established on the basis of the
dry-weight increase of flower buds. The tests in which the dry weight was measured
after forcing the flower buds at 200C were more precise in detecting the beginning
of the reactivation phase than those in which they were not forced. It was concluded
from the results that coinciding flowering periods of different cultivars is no
indication that their cold and heat requirements are similar, as a clone with low
cold and high heat requirements may flower at the same time as a clone with high
cold and low heat requirements. Heat requirement from blooming to maturing in
15 apricot cultivars grown in the Belgrade region was determined by the Growing
degree days method (GDD), using 8 year phenological data. The best lower
threshold temperature varied from 0°C in cultivar Kishiniev Early to 7°C in cultivar
Roxana. The number of days from full bloom to maturity ranged from 82 to 112.
When 3°C was taken as a base temperature (the mean value of all cultivars studied),
the accumulated GDD between these two phenological stages varied from 1056 in
Senetate to 1648 in Kecskemet Rose (Milatovic et al; 2008).
Banana : The growth and development potential of 6 banana cultivars (Dwarf
Cavendish (DC), Williams (W), Chinese Cavendish (CC), Valery (V), Grand
Nain Central America (GNCA) and Grand Nain Israel (GNI)) in the first crop
cycle were evaluated and it was observed that heat units above 14°C required
from planting to flowering and from flowering to harvest were 2152 and 1388,
respectively. The optimal mean temperature for relative growth rate for all 6
cultivars was 24°C, and for net assimilation rate the optimal mean temperature
was between 24 and 26.6°C, depending on cultivar. The cultivars DC and GNI
are more suited to warm areas and W, GNCA and CC are more suited to cooler
areas based on their various growth parameters (Morse et al; 1996).
 Heat Units and Thermoperiodism 61

Pineapple : Based on heat unit model two phases of fruit development were
distinguished: the time from induction of fruit development with a growth regulator
(forcing, DAYFOR) to opening of the first flower (DAYFF), and from DAYFF
to harvest (DAYH50, the date when 50 per cent of the fruits were one-third
yellow). From DAYFOR to DAYFF, predictions were based on the accumulation
of heat-units (primarily on air temperature). After DAYFF, heat units were
accumulated from estimated fruit temperature. The decrease in development rate
at above-optimum temperatures during the day was also simulated. The model
was calibrated and tested using data sets from several important producing countries
with different environments (south eastern Queensland, Australia, 26 degrees S;
Cote d’Ivoire, 5 degrees N; Hawaii, 20 degrees N; and Thailand, 13.5 degrees N).
The model predicted harvest date with a mean error of 11, 3, 12, and 5 days in
Australia, Cote d’Ivoire, Hawaii, and Thailand, respectively (Malezieux, 1994).
Olive : The full bloom dates were analysed in the world collection in Cordoba,
Spain for 10 years between 1973 and 1989 using 3 methods to develop a model
predicting flowering time. The most accurate model was obtained using the method
of heat units accumulated before flowering. The heat accumulation periods were
determined from phenological and temperature data and 12.5°C was found the
most appropriate threshold temperature for heat accumulation (Alcala and Barranco,
1992).

THERMOPERIODISM IN FRUIT CROPS

Thermoperiodism is a phenomenon in which growth and development is promoted
by altering the day and night temperatures. Plant productivity is higher under
thermoperiodic environment. Many thermoperiodic responses interact with the
light environment, typically via photoperiodism and probably balances in the
phytocrome system. Growth rates of vegetative parts of a plant are always strongly
influenced by temperatures, sometimes plant form is changed in rather specific
ways, and often the responses are delayed. This is often most noticeable in the
vegetative structures associated with flowering. For proper growth and development
of the entire plant, the temperature range within the day should include near
optimal temperatures for the growth of all necessary tissues. Plants may have
differing cardinal temperatures for roots and shoots because of different soil and
air temperatures.
Plant growth and flowering are optimal when the night temperature is lower
than the day temperature for the majority of flowering and fruiting plants produced
hydroponically. Most plant species exhibit these diurnal rhythms where certain
plant process such as the rate of growth of the flower buds, stomata opening, and
discharge of perfume from flowers, cell division and metabolic activity occur
62 Fruit Tree Physiology

more rapidly at a certain time within a 24 hour period. For example, just before
noon photosynthesis in most plants is known to reach a maximum, while just
before dawn cell division also seems to reach a maximum. Many species flower
are grow well only when temperatures during the part of the diurnal cycle that
normally comes at night are lower than temperatures during the day. Interuption
of dark period inhibits some plant processes in long night or short day plants.
Some plants are more sensitive to alternation of day and night temperatures
and produce more flowers when night temperatures are lower than day temperatures
- this effect in plants is called thermoperiodism, and is common amongst many
plant species e.g tomatoes and pepper. Pepper plants also require lower night than
day temperatures for proper production, and many more buds on pepper plants
develop into open flowers when night temperatures are at least 60 C (11oF) lower
than day temperatures. Flowering and fruiting can be adversely affected where
day and night temperatures remain at similar levels on a long term basis, particularly
where temperatures are warm. Production of crops such as tomatoes and peppers
under tropical conditions are poor due to abscission of bud, flower and fruitlet
when they do not receive lower night temperatures. For most plants around 180C
(650F) to 240C (750F) lower than day temperatures, are optimal for photosynthesis.
At night transport of sugars into sinks is promoted. Sinks in most plants are the
developing flower buds, flowers and fruit which have the greatest affinity for the
sugars produced by the plant. The source is the producer of the assimilates, usually
the leaves, but sometimes also the stem in some plant species acts as source. So
sinks get more assimilate pumped into them at night than if they remained as
warm as they were during the day light hours. Lower night temperature setting has
other beneficial effects on plant processes such as root pressure is greater at night
under cooler conditions which increases the pressure in the xylem vessels, so that
calcium and other plant growth compounds which are carried in the xylem stream
are forced out to the leaf tips and into developing buds, flowers and fruits. This
turgor pressure is often essential in the prevention of tip burn as it ensures calcium
is carried to the very edges of the leaves. It is this root or xylem pressure which
also acts to pump up the plant during the cooler night temperatures particularly
after a day when transpiration rates and warm temperatures have resulted in some
wilting and loss of turgour. Respiration rates also get slowed down by lower/
cooler night temperatures and hence the photosynthates are not exhausted and can
be used for growth and development by the plant.

EFFECT OF THERMOPERIODISM IN FRUIT CROPS

Mango : Different temperature regimes like 15/5, 20/10, 30/20°C for 2 weeks in
mango under glass house revealed that the inflorescence development did not
 Heat Units and Thermoperiodism 63

progress when temperature was 15/5°C. Cooler temperature (20/10°C) delayed

the start of anthesis compared with trees grown at 25/15°C and 30/20°C (Sukhvibul
et al; 1999). Mango cultivars exposed to different day/ night temperatures (15/
10, 20/15, 25/20 and 30/25°C) for 20 weeks showed that vegetative growth
increased with increasing temperatures. All cultivars grew vegetative at 25/20 and
30/25°C. The rise in temperatures increased the average number of growth flushes
in responsive cultivars from 0.48 at 15/10°C to 3.21 at 30/25°C (Whiley et al;
1989). The effect of diurnal maximum/ minimum (20/10 or 25/15 °C) temperatures
on seed and fruit development of Irwin, Kensington and Nam Dok Mai mangoes
(Mangifera indica L.) was studied by Sukhvibul et al. (2005). Exposure to low
temperatures (20/10 °C day/ night) 3 days after hand pollination significantly
increased the percentage of stenospermocarpic fruit (nubbins), in which embryos
were aborted at some stage during early fruit development. There were significant
differences between cultivars in the percentage of nubbins produced out of the
total fruit set following overnight exposure to 10°C with 21 per cent for Nam
Dok Mai, 11 per cent for Kensington and 3 per cent for Irwin. At 45 days after
pollination, nubbin fruits were much smaller in size and weighed 50 per cent less
than normal fruits. The lower percentage of nubbin fruits in Irwin implies a
greater adaptation to cool temperatures by this cultivar during fruit set and early
embryo development.
Strawberry : The effects of photoperiod (12, 13, 14, 15 or 16 h), day
temperature (12, 15, 18, 24 or 27°C) and night temperature (6, 9 or 12°C) and
their interactions on flower and inflorescence emergence were investigated by
exposing 4 week old runner plants of strawberry cvs. Korona and Elsanta during
a period of three weeks. A daily photoperiod of 12 or 13 h resulted in the highest
number of plants with emerged flowers. A photoperiod of 14 h or more strongly
reduced this number, while no flowers emerged at a photoperiod of 16 h. Plants
exposed to photoperiods of 12 or 13 h flowered earlier and had longer flower
trusses. A day temperature of 18°C and/ or a night temperature of 12°C were
optimal for plants to emerge flowers and resulted in the shortest time to flowering.
A night temperature of 6°C strongly reduced the number of plants that emerged
flowers, especially when combined with lower day temperatures. Photoperiod
and temperature had no effect on the number of inflorescences, all flowering
plants produced on average one inflorescence. The number of flowers on the
inflorescence increased with decreasing day temperature and when photoperiod
was raised from 12 to 15 h. In general, Korona was more sensitive to photoperiod
and temperature as Elsanta and had a lower optimal day temperature for flower
emergence. These findings will help to produce high quality plant material or to
define optimal conditions when combining flower induction and fruit production
(Verheul et al; 2007).
64 Fruit Tree Physiology

Apple : High temperature has been found to result in dwarfing of the sensitive
plant types. Increasing the difference between night and maximum day temperature
resulted in short internode dwarf plants with small leaves similar to orchard
grown dwarf trees (Steffens et al; 2006). Starting at full bloom, 4 temperature
treatments were applied to 3 year old Golden Delicious and Cox’s Orange Pippin
trees. Either 17 or 24°C were applied in 3 successive periods of 5–6 weeks each.
In Golden Delicious, exposure to 24°C during the first 5 weeks after full bloom
enhanced shoot growth and reduced flower bud formation in spur buds. The
difference in temperature regime during the third period did not affect either
growth or flowering. Almost all apical shoot buds became floral, irrespective of
treatment. Cox’s Orange Pippin trees maintained at 24°C throughout grew more
vigorously than did those kept at 17°C continuously, but flowering abundancy
was the same. Lowering of the temperature in the last period before harvest did
not influence shoot growth, but markedly reduced flowering of both spur buds
and apical shoot buds. When a night temperature of either 20 or 10°C was applied
in 2 successive periods to 3 year old Cox’s Orange Pippin trees kept at a day
temperature of 20°C throughout. Lowering of the night temperature in the middle
of the season reduced flower bud production, but there was no difference in
growth vigour compared with 20°C continuously. It is postulated that temperature
affects flowering in two opposite ways, whose relative importance determines the
net result (Tromp, 1980).
Litchi : In litchi cultivars, when trees were exposed to day/ night temperatures
of 30/25°C and 25/20°C did not flower and temperatures of 20/15 and 15/ 10°C
gave rise to variable proportions of vegetative, leafy panicles and leafless panicles
depending on the cultivars (Menzel, 2003).
Citrus : Vegetative growth in citrus is enhanced by higher temperatures in
citrus, but after certain limit, it retards the shoot elongation e.g higher temperature
(38/ 28°C; day/ night temperature) for 10 weeks in Sour orange, Troyer citrange
and Valencia oranges showed that seedlings were with short internodes and leaves
were markedly shorter as compared to normal ambient temperature (28/ 22°C).
At cooler temperature (20/ 15°C day/ night) more leafless floral shoots are produced
and at higher soil and air temperature there is enhanced production of leafy floral
shoots. Carotenoid content increased substantially under cool environment (20/
15°C; day/night temperature) while warm temperatures (30/ 15°C) reduced the
caroteniod content but increased the chlorophyll content in Valencia oranges .
The effect of early season day/ night temperatures on vegetative growth,
flowering and fruiting of 2 year old potted cultivar Tosa Buntan pummelo trees
on trifoliate orange root stock was investigated by Susanto et al. (1992). The
growth chamber day/ night (12 h/12 h) temperature regimes were 20/ 5°C,
 Heat Units and Thermoperiodism 65

20/ 10°C, 20/ 15°C, 25/ 10°C, 25/ 15°C and 25/ 20°C, from early December 1988
to early April 1989. The higher day/ night temperatures shortened the time to
shoot initiation and flowering, with day temperature having a stronger effect than
night temperature. The trees from the 25°C day temperature regime produced
fewer new shoots, leaves and flowers, but had increased shoot length, leaf size
and fruit set compared to those grown at 20°C. Fruit from the 25°C day temperature
regime were markedly larger and had a more pyriform shape, thicker peel, higher
total soluble solids and lower juice citric acid contents in early December 1989.
There were no significant differences in chlorophyll, N, P, K, Ca, Mg and
carbohydrate contents in new leaves at the termination of temperature treatments.
Pineapple: Low (23 and 15°C) or high (29 and 20°C) temperature for
3 months prior to harvest increased the translucency. Fruits on plants grown at
26/ 22°C (day/ night) matured in 198-204 days from forcing, while fruits matured
in 230 days in 34/ 18°C; which might be due to above optimum day / night temp
(Bortholomew and Malezieux, 1994). The incidence of the disorder was correlated
with both higher (28/ 13°C, max/ min temperature) and lower temperatures
(23/ 15°C) 3 months preceding the harvest. There was marked increase in shoot
NR activity (NRA) during the first half of the light period in plants grown under
constant temperature (28°C light/ dark) whereas significant elevations in the
NRA were detected only in the root tissues at night under thermoperiodic conditions
(28°C light/ 15°C dark). Under both conditions, increases in NR transcript levels
occurred synchronically about 4 hr prior to the corresponding elevation of the
NRA. Diurnal analysis of endogenous cytokinins indicated that transitory increases
in the levels of zeatin, zeatin riboside and isopentenyladenine riboside coincided
with the accumulation of NR transcripts and preceded the rise of NRA in the
shoot during the day and in the root at night, suggesting these hormones as
mediators of the temperature induced modifications of the NR cycle.
Grapes : Anthocyanin accumulation in berry skin is reduced by high night
temperature (30°C continuous) while low night temperature (30/ 15°C: D/ N
temperature) increase the anthocyanin content (Mori et al; 2005). The impact of
fruit temperature on the phenolic metabolism of grape berries (Vitis vinifera L. cv.
Merlot) grown under field conditions with controlled exposure to sunlight were
conducted by Cohen et al. (2008). Diurnal temperature fluctuation was damped
by daytime cooling and night time heating of clusters. Daytime only and night
time only temperature controls were applied for comparison. The results indicated
that damping the diurnal temperature fluctuation advanced the onset of ripening.
Those berries were larger (double-damped: 0.753 ± 0.015 g berry–1 vs control:
0.512 ± 0.034 g berry–1) and more colored than all others. Damping the diurnal
temperature fluctuation reduced proanthocyanidin mean degree of polymerization
66 Fruit Tree Physiology

(double damped: 21.8 ± 1.0 vs control: 28.0 ± 1.7). Proanthocyanidin accumulation

at veraison was linearly related to heat summation over the developmental period
with night time heating yielding the highest concentration and day time cooling
yielding the lowest (night-heat: 1.46 ± 0.13 mg berry –1 vs day cool:
0.97 ± 0.09 mg berry–1). Damping the diurnal temperature fluctuation had a marked
effect on the rate of fruit development whereas total heat summation had more
of an effect on phenolic metabolism alone. The results provide insight on the
direct effect of temperature on phenolic metabolism.
Annona : The effects of warm (30/ 25°C) and cool (20/ 15°C) day/ night
temperatures on fruit set and fruit growth of potted cherimoya (Annona cherimola
Mill.) trees were investigated under glass house conditions. It was observed that
warm temperature adversely affected pollen germination percentage, reduced fruit
set due to both pollen and stigmatic damage from heat stress. Fruit at warm
temperatures grew slowly and required more days to mature than those at cool
temperatures. Warm temperatures reduced fruit growth rates at the initial and
final growth stages. The intervening growth deceleration period was longer at
warm temperatures. Warm temperatures produced asymmetrical and small fruit
containing a small number of seeds, caused by low viability pollen. Temperature
had no significant effect on the total soluble solid content in fruit (Higuchi et al;
1998).

CONCLUSION
Temperature plays an important role in growth and development of plants. Growing
degree days (GDD) also called growing degree units (GDUs), are a measure of
heat accumulation used by horticulturists, gardeners, and farmers to predict the
date that a flower will bloom or a crop reach maturity. The transition from one
developemental stage to another requires energy which is provided by heat and
the amount of heat required varies from species ro species. Heat units are calculated
based on the difference between the average of the daily maximum and minimum
temperatures and a certain critical basal or developmental threshold temperature.
Heat units are very helpful in selecting the suitable site for fruit crops, selection
of varieties, predicting pest out break etc. Different fruit crops require different
heat accumulation units for proper growth and development. Some plants are
more sensitive to alternation of day and night temperatures and produce more
flowers when night temperatures are lower than day temperatures and this effect
in plants is called thermoperiodism. If there is no variation in diurinal temperature
flowering and fruiting in most of the plants is adversly affected. Lower night
temperatures are essential for better performance of crops because cooler sinks
get more assimilate pumped into them at night than if they remained as warm as
 Heat Units and Thermoperiodism 67

they were during the day light hours and respiration rates are also low. Further
the root pressure is greater at night under cooler conditions which increases the
pressure in the xylem vessels that helps the calcium and other plant growth
compounds carried via xylem stream to be forced out to the leaf tips and into
developing buds, flowers and fruits.

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 4

PHYSIOLOGY OF FLOWERING IN
HORTICULTURAL CROPS

F lowering is a cascade reaction consisting of several steps. Flower formation

is a transition phase in the life cycle of a plant. It is of immense importance
for perpetuation and origin of variability in the next generations. Flower initiation
takes place by the transformation of vegetative apex into a reproductive structure.
It signifies the transition from vegetative to floral state. The shoot meristem is
reduced to develop sepals, petals, stamens, carpals in place of leaves. The plant
must attain specific state of ‘ripeness to response’ before it flowers. Once the
stage is reached, then it can induce to flower.
The physiology of flowering involves the changes in the characters of cells
proliferating in the meristematic tissues of the shoot owing to the specific gene
action and change in the phyto-hormones level.
The flowering is closely linked to the diverse environmental conditions in
which each species has evolved. The effects of the large number of factors that
influence the proportion of buds giving rise to flowers have generally been
interpreted in terms of an inbuilt propensity to flowering and interference with
attainment of this. The controlling factors may involves hormonal balance
(Luckwill, 1974; Hoad, 1984), availability of nutrients, especially carbohydrates
and interaction of these (Ryugo, 1986).

Vgetative ⎯→ Transition ⎯→ Reproductive ⎯→ Plant maturation

(Juvenile) Becomes reproductively (Adult)
competent

FLOWERING PHYSIOLOGY
Plants come to flower after a certain period of vegetative growth. The process
until plants come to flower is classified into three stages, that is:
 Physiology of Flowering in Horticultural Crops 71

1. Differentiation of flower buds.

2. Development of flower buds.
3. Opening of flower.
The differentiation of flower buds is the most important turning point in this
process, and the plant’s life cycle switches from vegetative growth to reproductive
growth. The switching from vegetative growth to reproductive growth is called
flowering. Flowering physiology is the study aimed to clarify the mechanism
switching the mode of growth from vegetative to reproductive.
Flowering occurs when flowering genes are activated. The activation of the
flowering genes occurs autonomously or is triggered by environmental factors.
Autonomous flowering occurs when plants reached a certain age. The
environmental factors triggering the activation of flowering genes are photoperiod,
low temperature and various stresses. Photoperiodic flowering regulated by
photoperiod and vernalization regulated by low temperature have been well studied.

Physiology of Flowering in Horticultural Crops

It can be studied under the following heads:
1. Events in the bud leading to flowering : Three events were proposed to
occur during the transition of buds to flowering i.e. induction, evocation and
initiation (Metzger, 1987).
Induction : The step of flowering at which the flowering stimulus is generated
is called floral induction. Searle (1965) defined induction and induced state
as a physiological condition initiated in tissues by external influences such
as water stress, chilling temperature or photoperiod, progressively towards
floral expression even under subsequent non-initiating conditions. This event
is translated to production of a putative floral stimulus or alternative in the
ratio of florigenic and antiflorigenic components which may be translocated
to target cells in meristems.
Evocations : The step at which the shoot apical meristem has received the
flowering stimulus, and is irreversibly committed to form flower bud primordia
is called floral evocation, so these are those processes occurring in the apex
essential for the formations of floral primodia (Metzger, 1987). Immunological
on electrophoretic analysis of evoked apices of photoperiodic plants
demonstrated alternation of protein complement after induction and prior to
initiation of flower primordia (Lyndon et al; 1983). Floral evocation involves
two development states: competent and determined. In general terms, a cell
or group of cells is said to be competent if it can respond in the expected
72 Fruit Tree Physiology

manner when given the appropriate developmental signal. For example, if a

vegetatives shoot (scion) is grafted onto a flowering stock and the scion
flowers immediately, it is obviously capable of responding to the level of
floral stimulus present in the stock and is therefore competent. Failure of the
scion to flower would indicate that the shoot apical meristem has not yet
attained competence. This is another way of stating that the apical meristem
is in the juvenile phase.
Flowering involves several processes that occur in different parts of the
plant. In photoperiodic plants, the inductive photoperiod causes the leaves to
export a floral stimulus to the rest of the plant that alters the developmental
fate of the cells of the shoot apex. Once the meristem becomes committed
to the new development program (flowering in this case), it is said to be
florally determined. The meristem is determined if it follows the same
developmental program even after being removed from its normal physical
and environmental context.

Initiation : In this process the evoked bud becomes recognizable as a flower

bud and is thus committed to reproductive development. This evident the
broadening and flattening of the growing point with concurrently developing
lobes.
2. Differentiation of the growing points : The transition from vegetative to
reproductive development is marked by an increase in the frequency of cell
division within the central zone of the shoot apical meristems. This leads to
increase in the size of meristem and the shoot meristem is induced to develop
sepals, petal, stamens and carpels in place of leaves. The first detectable
change in the growing points of the bud after induction of flowering is
 Physiology of Flowering in Horticultural Crops 73

increasing synthesis of DNA and RNA (Buban and Faust, 1982; Faust 1989).
The first visible sign of differentiation is when the flat apical meristem
becomes domed, then the central meristem is partitioned and the pitch
meristem develops. The meristem becomes a block-like structure and its
subsequent development is relatively rapid. The stage of flower bud
development at any given time during summer and autumn varies with type
of bud (basal, middle, upper or terminal shoot suds) and with species and
cultivars (Crabbe, 1984)
In temperate fruits, normally 8-14 days are required from the beginning
of histological differentiation to the appearance of lower meristem. The
subsequent development of the flower meristem is relatively rapid. The entire
process of flowers and differentiation may take 54-112 days depending on
the species.

Changes Occuring in Target Tissues

1. Biochemical changes :
(i) Development of chloroplast inhibitors
(ii) Moving out of floral stimulus from leaves
(iii) Increase in total RNA synthesis and chromatins dependent RNA synthesis
(iv) DNA synthesis increases
(v) Mitotic activities increases
(vi) Number of mitochondria increases
(vii) Starch content decreases
2. Metabolic changes :
(i) Meristem height increases
(ii) Cell size increases
(iii) Lengthening of mitotic phase (Second mitotic peak)
(iv) Entry of cells from photosynthetic to mitotic phase
(v) Flower bud initiation
De-novo synthesis of DNA and rise in mitotic activity leads to production
of cells which are responsible for formation of flower bud primordia.
Morphogenesis occurs when the flower bud primordia develop also called
morphogenetic phase.
74 Fruit Tree Physiology

MECHANISMS OF FLOWERING
Various theories have been proposed to explain the mystery of flowering:
1. Phytocrome theory
2. Bunnings hypothesis
3. Chailakiyans hypothesis
1. Phytochrome theory : Phytochrome is a non-photosynthetic photo-receptor
pigment that exists in two forms, Pr form and Pfr form. The former absorbs
light of wave length P660 and the latter of P730. The two forms are inter
convertible.
It is believed that phytochrome controls flowering in plants. P730
suppresses flowering in short day plants and promotes flowering in long day
plants. On the other hand, P660 promotes flowering in short day plants and
inhibits in long day plants. Sun light functions as red light. At the end of
light i.e. in evening after sun set, Pfr form pre-dominates while at the end
of dark period, i.e. before sunrise, Pr form predominates.

During long night Pfr form is converted to Pr form. If the long dark
period is interrupted with red light, sufficient amount of Pr form is not
formed hence flowering is inhibited in short day plants. Flowering in short
day plants depends upon Pr/Pfr ratio. Low ratio of Pr/ Pfr inhibits flowering
in short day plants and promotes flowering in long day plants. High Pr/ Pfr
ratio favours flower formation in short day plants and inhibit in long day
plants (Takimoto and Saji, 1984).
2. Bunning’s hypothesis : Bunning (1958) assumes the presence of endogenous
rhythms (oscillator) which consist of two half cycles. The first half cycle
occurs in day and is called photophilous phase. During this, anabolic process
predominates including flowering in plants. The other half cycle is dark,
sensitive and is called as skotophilous phase. In this, catabolic process
(dehydration of starch) predominates.
Short day plants have a critical day length of 9 hours. This period falls
within the photophilous phase. Light during skotophil phase will inhibit
photoprocess initiated during photophase. The long day plants have a critical
day length of 15 hours and some light falls in the skotophilous phase. Under
 Physiology of Flowering in Horticultural Crops 75

these conditions long day plants will flower. In short day plants oscillator is
present close to skotophilous phase, while in long day plants, it is close to
photophilous phase.
3. Chailakhyan’s hypothesis : This hypothesis assumes that flowering hormone
– florigen is a complex of two types of substances – gibberellin and anthesins.
Gibberellin is essential for growth of the plant stems and anthesins are required
for flower formation.
According to Chalikiyan (1937), flowering in all annual seed plants
requires two phases: (i) Floral stem formation phase (ii) Flower formation
phase. First phase involves increased carbohydrate metabolism and respiration
with increased content of GA in leaves. Second phase requires intensive
nitrogen metabolism, higher content of anthesins in leaves and nucleic acid
metabolites in stem buds. Long day conditions favour the first phase while
short day conditions favour second phase. In long day plants gibberellins are
critical, while anthesins are critical in short day plants. However, anthesin is
hypothetical; it has not been isolated as yet.
Chailakhyan (1937) proposed that vernalin is produced in plants at low
temperature. In long day conditions it is converted into gibbrellin. Anthesin
is present in long day plants. Anthesin along with vernalin cause flowering
in long day plants. But in short day conditions, the vernalin is converted to
gibbrellin, hence flowering does not occur. Addition of gibbrellin to long day
unvernalised plants in long day conditions leads to flower formation as these
plants contain anthesin. Gibbrellin is ineffective in producing flowers in
short day plants as they lack anthesin.

As postulated by Purvis (1961) the formation of a substance A from its

precursor. A is then converted to B after chilling. The substance B is unstable.
At suitable temperature, B is converted to a stable compound D called Vernalin.
At high temperature, B is converted to C i.e devernalisation occurs. At correct
photoperiod, vernalin is converted to substance F (florigen) - a flowering
hormone. Florigen induces flower formation in proper photoperiod.
76 Fruit Tree Physiology

Photoperiodism
Relative length of daily light and dark periods helps in transforming a vegetative
shoot meristem to a floral bud. The induction of flowering in response to the
relative length of daily light and dark period is called photoperiodism. It simply
means the duration of light. It is a physiological response of plants in relation to
duration of light. There are various photoperiodic responses in plants, however,
the most important is the initiation of flowering. The photoperiodic signal is first
received by phytochrome in the leaf, and the signal from phytochrome starts the
biological clock. After the biological clock measures a certain period of time
(inductive photoperiod), the production of the flowering stimulus in the leaf
starts. The flowering stimulus is transmitted from the leaf to shoot apex leading
to change in the growth mode of the meristem from vegetative to reproductive.
The shoot apical meristem produces primordia of floral organs viz. sepals, petals,
stamens and carpels, thus generating a flower bud.
Based on photoperiodic responses plants may be classified into three groups.
1. Short day plants (SDP) : These plants flower only when exposed to day
lengths shorter than a certain critical maximum. Critical day length is 11-15
hours. Most of the plants in tropics are short day plants e.g pineapple,
strawberry etc.
2. Day neutral plants (DNP) : These plants flower after a period of vegetative
growth regardless of the photoperiod. Flowering is not affected by day length
and it occurs any time during the year e.g guava and citrus.
3. Long day plants (LDP) : These plants begin flowering when exposed to day
lengths longer than certain critical minimum. Below the critical period, these
plants continue their vegetative growth. Most of the temperate zone fruits are
long day plants. Critical day length lies between 12-14 hours.
 Physiology of Flowering in Horticultural Crops 77

In the short day plants, when the long night period is interrupted by a brief
exposure to light, the plants fail to flower. Similarly, long day plants respond to
nights shorter than the critical dark period.
Short-Day Long-Day
8-Hour Light 16-Hour Light Plants Plants
A Flower Vegetative

B White
VEG. Flower
Light
C Flower
VEG.
Red Light
D Flower

E Flower VEG. Far Red

Light
VEG. Flower
F

Fig. : The effect of Red and Far-Red Light during the night break on the flowering
of LD and SD plants. The light is obtained either from sunlight or high-
intensity radiations.

Table: Response of SDP and LDP to photoperiods.

Day/night length Duration (hrs) SDP LDP

Day Night

Long day Short night 16 8 No flowering Flowering

Long day Long night 16 16 Flowering No flowering
Short day Short night 8 8 No flowering Flowering
Short day Long night 8 16 Flowering No flowering
Continuous day 24 – No flowering Flowering

Table: Difference between SDP and LDPs.

Short day plants Long day plants

Can flower under short day conditions Flower under long day conditions
Inhibition of flowering by interruption of No inhibition of flowering by interruption of
long dark period by flash of light long dark period by flash of light
Interruption during light period does not Interruption during light period inhibit
inhibit flowering flowering
Under continuous darkness flowering can Such conditions can not induce flowering
occur if food is supplied
78 Fruit Tree Physiology

Photoperiodic induction : It is defined as the initiation of flowering in a

plant after exposure to light for a required period of time. This is calculated for
a 24 hrs and the time of specific exposure to light within a day is called
photoinductive cycle. The number of these cycles may be one, two or many
depending on a particular plant species and together they are called photoinduction.
Site of photoinduction perception : Experiments conducted by various
scientists like Chailkhyan (1937), etc. have demonstrated that leaves are the sites
of photoinduction e.g no flowering is produced in defoliated plants even after
exposure to correct photoperiod.
Transmission of stimulus : The stimulus is transmitted from leaves to the
target region (in the axils or at the branch apices or flowering regions) through
phloem at 10-24 mm/ hr. Photosynthates, however, travel at 1000 mm / hr. For
LDP the rate of transmission is 30 cm/ hr and SDP is 2.4mm/ hr, however, it may
vary from plant to plant.
Nature of stimulus : Chailkhyan (1937) suggested that the stimulus is a
hormone and named it florigen based on following evidences:
● Transmission through phloem.
● Transmission of stimulus from a photoinduced branch to a non photoinduced
branch through grafting.
● Initiation of flowering in a non photoinduced plant by the leaf extracts of a
photoinduced plant e.g xanthium.
However, others like Sachs and Hackett (1983) opposed the florigen concept
and believed that diversion of nutrients through photoinduction towards meristems
leads to the floral initiation.

Physiology of flowering in xanthium (Lang, 1955)

 Physiology of Flowering in Horticultural Crops 79

Phytochrome and Flowering

Phytochrome :
● Phytochrome is a homodimer: two identical protein molecules each conjugated
to a light-absorbing molecule (compare rhodopsin).
● Plants make 5 phytochromes: PhyA, Phy B, as well as C, D, and E.
● There is some redundancy in function of the different phytochromes but
there also seem to be functions that are unique to one or another.
● Phytochromes exist in two inter convertible forms.
PR because it absorbs red (R; 660 nm) light
PFR because it absorbs far red (FR; 730 nm) light
● These are the relationships:
Absorption of red light by PR converts it into PFR.
Absorption of far red light by PFR converts it into PR.
In the dark, PFR spontaneously converts back to PR.

The behavior of phytochrome explains the experimental results with the

cocklebur:
● Sunlight is richer in red (660 nm) than far red (730 nm) light so at sundown,
all the phytochrome is PFR.
● During the night, the PFR converts back to PR.
● The PR form is needed for the release of the flowering signal.
● Therefore, the cocklebur needs 8.5 hours of darkness in which to convert all
the PFR present at sundown into PR, carry out the supplementary reactions
leading to the release of the flowering signal (florigen).
● If this process is interrupted by a flash of 660 nm light, the PR is immediately
reconverted to PFR and the night’s work is undone (C).
● A subsequent exposure to far red (730 nm) light converts the pigment back
to PR and the steps leading to the release of florigen can be completed (D).
80 Fruit Tree Physiology

● Exposure to intense far red light at the beginning of the night sets the clock
ahead about 2 hours or so by eliminating the need for the spontaneous
conversion of PFR to PR (E).
Working of phytochrome :
● When sunlight (660 nm) converts PR into PFR, the PFR moves from the
cytoplasm into the nucleus.
● There it binds to a protein called PIF3 (phytochrome-interacting factor 3).
● PIF3 is a helix loop helix protein as are many transcription factors.
● The complex of the two binds to and turns on promoters containing the
sequence
CACGTG
GTGCAC
● These promoters are found in genes that those selves encode other transcription
factors.
● These other transcription factors, in turn, initiate transcription of a variety of
genes that are expressed when the plant is exposed to light.
● Exposure to far red light converts the PFR back to PR which
– dissociates from PIF3, and
– returns to the cytoplasm.
The studies of the role of phytochrome in etiolation indicate that PFR is the
active form; PR inactive. However, flowering of long night (short day) plants like
the cocklebur requires PR.

The scheme shows that the last exposure is important for initiation or inhibition
of flowering. This indicates that some pigment system acts on the plants as
photoreceptors. Furthermore, photoinduction and deinduction in different wave
lengths also indicates involvement of two separate pigments i.e one for induction
and another for deinduction or two forms of the same pigment that undergo
conformational changes due to reception of different light qualities and influence
induction or deinduction accordingly. Butler et al. (1959) discovered phytochrome
which is widely distributed in green plants and is present within plasma membrane
of cells.
 Physiology of Flowering in Horticultural Crops 81

Light and phytochrome : Phytochrome a biliprotein pigment probably

localized in plant membranes. It changes in form and absorbance in response to
the light it receives. One form Pr (Phytochrome red) absorbs mainly red light
(660 nm) and is transformed to another form Pfr (Phytochrome far-red) which
absorbs mainly in the far-red region (730 nm). The precise role of phytochrome
in flower initiation is not clear. However, it may be added that this pigment is not
the flowering stimulus but possibly aids the stimulation of flowering stimulus.
In SDP, at the end of light period Pfr is in high concentration and Pfr and Pr
ratio is such that it prevents formation of flowering stimulus. If the dark period
is prolonged, then either Pfr is destroyed or it is reverted to Pr state. At this stage
the ratio of Pfr to Pr is such that it triggers processes leading to formation of
flowering stimulus. Under the situation where dark period is briefly interrupted
by red light, there is formation of Pfr from Pr and the ratio of the two is altered
and thereby preventing the formation of flowering stimulus.
On the other, LDP need high ratio of Pfr to Pr to trigger flowering stimulus.
This ratio is attained at the end of a long day. Once the night is long then Pfr is
revered back to Pr and the formation of flowering stimulus is prevented. Under
the conditions where night in interrupted by red light, then Pr is changed to Pfr.
Here the ratio of two pigments is high and flowering stimulus is produced.
Hendricks and Borthwick (1955) discovered that phytochrome is the
photoreceptive pigment that induces flowering. Four steps are involved in flowering
response:
1. Perception of the stimulus by phytochrome in the leaves.
2. Change to new pattern of metabolism in the leaves leading to the production
of flowering hormone, the florigen.
3. Translocation of florigen to the bud primordial.
4. Transformation of vegetative bud primordial into floral bud.

PHOTOPERIODISM AND FLOWERING IN FRUIT CROPS

In mango, flower induction is caused by cool temperature and not by short
photoperiods and that flowering is inhibited by warm temperature not by long
photoperiods. Variety like Neelum flowers twice in Kanyakumari but only once
under north Indian conditions and the common observation that the eastern side
of mango tree, which receives longer hours of sunlight flowers a few days earlier
than others, indicate the effect of light on reproductive process of mango.
In strawberry, flower formation is influenced by the duration of length of
daily light period. A decreased light period induces flower formation at the expense
82 Fruit Tree Physiology

of runners, while the longer day length tends to induce runner rather than flower
formation. Although short days favour flower formation, regardless of temperature,
yet temperature to a certain extent modifies the day length response. In general,
any decrease in temperature shortens the day length, which will permit the flower
formation. Konsin et al. (2001) found that 12 and 13.5h photoperiods were equally
efficient in produced yield in Korona strawberry, but more vigorous vegetative
growth was maintained during treatment in a 13.5 h photoperiod. Serce and
Hancock (2005) studied 8 strawberry cultivars and reported that cvs. CFRA 368
and Frederick 9 produced same number of flowers both under short day and long
day conditions. Day neutral genotypes like LHSO 4 and RH 30 produced runners
under short day conditions whereas ‘Fort Lanamie’ strawberry has floral heat
tolerance.
Teaotia and Singh (1967) studied the effect of seasonal variation on sex
expression of papaya cv. Coorg Honey Dew and reported that the production of
female flower was promoted by long day and high temperature. Similarly, Storey
(1986) observed that long day treatment, i.e. photoperiod cycle of 16 hour light
and 8-hour darkness favored femaleness in Co. 1 papaya.
Four systems of production viz artificial light/ irrigation/ shade; artificial
light/ irrigation; artificial light/ shade; artificial light and a natural condition
system were compared by Cavichioli et al. (2006). The results showed that artificial
light with and without irrigation increased the number of flowers, the number of
fruits and total yield of yellow passion fruit. The irrigation did not affect the
flowering, the fructification and the yield in the treatment with artificial light, but
reduced the number of flowers in the shaded treatment. The shading with and
without irrigation reduced the number of flowers.
The artificial light treatment increased the percentage of fructification. Chiou
et al. (1999) exposed Z. mauritiana plants to light for 15, 30 and 45 days.
Flowering of light treated plants was significantly advanced. Similarly, number
of flowers ad fruit set increased with light treatment.
In pineapple neither a diurnal temperature differential nor short days are
necessary for natural flowering. Rather, attainment of some minimum size and
cool night temperature, which regulates the natural flowering to occur. Pineapple
plant generally flowers after attainment of certain vegetables growth 11-12 months
after planting and formation of at least 40 leaves. A pineapple produces only one
fruit during life time. There is no quantitative involvement of photoperiod in
citrus flowering (Lenz, 1964). Similarly guava is a day neutral plant.
In fruit crops like apples, peaches, cherries and plums, floral initiation appears
to be unaffected by photoperiod, which nevertheless affects the vegetative growth
 Physiology of Flowering in Horticultural Crops 83

of these species. Tromp (1984), using controlled environment with artificial light
in apple found that reduced light intensity for 7 weeks flowering bloom halved
flowering in the following year, while reducing light levels from week 7 to 16
had only a smaller and non-significant effect.

FLORIGEN CONCEPT
Leaf is the site of photoperiodic perception and shoot apex is the site of floral bud
formation. This means that some information must be transmitted from the leaf
to shoot apex which is called as flowering stimulus. The transmission of information
in plants is generally due to movement of plant hormone like substances.
Chailakhyan proposed the florigen (flowering hormone) theory in 1937 that is
generated in leaves exposed to a suitable photoperiodic cycle, transmitted through
phloem, and activates the flowering genes at the shoot apex.

Activation of genes for lower

differentiation

P-660 P-730
P-660 P-730

Sythesis of
Genes for synthesis of florigen
florigen are inhibited

Fig. : Hypothesis regarding photoperiodic induction of short day plant. Synthesis

of florigen in dark and its translocation to the apex for the activation of gene
for lower differentiation. (Chailakhyan, 1937)

The biochemical nature of florigen is still unknown. Currently, the existence of

florigen can be presumed only by grafting experiments. One of the most important
goals in the study on flowering has been to isolate and characterize florigen.
Florigen may not be universal : The florigen concept is now understood in
several ways resulting in confusion. It has been misunderstood that florigen was
proposed as a single component that is common in the plant kingdom. The grafting
84 Fruit Tree Physiology

experiments showed that the flowering stimulus could be transmitted between

different plant species suggesting that florigen was common among species.
Although, gibberellins can induce flowering of some long day plants under
noninductive conditions but gibberellins are not considered as florigen, because
they do not induce flowering of short day plants. However, there is no reason to
consider that florigen needs to be ubiquitous. The florigen of long day plants could
be a gibberellin and the florigen of short day plants might be a chemical other than
a gibberellin. The results of grafting experiments mentioned above should not be
generalized, because grafting is possible only between taxonomically close relatives.
A possible role of GA- like hormones in the induction of flowering was
postulated by Brain (1959). According to him, CO2 is converted into an inactive
or less active processor under the influence of red light or infrared light produces
a GA like hormone which may produce florigen. The GA-like hormone can be
converted into its precursor with exposure to far red light. The same can be
achieved by dark treatment but the process is very slow. This hypothesis of Brain
is based on the effect of darkness or red and far red light and the GA treated long
day plants maintained in short day conditions.

Fig. : GA the flowering response.

Flowering has been induced by growth regulating chemicals in number of
fruits. It is generally believed that high level of gibberellin inhibits flowering
while high level of auxins or auxin like substances may promote flowering either
by reducing the effectiveness of GA or by decreasing the permeability of cell
membrane particularly the plasmalemma (Baxter, 1970).
Alternative ideas on florigen concept : Although, almost 70 years earlier
since its existence was proposed, florigen has not been isolated yet. Some plant
biologists say that florigen has not been isolated, because it does not exist, and
do not assume the existence of florigen to explain the regulating mechanism of
the flowering process. The alternative ideas include the presence of multiple
factors, a flowering inhibitor, and electrical signals.
Multiple factors : The florigen concept assumes one hormonal compound
acting specifically to induce flowering. However, several compounds which are
not specific to regulation of flowering can act as hormonal regulators. In some
 Physiology of Flowering in Horticultural Crops 85

plant species, endogenous levels of cytokinins, putrescine, sucrose and calcium

ion increase and they move to the shoot apex under a flower-inducing condition.
These facts suggest that flowering occurs when these compounds work together
in the shoot apex. Therefore, all of these compounds could be factors necessary
for the induction of flowering. This is called the multifactorial concept.
The florigen concept does not necessarily assume a single universal compound.
It does not necessarily assume specific compounds. However, it could be a mixture
of two or more compounds, and they could be known compounds with multiple
functions as usual plant hormones are. In this sense, the multifactorial concept
may be considered as a variation of the florigen concept.

Importance of Photoperiodism
1. Yield can be increased by knowing favourable photoperiods in plants.
2. Plants like radish, carrot etc. can be made to remain vegetative for longer periods.
3. Annuals can be grown twice or thrice a year.
4. Prevention of winter dormancy and autumn leaf fall.
5. Increased stolon formation through long days in strawberry.

VERNALIZATION AND FLOWERING

There are environmental factors regulating flowering other than night length.
Temperature is also important. Some plants require exposure to a low temperature,
0 to 10°C, for a few days to a few weeks for flowering. Such an induction of
flowering by a low temperature is called vernelization. It is observed in many
long day plants, but is not correlated with photoperiodism. The length of chilling
may vary from species to species ranging from four days to few months and the
most effective temperature is found to be 4°C.
Site of vernalization : Active apical meristem and young leaves.
Stimulus of vernalization : In annuals, embryo axis is the site of stimulus
reception while in biennials and perennials it is located in apical meristem of the
shoot. In biennials, vegetative growth takes place during the first year and flower
in the following summer after vernalization during winter. However, perennials
need vernalization cycle every winter. Meristems are the sites of stimulus and
transmission. It has been postulated that vernalin is transmitted from vernalised
plant to non vernalised through grafting.
Mechanism of vernalization : Two theories have been postulated to explain
the mechanism of vernalization
1. Phasic development hypothesis
2. Hormonal hypothesis
86 Fruit Tree Physiology

1. Phasic development hypothesis : The development of annual seed plants

consists of several physiological stages / phases which occur in pre determined
sequence i.e when one phase terminates the next begins. The main stages
have been identified as:
(i) Thermostage : It is a vegetative phase and low temperature (0.14OC),
moisture and aeration are necessary for the completion of this stage
wherein the length depends on plant and environment.
(ii) Photostage : It requires high temperature and is also called
photoperiodism. Here vernalin helps in the synthesis of florigen.
(iii) Third stage : It is necessary for the formation of sex organs and gametes.
2. Hormonal hypothesis : Same as Chalikhyan hypothesis.
Significance of vernalization :
1. Production of more than one crop in a year by shortening the vegetative period.
2. Protection from freezing injury by sowing winter crops in spring.
3. Tropical plants (requiring long summer) can be grown in temperate areas
(short summer).
Molecular and genetic aspect of vernalization : Seeds contain specific
proteins and its associated components which bind to loci that are involved in
flower induction and silence the chromatin by heterochromatization.
Heterochromatization is achieved by histone methylation and histone deacetylation
at specific loci. One such protein is known as flowering locus C (FLC). FLC acts
as a repressor. Cold treatment in fact induces few FLC antogonizing genes and
FLC dissociates from the loci and get degraded or remain free. There are several
genes that are involved VRN1, VRN2, VRN3 and VIN3. Even Frigida (FRI)
proteins are involved. Frigida promotes FLC and FRI antagonizes FLC. Once the
chromatin is free from FLA and its associated protein, they interact with floral
integrators such as SOC, CO, FT and LFY, which inturn activate genes for floral
parts. In certain cases GA can overcome vernalization (e.g Petkus rye seeds).

Izawa et al. (2003)

 Physiology of Flowering in Horticultural Crops 87

TRANSLOCATION OF FLOWERING STIMULUS

Flowering stimulus from leaf to apical apices is translocated through phloem
cells. Effect of photoperiodic stimulus induction may be localized or generalized
depending upon nature of plants. Transmission of stimulus also occurs from one
plant to another e.g through grafting.
88 Fruit Tree Physiology

Factors for Induction of Flowering in Fruit Crops

Major factors regulating induction of flowering could be broadly classified into
five major groups:
Environmental factors : Photoperiod, temperature, water relations and
location etc.
Internal factors : Nitrogen metabolism and carbohydrate status, nutritional
factors and phytohormones.
Soil factors : Nutrient status, soil structure, texture, moisture content etc.
Horticultural traits : Age, crop load, root stocks, type of shoots etc.
Cultural practices and use of chemicals : Smudging, girdling, root and
shoot pruning, use of TIBA, SADH, PBZ, CCC etc.
The important factors which effect flowering in fruit crops are discussed
hereunder:
1. Temperature : Various dominant effects of temperature are - effects on
emergence from dormancy, i.e. the meeting of winter chilling and thermal
time requirements to achieving bud break. Hield et al. (1966) reported that
in grapefruit temperature of 20 to 26°C (day) and 8 to 12°C (night) permitted
expression of up to 100 per cent of terminal flowers whereas day temperature
of 32 to 39°C prevent flowering. In Washington Naval oranges high
temperature (30°C day/ 25°C night) inhibited flower formation, whereas low
temperatures (15°C/ 10°C) favored the flower formation because high day
temperature of 30°C impaired the flower formation leading to irregular
flowering (Lenz, 1964). Exposure of Satsuma mandarin trees to a low
temperature in the fall induced flowering and the buds after floral evocation
develop floral buds within a few days at 32°C/ 80 RH (Okuda et al; 2004).
Temperature greatly influences the development of floral buds in mango
(Singh et al; 1974). Initiation of growth in shoots at cool temperatures (18°C
day/ 10°C night or 15°C day/ 10°C night) leads to development of
inflorescence in mango. Cool night temperatures between 8 and 15°C in
combination with day temperatures below 20°C typically induce flowering
if shoot initiation occurs when plants are exposed to those conditions. During
panicle development, Singh et al. (1966) observed the maximum and minimum
temperatures have pronounced effect on the percentage of perfect flowers.
Sukhvibul et al. (1999) reported that temperature had an inverse effect on the
total number of flowers per inflorescence with 619.6±108 flowers per
inflorescence at 20/ 10°C, decreasing to 431.3± 80.5 flowers at 30/ 20°C in
four mango cultivars. Floral expression may depend on the ecotype
 Physiology of Flowering in Horticultural Crops 89

(polyembrayonic; Nam Dok Mai, Kensington and monoembryonic; Irwin,

Sensation) and low temperature of 15/ 5°C strongly inhibit inflorescence
development. The percentage of hermaphrodite flowers decreased in
polyembryonic cvs. but, increased in monoembryonic cvs. at 20/ 10°C.

Table : Effect of temperature on no. of days from first-untill 50 per cent anthesis
of male and hermaphrodite flowering in mango (Sukhvibul et al; 1999).

Cultivar Days from Ist anthesis to Days from Ist anthesis to mid-
mid-anthesis for male flowers anthesis for hermaphrodite
temperature (°c) 20 weeks flowers temperature (°c)
20/10 25/15 30/20 20/10 25/15 30/20
Nam Dok Mai 16.8 14 5.5 5.5 11.0 4.3
Kensington 24.3 13.5 7.0 25.8 10 3.5
Irwin 20.8 20.5 13.0 5.8 23 10.3
Sensation 20.8 11.5 6.5 22.3 9.8 4.3
Mean 20.6 13.0 6.2 19.2 10.3 4.1

The optimum temperature was found to be high (21°C) under short day
conditions and low (15°C) under long day conditions for floral initiation in
strawberry. Manakasem and Goodwin (2001) studied the responses of day
neutral and June bearing (short day) strawberries to temperature and day
length. Floral initiation was repressed in long days, with poor fruit set and
development compared with flowers initiated under short day conditions.
The day neutral cultivars were less affected by day length, but each had
specific temperature and day length combinations during floral bud initiation
for optimum fruit development. The optimum temperature during floral
initiation for the reproductive development of all cultivars was between
18/ 13°C and 21/ 16°C, with sharp decrease in flower number, fruit set and
fruit weight at lower or higher temperatures was observed.
In litchi reversion of some reproductive buds to vegetative ones occured
due to high day/ night temperature (20/ 15, 25/ 20, 30/ 25 vs 15/ 10°C) and
was attributed to reduction in the proportion of female flowers (Menzel and
Simpson, 1991). High temperature and water stress after panicle emergence
have strong effects on reproductive development and sex ratio in litchi
(Tables a and b). Menzel and Simpson (1988) reported that moderate day/
night temperature (20°/ 15° & 15/ 10°C) increased vegetative growth and
reduces flowering in seven litchi cultivars and the temperature of more than
25/ 20°C totally eliminate flowering.
90 Fruit Tree Physiology

Table (a) : Low temperature effect on flowering in litchi (Menzel and Simpson, 1991).

Cultivar Temperature 15/ 10 (°C) Temperature 20/ 15 (°C)

VEG LP LLP VEG LP LLP

Tai So 0 93 7.0 50 49.8 0

Bengal 0 70.7 29.4 49.2 32.0 8.6
Souey Tung 0 39.4 60.7 58.3 3.2 28.3
Kwai May Pink 0 54.5 45.6 20.0 48.7 8.2
Kwai May Red 0 7.0 93.0 80.0 3.0 5.3
Salathial 0 0 100 16.5 19.5 58.1
Wai Chee 0 0 100 4.0 13.5 80.5

Table (b) : Low temperature effect on flowering per branch in litchi. (Manzel and
Simpson, 1991).

Cultivar Total number of flowers % of Female flowers

15/ 10 20/ 15 25/ 20 30/ 25 15/ 10 20/ 15 25/ 20 30/ 25

Tai So 45 108 117 207 74 51 31 11

Bengal 96 127 NF NF 98 68 NF NF
Souey Tung 383 44 NF NF 59 55 NF NF
Kwai May Pink 72 205 176 103 64 24 28 23
Wai Chee 116 148 97 111 65 48 78 19

High temperature enhance maleness (due to ↑ in GA and ↓ in cytokinin)

Apart from development of various floral parts, temperature also
determines the type of inflorescence e.g exposure of litchi trees to day/ night
temperatures of 30/ 25 oC and 25/ 20oC suppressed flowering and temperatures
of 20/ 15 and 15/ 10 oC gave rise to variable proportions of vegetative, leafy
panicles and leafless panicles depending on the cultivars (Menzel, 2003).
In temperate fruits, normally the time of flower bud initiation is closely
related to the growth cessation of shoots on which they set. The period
ranges from growth cessation (low temperatures) up to appearance of floral
primordia in the period of physiological differentiation. Among the temperate
fruit crops, chilling requirements of buds must be properly fulfilled in order
to have regular flowering. In apple cvs San Miguel and Cella, known for
their late flowering have very high chilling requirements of 1400 h and 1600
h, respectively at 7.2°C (Tabuenca, 1983). When these requirements are not
properly met during mild winters, anomalies in flowering such as deformed
and non viable floral parts, flower bud abscission and delayed bloom, are
 Physiology of Flowering in Horticultural Crops 91

observed leading to poor crop. Normal development of reproductive growth

in peach cv. Hakuho is impaired by high temperature of 25°C (Kozai et al;
2004). Frosts can reduce the number of buds, flowers and fruits in apricot.
An increase in temperature during the time of flower bud growth induced
earlier flowering thus reducing fruit set. Warm conditions during pre bloom
resulted in reduced pistil weight and short stylar length. During flower bud
development the sensitive period to temperature is from separation of bud
scales to anthesis and high temperature during this time leads to abnormal
flowers with short styles and un swelled ovaries hence reduced fruit set
(Rodrigo and Herrero, 2002).
An adequate fruit set on sour cherries has always been a problem because
it blooms somewhat later than most stone fruits when temperatures are high
and at these temperatures the blooming period becomes shorter. Uncertainties
in pollination and fertilizations also occur e.g honeybees do not visit sour
cherry (Nyeki et al; 2006).
2. Water relations : The tropics have distinct rainy and dry season. Water stress
substitutes for chilling. Excessive rainfall or irrigation which maintains
extension of growth late into summer will often reduce flower bud formation,
but this is inhibited under moderate drought and nearly completely prevented
under severe stress. Plant water stress has generally been presumed to provide
the stimulus for flowering (Schaffer et al; 1994). The proportion of apple
buds becoming floral from 12 to 54 per cent was increased by exposure to
low (45- 55%) instead to high (80- 100%) relative humidity for 9 weeks
from full bloom (Tromp; 1984). According to Pongsomboon et al. (1991), in
the absence of cool temperatures (< 15°C) mango trees in the tropics may
flower in response to irrigation or rains following water stress of 6-12 weeks
or more. The primary impact of water stress on mango is to prevent vegetative
flushing during the stress period. The accumulating age of stems is greater
in water stressed trees than in trees maintained under well-watered conditions,
which can vegetatively flush more frequently (Davenport, 1993). This delay
in flushing may provide more time for accumulation of a proposed flora
stimulus or reduction in the level of a putative vegetative promoter.
Several waves of bloom are observed in Eureka lemon by withdrawing
irrigation at a suitable time of the growing season and may be induced to
concentrate their bloom production at a season suitable for the production of
summer lemons (Monselise et al; 1981). Induction of flowering through
water stress was also practiced in some citrus species. The summer production
of lemons following water stress is known as Vardelli in central India. A
water stress period of 30-40 days was found sufficient for successful induction
of ‘Ambi’ flowering in Nagpur mandarin grown in medium deep black clay
92 Fruit Tree Physiology

soils. Similarly, moderate drought reduced shoot growth by 20–30 per cent
and increased number of flowers per lateral by 40 per cent as compared to
well watered control, however, overall fruit set was not adversely affected.
It is hypothesized that there is accumulation of ammonia during stress
resulting in increased biosynthesis of arginine, polyamines and subsequently
in increased rate of cell division following release from stress, ultimately
leading to floral initiation.
3. Nutritional factors : According to Kraus and Kraybill (1918), the
concentration of sugars should be greater than that of nitrogenous compounds
for flowering. Limiting factor for flower formation in the off year of alternate
bearing cultivars of C. reticulata has been attributed to levels of stored, non
structural carbohydrates in the form of starch (Goldschmidt and Golomb,
1982). Reduced flower production occurs when nitrogen fertilizer application
prolongs the period of extension shoot growth and delays terminal bud
formation.
Ringing of the branches in late summer or early winter in mango trees
can induce flowering in the “off” years and increase flowering in the “on”
years, also gives supporting evidence to the fact that nitrogen and carbohydrate
reserves play an important role in floral initiation in mango (Panday, 1989
and Malik, 1951) by increasing the C/ N ratio of shoots.
Potassium has a positive effect on flowering as it enhances amino acid
formation, which stimulates the formation of IAA oxidase, thereby flower
induction as well as carbohydrate levels. K might be effective through pyruvate
chinase, which in turn would determine the levels of several amino acids
(Fabbri and Benelli, 2000).
However, C/ N ratio theory is inadequate to explain floral bud
differentiation in fruit trees (Baxter, 1970). He further emphasized that the
wood of perennial trees was rich in carbohydrates and they also stored large
quantities of amino acids and amide nitrogen. It was difficult to conceive that
either of these could ever be inefficient for floral initiation, unless the trees
were nitrogen deficient. Normal flower development requires adequate mineral
elements in the proper balance. Deficiency of nutrients like Zn, Cu and B is
detrimental to flowering e.g boron deficiency in pear during anthesis causes
withering of flowers. 1-2 per cent NH4NO3 or 2- 4 per cent KNO3 have been
found to be effective for initiating floral buds in mango (Nunez-Elisea, 1985).
Maximum flowering was obtained by spraying of urea (2.5%) and etheral
(200 ppm) in guava by Chandra and Govind (1994).
 Physiology of Flowering in Horticultural Crops 93

Available nitrogen, potassium, phosphorus, calcium and magnesium of the

medium were increased by the application of natural zeolite. Net
photosynthetic rate, stomatal conductance, water use efficiency, mesophyll
efficiency, petiole length, leaf area, specific leaf weight, fresh and dry weights
of shoots and roots, fruit weight and number of achenes of strawberry were
also increased by zeolite application (Abdi et al; 2006). The methods to
induce alternate bearing in Golden Delicious cultivar of apple is depicted in
the figure.

Fig. : Fertilizing, ringing, fertilizing + ringing effects on poor bearing year bud
formation.

4. Plant growth substances :

(i) Gibberellins : GA is a primary inhibitor of floral initiation. A specific
stage must be attained for a bud to be induced and begin to initiate
flowering. Initiation at the apex begins only after a critical node number
is reached. This critical number is 20 for apple cv. Cox’s Orange Pippin
and 16 for Golden Delicious. The time interval between successive nodes
is termed as plastichron. The plastichron for Cox is 7 days (Fulford,
1966). GA extends the plastichron and the bud never reaches the receptive
stage, thereby indirectly inhibiting flowering (Luckwill, 1974). Moreover,
GA makes trees over vegetative and flowering may be less than desired.
GA also influences sex expression in papaya. Application of GA either
once or twice before flowering @ 0, 50 and 100 mg/ litre for two
consecutive years in mango revealed that treated trees had two flowering
dates (January to March and April to May) and two harvesting dates
(June and July) while control trees showed only one flowering date
(January to March) and one harvesting date (June). Treated trees produced
94 Fruit Tree Physiology

about 16 per cent while untreated trees could not produce mixed panicles.
GA delayed flowering and harvesting dates by 90 and 42 days,
respectively. However, fruit yield was not affected by GA3 applications.
Application of gibberellic acid (GA3) on citrus consistently reduced flower
formation, but had a variable effect on the amount of first grade fruit in
the early harvest of Clausellina Satsuma (Citrus unshiu Marc.), but these
applications had no significant effect on the value of crop in the long
term (Duarte et al; 2006).
The application of gibberellic acid during flower bud induction
significantly reduced flowering of Black Diamond and Black Gold
Japanese plums. Flowering was reduced upto 40 per cent by GA3 50 mg
/ l and upto 75-90 per cent by GA3 75 mg / l or higher concentration in
mixed shoots. With regard to spurs, reduced flowering intensity was
reduced by 40 and 25 per cent in Black Gold and Black Diamond,
respectively by GA3 50 mg / l in spurs, while GA3 75 mg / l or higher
concentration reduced flowering by 70 and 50 per cent, respectively.
This partial inhibition of flowering significantly reduced the cost of
manual thinning. The best GA3 concentration was found to be 50 mg/ l,
since it reduced the cost of thinning by 45-47 per cent and increased
final fruit weight by 7-33 per cent for Black Diamond and Black Gold,
respectively (Gonzalez et al; 2006). He also reported that the application
of gibberellic acid during the flower bud induction period significantly
reduced flowering in peaches and nectarines. Peach was higher sensitive
to GA3 in comparison with nectarine. Flowering was reduced by 50 per
cent in both peaches and necterines which reduced the cost of thinning
by 50 per cent in both crops without affecting the yield when GA3 was
applied at 0.5-1 mg/ tree.
(ii) Auxins : Auxin is mainly used for floral induction in pineapple. The
induction of flowering in pineapple with auxin was first reported as early
as 1939. Among the auxinic substances, NAA and NAA based compounds
like planofix and celmone have been reported to be effective in induction
of flowering @10- 20 ppm. Auxin stimulates ethylene biogenesis to
induce flowering in pineapple (Burg and Burg; 1966). Increased ratio of
perfect to staminate flowers has been obtained in mango with the
application of NAA @ 100-200 ppm about a week following auxin
treatment (Singh et al; 1965). Ubi et al. (2007) determined the effect of
three forcible hormones calcium carbide, NAA and and B-hydroxyethyl
hydrazine on the flowering and fruiting of two pineapple cultivars and
found that B-hydroxyethyl hydrazine at 0.50 g/ litre produced flowers
within 8 days and was significantly superior to all other treatments.
 Physiology of Flowering in Horticultural Crops 95

Calcium carbide applied at 30 g/ litre was more outstanding in inducing

flowering and fruit production than 25 or 35 g/ litre concentrations.
Increased production of flowers was obtained by increasing the
concentration of NAA from 0.25 to 0.50 g/ litre and from 0.50 to 0.75
g/ litre. However, increased B-hydroxyethyl hydrazine from 0.50 to 0.75
g/ litre lead to 51.5 per cent drop in flower production and 34.1 per cent
drop in fruit production
The size to which the fruit develops is directly dependent on the
number of leaves on the plant at the time of flowering in pineapple but
here the problem is to obtain flowering at the proper time. In Cabezena
variety of pineapple, which flowers poorly when left to itself, can be
made to flower at any time of the year by a single application of an auxin
- naphthalene acetic acid or 2, 4- dichlorophenoxyacetic acid. Uniform
fruits of a selected size can be produced by applying the auxin to plants
having appropriate number of leaves. Flowering in pineapple can be
induced by pouring NAA (10 ppm) + urea (2%) @ 50 ml per plant into
the crown.
(iii) Cytokinins : The precocious bud break and flowering in mango shoots
in response to an early October application of 100mg/ l 6-BA was
described by Chen (1985). It also resulted in full flowering in one month
after application as compared with 3 months later on non fruited trees.
Naunez-Elizea et al. (1990) while using synthetic cytokinins thiadizuron
also reported similar results. Highest level of putative transzeatin and its
ribosides were measured during the early flowering and full bloom stage
while the lowest level were translocated from roots during the vegetative
growth and resting state.
It is generally seen that upward flow of cytokinins in xylem sap of
trees reaches a peak in spring about the time of full bloom. There was
increased percentage of hermaphrodite flowers in mango when 50 ppm
BA plus 2 per cent Ca2+ was applied through foliar application (Singh
and Rajput, 1990). The elevated cytokinins level found prior to and
during flowering and the flowering response to applied BA leads to the
conclusion that cytokinins are involved in flowering in mango (Chen,
1985).
(iv) Ethylene : Numerous investigations show that ethephon is an effective
floral promoter of some mango varieties under specific conditions found
in the low latitude tropics (Das et al; 989). Exposure of trees to smoke
from burning leaves has been utilized to stimulate flowering in mango
in Phillipines (Pantastica and Manuel, 1978). Flowering was induced
96 Fruit Tree Physiology

during “off” year in mango cv. Alphonso by spraying ethrel (2-chloroethyl

phosphonic acid) @ 200ppm. Rabelo et al. (1999) reported that ethephon
@ 0.25ml/ l accelerates the flowering on both ringed and non ringrd
mango trees in cv. Haden. Flowering in pineapple and plant species
belonging to the Ananaceae and the bulbious plants is induced by ethylene.
Ethylene is commercially used as a flower inducing agent in pineapple
cultivation and acts as a florigen of the Ananaceae. On the other hand,
ethylene inhibits flowering in many other plant species. However, it is
not clear whether ethylene synthesis is induced under inductive
photoperiods, or whether ethylene is actually involved in the regulation
of flowering.
More than 90 per cent flowering was induced in pineapple by etherel
or ethephon @ 25 ppm in combination with urea (2%) and CaCO3 (0.04%)
after 50 days of application (Dass et al; 1989). Aldrich and Nakasane
(1975) reported 100 per cent flowering by the application of dry calcium
carbide powder or with water into the core of plants in cv. Giant Kew.
The flower and fruit number was increased by spraying ethereal @ 1ml/
l due to greater ethylene forming enzyme activity in leaves leading to
maximum number of flowers and fruits in guava (Castelen and Becerril,
1994). Etherel has been tried to induce flowers in “off” season in different
cultivars of mango but the results obtained were erratic. However, results
have indicated that ethylene may promote flowering in mango only when
conditions for bearing are not otherwise unfavourable (Sant Ram, 1977).
5. Other chemicals : Plant growth retardants like Alar, cycocel (CCC) and
triazoles like paclobutrazol, uniconazole are used to induce flowering in fruit
crops. Cycocel @ 1000 ppm and SADH @ 500- 1000 ppm favours floral
initiation in pear and apple, respectively. CCC and TIBA advanced flowering
by 9 and 4 days, respectively in Co- 2 papaya was reported by Bhattacharya
and Madhav Rao (1992).
Among triazoles, paclobutrazol is commonly used to stimulate flowering.
Paclobutrazol @ 2.5 to 15 g a.i./ tree significantly increases the percentage
of reproductive shoots in mango and also causes extension of the flowering
period (Ferrari and Sergent, 1996). Paclobutrazol @ 10g per tree had a dual
effect i.e reduction of tree size and induction of early flowering and cauliflory
(Kulkarni, 1988). It was suggested that under rain fed conditions, a single
application of 10g PBZ/ tree should be followed by further application every
two years. According to Rao and Srihari (1998), foliar sprays of 100 ppm
TIBA or 2000 ppm PBZ or soil application of 10g PBZ in mango promoted
flowering directly on fruited shoots in the “off” year by passing the need for
 Physiology of Flowering in Horticultural Crops 97

vegetative phase. Soil application of paclobutrazol (10g ai.tree-1) significantly

increased the ratio of perfect to staminate flowers in mango (Kurian and Iyer,
1993). Biennial bearing in apple, pear and cherry can be overcome by
compounds like SADH and TIBA through maintaining the hormonal balance
rather than C: N ratio. Application of CCC @ 500 ppm resulted in earliest
flowering and maximum number of flowers in guava (Brahmachari et al;
1996). Paclobutrazol at 2 g/ plant increased flower bud number in mango
(Ataide et al; 2006). Other chemicals like benzoic acid, nicotinic acid,
nicotinamide and L-pipecolic acid act as flower inducing substances, however,
their role as endogenous regulators of flowering has not been proved yet.
6. Cultural practices : Several cultural practices like spreading of branches,
trunk or branch girdling, defoliation and deblossoming, chemical treatments
and pruning are used to provide vigour control and initiate blossom bud
formation in fruit crops.
(i) Spreading, orientation, bending and shoot type : Spreading of branches
in apple towards more horizontal position reduces growth and promotes
flower bud formation. The major factor controlling floral initiation on
long shoots of Williams Bon Chretien pear appeared to be the pattern of
growth. The number of fruit buds which developed per shoot was inversely
correlated with the relative growth rate of the shoot before harvest.
Training, bending or placing apple shoots horizontaly checks their growth
and increases the proportion of the buds that become floral (Robbie et
al; 1993). Shoot bending led to an increase in the sorbitol, amino acid,
IAA and cytokinin content of lateral buds and a decrease in these in
shoot tips. Bending was accompanied by a decrease in the gibberellin
content of both shoot tips and axillary buds. The shoots that placed
horizontal after cessation of their growth produced more flowers than
when left vertical (Tromp, 1984). The bending of branches of young
trees at an angle of 45° from the main trunk may induce flowers earlier.
Orientation of apple trees at 45°C from soil surface reduces vegetative
growth, tree canopy volume and promotes spur formation. The physiology
behind bending is that it restricts carbohydrate movement and auxin
from the upper portion of the limb towards the roots. An accumulation
of carbohydrates and slowing down of the growth beyond the bend is,
therefore, assumed to be favourable to flower bud formation.
(ii) Girdling : Trunk girdling has been used in mango trees to promote
flowering by Pandey (1989). The actual cause of flowering by girdling
cannot solely be attributed to carbohydrate accumulation but ethylene
produced may also play an important role as it is produced under stress
conditions (Leather and Abeles, 1972). It has been found to increase
98 Fruit Tree Physiology

flowering in the “off” year of alternate bearing cultivars, although

it either has no effect or is marginally beneficial in the “on’” year
(Rameshwar, 1989). Flower initiation in apple is also achieved through
girdling or scoring of trunk. Girdling increased the bud sprouting, flower
formation and reduced the number of vegetative shoots in the early and
late harvested trees in Satsuma mandarin (Lewis and McCarty, 1995).
Tree response is dependent on the width of the girdle. Narrow cuts result
in either short term or no response, whereas too wide girdles can kill
trees through root starvation. It is observed that in apple ringing and
scoring (making two or more parallel cuts to sapwood around limb without
removing the bark) can make the trees more susceptible to winter injury
during the following winter. Garcia and Martins (2006) observed that the
girdling in the main branches in litchi induced higher flowering, without
any effect on the flower panicle characteristics and there were no
differences in the fruit set, but with an increase in the flowering, the
production increased.
(iii) Defoliation : Defoliation alters the cytokinin to auxin ratio in the buds
and stimulates flowering. Because apical leaves are the primary source
of auxin, their removal results in initiation of new shoots, perhaps due
to increase in cytokinin to auxin ratio. Growth of dormant axillary buds
is stimulated by de blossoming and produces inflorescences if initiation
occurs under conditions conducive to floral morphogenesis. When leaves
in apple are stripped one month after harvest, permits the trees to initiate
flowers before the next cycle begins (Janick, 1974). Defoliation
experiments have shown that leaves promote flower bud formation. Ryugo
(1986) reported that no flower formed on apple spurs that had been
defoliated within 6-10 weeks after full bloom but the number of flower
buds formed on spurs defoliated after this period gradually increased.
The effect of leaves has been attributed to their effects on carbohydrate
supply and on the flow of cytokinins.
(iv) Use of chemicals : Flower induction has been achieved by several
chemicals like dinitro compounds, KNO3, thiourea, cyanamides and
mixture of cytokinin and gibberellins in fruit trees. Application of KNO3
@ 5-7 per cent prior to oil and DNOC spray improved flower bud
development and reduced the number of abnormal flowers in certain
peach cultivars. Hydrogen cyanamide is found to be useful for enhancing
bloom and synchronizing time of bloom for varieties and their pollinizers
in several temperate fruit crops. Promalin, a combination of BA and
GA4+7 is used to cause lateral bud break in sweet cherry, a species with
strong apical dominance.
 Physiology of Flowering in Horticultural Crops 99

Initiation of flower buds in mango has been achieved by spraying KNO3

@ 2-4 per cent or NH4 NO3 @ 1-2 per cent (Naunez-Elisea et al; 1988).
CaNO3 resulted in advanced flowering in mango cv. Irwin by one month.
Plants treated with 0.4 per cent Zn in combination with Fe and Boron @
0.1 per cent produce more flowers in mango cv. Fazli.
(v) Pruning and pinching : Pruning has been found to be beneficial to
promote flowering in many fruit crops. Increasing severity of pruning
usually decreases flowering and cropping (Forshey and Elfying, 1989),
and heading back methods of pruning may convert potential fruiting
spurs into shoots. Inadequate pruning may, however, result in excessive
vegetative growth within tree shade and inhibit flowering (Feucht, 1976).
Saidha et al. (1983) stated that though pruning had no significant effect
on time of floral bud formation, but it enhanced the leaf ethylene level
and percentage of flowering in cvs. Mulgoa, Neelum and Bangalora.
Pruning to a length of 30 cm resulted in the greatest flowering in mango
cv. Haden (Gill et al; 1998). One leaf pair pruning in guava cv. Sardar
has been found to be superior to initiate flower bud formation (Lal et al;
1996). In Ber, blossom initiation can be regulated by time of pruning.
Flowering was advanced by about 3 months i.e. during summer by pruning
during spring instead of at the normal time i.e 2-3 months later (Pareek,
1983).
Pinching out of the shoot tip in apple at right time helps in
development of buds was reported by Crabbe (1984). In Japanese pear
(P. serafina), most flower buds are produced on young wood, especially
in cultivars which commonly produce flowers from lateral buds, and
pruning systems are designed to maximize the proportion (Klinac et al;
1995).
7. Root stock : Flowering is also influenced by the choice of root stock. Young
trees on selected clonal/ dwarfing root stocks often flower earlier than those
on seedling root stocks e.g apple trees on vigorous M1 root stock flower
earlier and heavier than trees on vigorous seedling root stock. It has been
postulated that root stocks influence flowering through their ability to reduce
scion chilling requirements as observed in pear and apple. When pear cv.
Bartlett was grafted on low chilling Pyrus calleryana root stock it required
only 850 hr of chilling as compared when grafted on P. communis which
required 1500 hr (Westwood, 1964). Buds of P. calleryana have been found
to show a much higher rate of development during autumn, when temperatures
are declining, than buds of P. communis. This is attributed to the relatively
higher rate of alternate pathway (cyanide-resistant) respiration under low
100 Fruit Tree Physiology

temperature conditions in P. calleryana which require more energy for growth

and development. Apple cv. Rome Beauty required less chilling and flowered
earlier on M26 root stock as compared to MM 104 or MM 106, both of which
are derived from the long chilling Northern Spy apple (Couvillon et al; 1984).
Apple root stocks have a major effect on the proportion of buds that become
floral. Blasco (1976) reported 48, 38, 28 and 32 per cent of all Cox buds to be
floral on M9, M26, M7 and MM106 respectively in mature apple trees.
8. Crop load : Heavy cropping in one year can inhibit flower bud initiation and
so reduces flowering in the following year. Some cultivars consequently tend
to become biennial in cropping. In the year of a heavy crop (on year), the
demand on carbohydrate supply is such that few flower buds are formed for
the next year in case of mango and apple and hence leading to an off year
(alternate bearing). Cultivar effects on the pattern of flower and fruit
development may be related to differences in the pattern of floral
morphogenesis within developing floral buds in the previous season in apple
(Hoover et al; 2004).

Adult Phase is Necessary for Flowering

The juvenile period is the time following seed germination during which the
seedlings cannot be induced to flower by any means. Adult phase starts after
completion of juvenility. Leaf shape, specific leaf and node number, phyllotaxy,
thorniness, ability to form adventitious roots and other character may change at
or just before the time the plant is able to flower. So, the attainment of adult stage
for flowering is essential.
Neither photoperiod nor temperature induces flowering in papaya. Each plant
commences flowering only after a genetically determined number of nodes which
develop following a short period of juvenility. As flowering in papaya is related
with number of leaves produced. Short statured cv. Betty produced first
inflorescence at 24th node and cv. Solo (a tall statured) may produce at 48th node
(Storey, 1986).
Similarly, in banana, in central America differentiation of bunch in cv. Gross
Michel was found to occur only after the production of 45 leaves. However, for
cv. Dwarf Cavandish 23 leaves were considered critical and in cv. Grande Naine
flower initiation took place after 29 leaves. Neither a diurnal temperature differential
nor short days are necessary for natural flowering in pineapple. Pineapple plants
generally flower after attainment of certain vegetative growth and ripeness to
flower stage is attained 11-12 months after planting and formation of at least 40
leaves.
 Physiology of Flowering in Horticultural Crops 101

Molecular and Genetic Aspects of Flowering

Gregor John Mendel showed that the color of the flower is controlled by the
action of a pair of heritable factor, now called genes. They are independent
discrete units of heredity. Inductive stimulus should transform the entire shoot
apical meristem (SAM) into flower inducing meristem; there again from the same
meristem different organs like sepals, petals; stamens and ovary have to develop.
The flower is indeed a modified shoot with leaves representing different parts of
the flower. Stamen and ovary differentiation is more complex than sepals and
petals for the simple reason they have highly specialized structures and they are
gamete producing super-specialized reproductive organs. Whether it is light induced
pathway or vernalization pathway or GA induced pathway or autonomous pathway,
signals are produced and translocated to phloem and the base of SAM. The
important component that induces FT is CO which activates the expression of
FT which is a 20Kda protein. The FT in association with FD (TF) activates AP1
and LFY; this leads to the activation of floral organ genes.
Three types of genes regulate floral development. Mutations have identified
three classes of genes that regulate floral development (Coen, 1991). One group
of genes directly controls floral organ identity. The proteins encoded by these
genes are transcription factors that likely control the expression of other genes
whose products are involved in the formation and/ or function of these organs.
Genes of the second group act as spatial regulators of the floral organ identity
genes by setting boundaries for their expression. These have been called cadastral
genes (from the word cadastre, which is a map or survey showing property
boundaries for taxation purposes). The third group of genes is necessary for the
initial induction of the organ identity genes. These genes, which are the positive
regulators of floral organ identity, are called meristem identity genes.
Genetic Control of Flower Development

Taiz and Zeiger, 2006

102 Fruit Tree Physiology

(Taiz and Zeiger, 2006)

The figure shows the multiple developmental pathways for flowering in
Arabidopsis: Photoperiodism, the autonomous (leaf number) and vernalization
(low temperature) pathways, the energy (sucrose) pathway, and the gibberellin
pathway. The photoperiodic pathway is located in the leaves and involves the
production of a transmissible floral stimulus, FT protein. In LDPs such as
Arabidopsis, FT protein is produced in the phloem in response to CO protein
accumulation under long days. It is then translocated via sieve tubes to the apical
meristem. In SDPs such as rice, the transmissible floral stimulus Hd3a protein
accumulates when the repressor protein, Hd1, is not produced under short days,
and the Hd3a protein is translocated via the phloem to the apical meristem. In
Arabidopsis, FT binds to FD, and the FT/FD protein complex activates the AP1
and SOC1 genes, which trigger LFY gene expression. LFY and AP1 then trigger
 Physiology of Flowering in Horticultural Crops 103

the expression of the floral homeotic genes. The autonomous (leaf number) and
vernalization (low temperature) pathways act in the apical meristem to negatively
regulate FLC, a negative regulator of SOC1. The sucrose and gibberellin pathways,
also localized to the meristem, promote SOC1 expression.

Three types of genes control floral identity :

1. Gene A activity controls the first and second whorls.
2. Gene B activity controls the second and third whorls.
3. Gene C activity controls the third and fourth whorls.
Transformation of some flowering genes in fruit crops to induce flowering.

Fruit crop Gene Induced character

P. trifoliata FT homolog Accelerate flowering

P. communis FT homolog Accelerate flowering
Citrus Arabidopsis API gene Induced early flowering
Citrus Arabidopsis LFY gene Juvenile phase reduced
Apple Suppression Md TFL I Reduced juvenile period
Apple FRUITFULL(FUL)- homolog Early flowering
104 Fruit Tree Physiology

Sen (1978) has suggested some molecular aspects of flowering such as

1. Gibberellins promote flowering in some plants.
2. Steroids promote flowering in some plants.
3. Protein component of phytochrome undergo some conformational changes.
4. RNA- is synthesized.

Mango flowering model (single lines are promotive and double ones are inhibitory).

Genes regulating flowering in some plants :

● elf 3 (early flowering 3)
● eafl (early flowering 1)
● tfl (terminal flower)
● HST (Hasty gene)

CONCLUSION
1. Flower bud development involves the transformation of the vegetative apex
to a reproductive structure.
2. Three events occur during the transition of buds to flowering i.e. induction,
evocation and initiation.
3. First visible sign of differentiation is when the flat apical meristem becomes
domed, then the central meristem is partitioned and the pith meristem develops.
 Physiology of Flowering in Horticultural Crops 105

4. Photoperiodism along with hormonal balance is critical factors for flowering

rather than C: N ratio in fruit crops. Low temperature regulates flowering in
long day plants and water stress enhances flowering in mango and citrus.
5. Various theories have been put forth by many workers to explain the
mechanism behind flowering but none is still holding good.
6. Flowering in fruit crops can also be influenced by various cultural practices
like bending of branches, defoliation, chemical treatments, nutrient regulation,
girdling and pruning.
7. Growth regulators have striking effect on flowering. While auxins promote
it, GA inhibits it. NAA is being used in mango and pineapple to induce
flowering. Ethephon is a major growth regulator regulating flowering in
pineapple. Various growth retardants i.e., Alar, Cycocel, TIBA (antiauxin),
SADH and PBZ stimulate flowering by inhibiting GA synthesis. Paclobutrazol
has been widely used in mango to induce flowering during the “off” year.
8. Molecular mechanisms are still not well understood, however, many genes
that regulate flowering have been isolated. Hence, flowering is a complex
phenonmenon regulated by number of factors.

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110 Fruit Tree Physiology

Singh RN, Majumder PK, Sharma DK and Sjinha GC (1974) Effect of deblossoming
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Westwood MN and Chestnut NE (1964) Rest period chilling requirements of Bartlett
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Further Suggested Readings

Sinha RK (2004) Modern plant physiology, Narosa Publishing House, New Delhi.
Taiz L and Zeiger E (2006) Plant physiology. Sinauer Associates Inc. Publishers
Sunderland, Massachusetts 5th Edition.
 5

FRUIT GROWTH AND

DEVELOPMENT

F ruit is the structure that arises from an ovary (or several ovaries and in some
cases associated floral parts) after fertilization and supports the developing
seeds. It is an evolutionary adaptation unique to flowering plants and is designed
to aid in the dispersal of seeds by various agencies.

STAGES OF FRUIT DEVELOPMENT

In most species, fruit growth can be represented by a sigmoid curve or a double
sigmoid curve with a second burst of growth during the ripening period while
Kiwi fruit follows a triple sigmoid growth curve. Physiologically and biochemically
fruit development can be divided into four phases, which although continuous,
are separated on the basis of major activities.
Phase-I : It includes ovary development in the flower, and flowering anthesis,
a decision to abort or proceed with further development (i.e rupture
of anthers to release mature pollen).
Phase-II : This phase involves a period of most rapid cell divisions.
Phase-III : It is the period of most rapid growth when cell divisions more or
less cease, and growth is almost exclusively by cell enlargement. In
this phase, food reserves are accumulated and most fruits attain their
final shape and size before the on set of ripening.
Phase-IV : Ripening phase.
In some fruits, such as avocado, cell division may continue well in to
phase-III.
112 Fruit Tree Physiology

Fruit crop Period of cell division

Ribes and Rubus Cell division ceases at anthesis

Cherry Ceases about 2 weeks after anthesis
Plum 4 weeks
Apple 4-5 weeks
Pear 7-9 weeks
Avacado and Strawberry Continues upto maturity

Some times during cell division period, cell enlargement begins and proceeds
at a rapid rate. At blossom time, intercellular air spaces are absent or very small.
Concurrent with cell enlargement, air spaces increase to a maximum, then remain
constant for the remainder of the season. Early in the cell enlargement phase,
vacuoles form in the cells and increase in size as the cell enlarges, ultimately
occupying most of the space in the centre.
The combined growth resulting from cell division, cell enlargement and air
space formation results in a general sigmoidal (S-shaped) curve. Some fruits such
as stone fruits- current, pistachio and seeded grape have a double sigmoidal
growth curve. The first slow growth period of stone fruits coincides with the
period of pit hardening during which lignifications of endocarp (stone) proceeds
rapidly while mesocarp (flesh) and seed (kernel) growth is suppressed. Near the
end of pit hardening, flesh cells enlarge rapidly until the fruit is firm ripe, after
which growth slows and stops.

Fruit Development Stages

1. Cell division :
● Predominates after bloom.
● Smaller fruited crops generally have a shorter period of cell division.
● Extended in some by blossom thinning.
● 2 million cells in flesh of apple at anthesis require 21 doublings of cell
number.
● 40 million cells at harvest require only 4.5 doublings.
● Most post-anthesis cell divisions occur in first few weeks after bloom,
but as long as 100 days.
2. Pit hardening (stone fruits only) : Lignification of endocarp
 Fruit Growth and Development 113

3. Cell enlargement :
● Predominates later in fruit development (and after pit hardening in stone
fruit)
● Begins soon after pollination, continues through cell division stage, then
at diminishing rate until harvest
4. Fruit maturation : Final weeks (days) of fruit development

(Jackson, 1975)
114 Fruit Tree Physiology

Key Growth Stages

Dormant : Fruit buds relatively inactive. This is the over wintering stage, (applies
to all fruits).
Silver tip : Applies only to apple. Fruit bud scales separated at tip, showing
light gray tissue.
Swollen bud : Equivalent to silvertip stage in apple. Fruit buds swollen,
scales separated to expose areas of lighter colored tissue (applies to all fruits
except apple).
Green tip : Applies only to apple. Fruit buds broken at tip, showing about
1/ 16 inch (1-2 mm) green.
Bud burst : Equivalent to green tip stage in apple. Fruit buds broken at tip,
showing tips of blossom buds (applies to pear, sweet and tart cherry, plum and
prune).
Half-inch green : (centimeter green): Applies only to apple and peach. In
apple, when about 1/2 inch (1cm) of leaf tissue is projecting from the fruit buds.
In peach, when the leaf bud occurring between a pair of fruit buds has produced
about 1cm of new growth.
Tight cluster : Applies only to apple. Blossom buds mostly exposed, tightly
grouped, stems short.
Green cluster : Applies only to pear, plum and prune. Blossom buds green,
mostly separated in the cluster, stems lengthened.
Pink : Applies only to apple and peach. For apple, all blossom buds in cluster
pink, stems fully extended. For peach, when the blossom bud shows a pink tip.
White bud : Applies to pear, sweet and tart cherry, plum and prune. Blossom
buds white, separated in the cluster and stems lengthened.
Bloom : Blossom buds open (applies to all fruits).
Petal fall : After about 75 per cent of the petals have fallen (applies to all
fruits).
Fruit set : A stage ranging from about 4 (cherry) to 10 (peach) days after
bloom when the blossoms that have or have not set fruit, initially, are clearly
evident (applies to all fruits).
 Fruit Growth and Development 115

Apple fruit growth stage (Chapman and Catin, 1976)

116 Fruit Tree Physiology

Factors Affecting Fruit Growth and Developement

1. Effect of hormones : Auxin, GAs and cytokinins are involved in the early
stages of fruit growth, especially in Phase-I and Phase-II, although their
exact roles and inter relations among them are unclear. In Phase-I, sources
of hormones is material tissues, but in Phase-II and III, which coincides with
embryo growth and maturation, respectively, the source of hormones is
debatable. The hormones could be synthesized in situ or translocated from
other parts of the plant via the phloem stream. There are also instances where
fruit growth is affected by auxins and/or CKs produced by the embryo and
endosperm.
Role of auxins : The influences of seed on fruit growth were shown by
Nitsch (1950) in strawberry. It is an aggregate fruit. A fruit formed from
several single carpels in a flower. Each carpel has a single seed and produces
an achene (a joint structure of seed and the surrounding ovary wall). Achenes
are borne on the receptacle, which form the fleshy fruit. Nitsch removed
varying numbers of developing achenes from young fruit of strawberries and
showed a strong correlation between the number of achene left and the fruit
growth around them. Similar positive correlations between the number of
seeds and fruit growth have been shown for other fruits (e.g grapes, blackberry,
apple etc.). He showed that exogenous application of b-napthoxyacetic acid,
to the deachened receptacle could replace the stimulus of fertilization for
fruit growth, which suggested that young seeds with developing embryo and
endosperm could be a rich source of auxin and that this auxin was the trigger
for continued growth of fruit. Endogenous IAA in developing fruits of tomato
(Buta and Spaulding, 1994) shows highest level at the earliest time of fruit
growth. The level then drops steadily and again rises to an early stage of
ripening.
In addition to auxin; CKs and GAs also play a role in the growth of the
young fruit. Cytokinin levels specially zeatin and zeatin riboside are high at
earliest stage of several fruits and correlated with period of maximum cell
division. In kiwi fruit, the concentration of total cytokinins was high at the
earliest time after anthesis, dropped to low levels later (Lewis et al; 1996).
The cultivar Hass of avocado produces two populations of frutis one
significantly smaller than the other. The cell size in two phenotypes is same,
but the smaller phenotype results from a reduced number of cell divisions,
which is related to a reduced cytokinins and abscisic acid ratio in the smaller
as compared to the normal phenotype. The deficiency is correlated and fruit
size is restored to normal size by an exogenous application of cytokinins,
isopentenyladenine in phase I (Moore-Gordon et al; 1998).
 Fruit Growth and Development 117

2. Fruit size and shape : The shape and size depend on the number of cell
divisions and directionality of growth of daughter cell in various sectors of
fruit in phase II and III. The growth of banana requires cell growth mostly
in a longitudinal direction and growth of pear fruit would be required altered
patterns of cell division and growth in different sectors of the fruits.
Factors that affect cell size and thereby the fruit size

Increase in cell size Decrease in cell size

Few cells per fruit Many cells per fruit

Light bloom at set Heavy bloom at set
Adequate soil moisture Low soil moisture
Strong fruiting spurs Weak fruiting spurs
Centre bloom fruits Side bloom fruits
High leaf: fruit ratio Low leaf: fruit ratio
Healthy leaves Choloretic or diseased leaves
Moderate to excess thinning No thinning

http://fruitgrowth.tfrec.wsu.edu/
118 Fruit Tree Physiology

http://fruitgrowth.tfrec.wsu.edu/

3. Leaf ratio : Leaf ratio plays an important role in fruit growth and
development. It varies from crop to crop e.g 30-40 leaves are considered
optimum for apple.

http://fruitgrowth.tfrec.wsu.edu/
 Fruit Growth and Development 119

4. Photo assimilates : During the rapid growth in Phase II and III, fruits act
as strong sink and import massive amount of photo assimilates from
photosynthesizing organ. Such translocation occurs in the phloem tissue and
the translocated materials are mostly sucrose, although in some species
oligosaccharides (rafinose) or hexitols (manitol, sorbitol) may be the
predominant sugar/ sugar alcohol. Hormones have been implicated in
regulating the partitioning of photo assimilates between competing sink. In
many studies, ABA has been suggested as the hormone that facilitates
unloading of the photo assimilates at the sink site, but evidence is equivocal.

FRUIT GROWTH AND DEVELOPMENT IN

SOME IMPORTANT FRUIT CROPS
Mango
The flowering period in mango is usually of two to three weeks. It is achieved by
low temperature, whereas high temperature shortens it. It bears mainly two types
of flowers – male and perfect, though neutral flowers are also found occasionally.
The percentage of perfect flowers varies and is influenced by the environment in
which these are grown. During the period of panicle development, low temperature
increases proportion of male flowers. The fruit set is directly related to proportion
of perfect flowers initially but final retention appears to be genetic characteristic
of the cultivar e.g the initial fruit set in Langra is very high, but ultimately retention
per panicle is better in Dashehari which has a relatively lower proportion of perfect
flower (Majumder and Sharma, 1996). Mango is highly cross pollinated crop.
Pollination takes place through insects (common house fly, Melipona sp and
Syrphidae). Dashehari, Langra, Chausa and Bombay Green are self-incompatible.
Cluster fruit set : The incidence of cluster setting in cv. Dashehari was
studied by Ram and Rajput (2004) during 1982-1998 and they reported that
during this period, 9 “off” years and 8 “on” years occured. Cluster setting was
observed in 7 years, whereas normal fruit setting was observed in the remaining
10 years. The incidence of cluster setting was low in 1982, 1986 and 1998, but
severe in 1984 (85%), 1990 (70%), 1992 (75%) and 1993 (80%). About 95 per
cent fruitlets from the cluster had no seeds and had brown shriveled embryos. The
majority of the fruitlets were misshapen. The clustered fruitlets grew slowly and
hardly developed beyond the marble stage, eventually dropping off from trees. In
1998, cluster setting was accompanied by excessive vegetative growth, and this
incidence reduced yield (608.30 kg/ ha), which was significantly lower than the
yield obtained during the preceding “off” year (2563 kg/ ha). During this year,
54 per cent of the trees did not bear fruits, and the yield of 30 per cent of the
trees was low. Crop loss due to cluster setting varied from 65.30 to 74.80 per cent
120 Fruit Tree Physiology

depending on the severity of the disorder. Crop loss reached 60-80 per cent in
1993 and 72.26 per cent in 1998.
Fruit growth and development : The fruit growth followed a sigmoid pattern
and in general, growth is rapid between 30 to 90 days after fruit set. The fruit
volume is initially more than that of fruit weight and the difference of fruit
volume and weight is narrow up to 70 days after fruit set. Afterwards, the fruit
weight was greater than that of the volume. With the advancement of maturity the
number of lenticels decreases. The mango seed also follows a sigmoid growth.
The rapid increase in length and diameter of seed was recorded up to 60 days
after fruit set (Dutta and Dhua, 2004).
Higher pulp content at harvest was found in the fruits of Mallika and Sunderja
than Langra and Sunderja. Specific gravity of fruits gradually decreased up to 45
days after fruit set and a linear increase in specific gravity was observed upto
maturity in fruits. Titratable acidity increased after fruit set and slowly decreased
towards maturity while ascorbic acid and moisture content of fruits decreased
after fruit set to maturity, Langra had higher values for these parameters. After
fruit set to maturity, TSS and total sugars content increased and were highest in
Langra and Sunderja, respectively (Rajput and Pandey, 1998).
There are three fruit set stages in mango i.e 1. Mustard; 2. Pea; and
3. Marble. Marble stage is related with the retention of the fruit. However, in
Dashehari, the fruit set could be considered when it is of mustard size. Sometimes
after pea or marble size, further growth of fruits is retarded and they remain in
panicle for a considerable period. The developemental physiology of mango fruit
involves changes in size, weight and in several major enzymes.
Endogenous plant growth substances : Mango fruit growth and development
is associated with changes in endogenous plant growth substances. The levels of
auxins, gibberellins, cytokinins, abscisic acids and ethylene have been determined
during growth and development by Murti and Upreti (1995). However, biosynthesis
and distribution of polyamines (PAs) in relation to mango fruit growth is unknown.
PAs are implicated in fruit growth and development because of their ubiquitous
presence in all cells (Smith, 1985). Exogenous application of PAs prior to anthesis
or at full bloom has been reported to increase the fruit set and retention in the
self incompatible mango cultivars likes Langra and Dashehari. The aliphatic PAs
(putrescine and spermidine) may play a key role in the fruit set and development
of mango (Singh and Singh, 1995).
The changes in endogenous free polyamines from fruit set were also reported
by Malik and Singh (2004) until one week before the expected harvest time in
Kensington Pride and Glen and found that PA contents of the pericarp declined
 Fruit Growth and Development 121

between fruit set and maturity from 788 to 101 mmol/ g of fresh weight (FW)
in Kensington Pride and from 736.6 to 89.6 nmol/ g FW in Glen during fruit
development. During the initial phase of fruit growth, spermidine (SPD) and
spermine (SPM) were higher than putrescine (PUT). The highest levels of free
PAs, especially SPD and SPM, at the initial stages of fruit suggest a potential role
during the cell division phase and not in subsequent fruit development. The ovule
seems to be rich source of PAs as evident from 2.3 and 2.7 fold higher PAs than
pericarp tissues in Kensington Pride and Glen, respectively.
Changes in jasmonate : Kondo and Yazama (2004) reported higher JA
concentrations in cvs. Nam Dok Mai and Narg Khangwan in the early growth
stages of skin pulp development and decreases after full bloom.
Factors affecting fruit set, growth and development in mango:
1. Temperature : The number of pollen tubes reaching the ovule in self pollinated
flowers was greatest at 20 and 25°C, but declined at 30°C and did not occur
when flowers were held at 10°C. The percentage of stenospermocarpic
(nubbin) fruits was increased by low temperatures, 20°C day/10°C night,
3 days after pollination significantly increased. Low temperature exposure
increased the percentage of numbins in different cvs as: 38.3 per cent in Nam
Dok Mai, 21.1 per cent in Kensington and 6.8 per cent in Irwin. The lower
percentage of nubbin fruit in Irwin implies a greater adaptation to cool
temperatures by this cultivar during fruit set and early development (Sukhvibul
et al; 2000).
2. Light : Fruits which receive optimum light are of maximum size and have
better physico-chemical qualities in comparison to those that receive
insufficient light.
Table : Effect of fruit position on the canopy on quality of Kensington mango fruit
(Hofman et al; 1995).

Fruit Fruit Dry Total sugars (ºBrix)

position mass (g) matter (%)

Upper 360 13.1 15.9 11.6

Middle 370 13.0 14.5 11.6
Lower 346 11.6 13.4 10.5

3. Water availability : If irrigation is cut off between flowering and the first
half of the growing period, water stress occurs and affects fruit growth rate
and final fruit size (Simmons et al; 1995). However, no effect on fruit size
was observed due to water shortage close to harvest (1.5 weeks before harvest).
Early water stress influenced final fruit size through an effect on the cell
122 Fruit Tree Physiology

number. Even when water was withheld during the second month of its
development, final fruit size was 34 per cent smaller than from non stressed
trees. When water stress occurs from the end of the first half of the growing
fruit period it alters the final fruit size by effecting the cell size as a result
of decrease in carbon assimilation and in water fluxes entering the fruit
(because of the lower leaf conductance and leaf water potential, respectively).
4. Pre flowering irrigation : Pre-flowering water deficit has a detrimental effect
on the photosynthetic activity of trees at the time of flowering, pollination
and fruit set although it may enhance the potential for development of floral
structures, which might negatively affect productivity. During wet months
the light saturated leaf carbon assimilation is highest and lowest during the
dry months on irrigated trees. If pre flowering water deficit further reduces
CO2 assimilation lack of photo assimilate increases the overall photosynthetic
activity of the tree at the time of flowering and results in more viable flowering
and more yield per tree (Gonzalez et al; 2004).
5. Nutrient sprays : Application of potassium nitrate (KNO3), alone and in
combination with urea at different concentrations on flowering, fruit set and
fruit quality of Tommy Atkins mango resulted in erratic flowering, continuous
and high intensity of vegetative growth as well as irregular bearing (Yeshitela
et al; 2005). They observed that KNO3 especially in combination with urea
(5 liter solution of 4% KNO3+0.5g urea tree-1 and 5 liters of 4% KNO3+1
g urea tree-1) produced better results for most of the flowering and yield
parameters. The qualitative parameters between the treated and non treated
trees were not affected significantly. Maximum flowering and yield due to
KNO3 was attributed to the supplimentation of nitrogen from these fertilizers.
6. Growth regulators : Application of NAA, GA3 and CPPU (forchlorfenuron),
applied 14 days after blooming, on fruit retention, yield and fruit quality of
mango cv. Arumanis showed that CPPU (10 ppm) gave the best results in
terms of increasing fruit retention, number of fruits per cluster and per plant,
weight per fruit, fruit volume and leaf area (Notodimedjo, 2000). Benjawan
et al. (2006) also observed increased fruit yield, flesh content, fruit weight
and fruit length in cv. Srisaket 007 with GA application.

Changes that Occur During Fruit Development in Mango

1. Hormonal changes :
● High concentration of auxins after fertilization in ovule and the
concentration was higher after 20-32 days after pollination.
● Cytokinin and GA like substances during cell enlargement and higher
concentration of GA was found in seeds rather than in pericarp.
● ABA was high during slow fruit growth and maturation.
 Fruit Growth and Development 123

2. Sugars : The soluble sugars of the fruit pulp mainly consist of glucose,
fructose and sucrose. During early fruit development only glucose and fructose
were detected and sucrose was present in traces which increased further with
fruit growth. The concentration of glucose and sucrose towards ripening
remained pre dominant. Starch almost disappeared during ripening which
was converted to soluble sugars.
3. Proteins : Decrease in soluble protein content till 44 days after fruit set
(DAFS) which increased again up to 96 days. The increase may be due to
the de novo synthesis of enzymes.
Enzymes like catalase and peroxidase activity was also studied and following
observations were found:
● Low activity phase
● Rapid activity phase
● High activity phase
● Steep rise to maximum activity phase
● Rapid decline to minimum phase
Maximum catalase and peroxidase activity correspond to mature stage of the
fruit.
4. Vitamins : Principal vitamins of mango are vit C, B carotene and small
amount of B complex. B carotene appears mostly during ripening, while vit
C at early stages of fruit growth.
5. Pigments : Chlorophyll disappears during fruit growth and subsequent
ripening. The principal pigments are chlorophyll, carotenes, xanthophylles
and anthocyanins.

Citrus
Citrus species usually produce a large number of flowers over the year. The floral
load depends on the cultivar, tree age and environmental conditions (Monselise,
1986). It has been reported, for example, that sweet oranges (C. sinensis) may
develop 250,000 flowers per tree in a bloom season although only a small amount
of these flowers (usually less than 1 %) becomes mature fruit (Erickson and
Brannaman, 1960., Goldschmidt and Monselise, 1977). Thus, flowering represents
a great input for citrus trees and to some extent even a waste of resources. For
some authors, however, this reproductive pattern may be linked to a survival
strategy (Bustan and Goldschmidt, 1998).
Under unfavourable environmental conditions, such as late frost or drought
or excessive rain etc. most of the citrus species produce poor fruit set and high
124 Fruit Tree Physiology

fruit drop results in poor yield. Effects of light, temperature and humidity on fruit
drop were reported by Yi et al.(1995) in bearing trees of Citrus reticulata and C.
sinensis and noticed that high or low temperatures and rainy days increased fruit
drop. Application of plant growth regulators and foliar fertilizers during adverse
climatic conditions from full bloom to the fruitlet development stage could reduce
the fruit drop. Increased fruit number per tree is achieved indirectly through an
increase in flower number as a result of better flower initiation or by a direct
effect on fruit set. Improved fruit size is brought about directly by stimulating the
growth of fruit tissues or indirectly by reducing fruit number by partial inhibition
of flower initiation or by subsequent fruit removal. Application of 2,4-D and
2,4,5-T has been found to increase fruit set markedly, improved the size and
quality of fruits in mandarin, increased titratable acidity, ascorbic acid and sugars
content of fruit.
In lemon (Citrus limon) fruit growth (polar and equatorial) follows a sigmoidal
curve. In fruits of winter flush the increase in the fruit size is slow but the fruits
remain attached to the plant for the longest duration, while fruits of rainy flush
are the earliest to attain maturity. With the advancement of the season specific
gravity and peel thickness of fruits declines gradually, although no definite trend
is discernible. With the age of fruit, juice accumulation significantly increased
and rainy flush fruits recorded the highest juice accumulation, and are closely
followed by winter flush fruits (Sema and Sanyal, 2003).

Factors Affecting Fruit Growth

Growth regulators : A complex set of hormonal interactions occur during fruit
development with gibberellins (GAs) and cytokinins generally considered to have
positive effect on fruit growth while auxins have been reported to act as stimulators
of growth and also as abscission agents. Abscisic acid (ABA) and ethylene have
been implicated in several ways in abscission.
(i) Gibberellins : Gibberellins activate cell division and cell enlargement
processes in vegetative organs (Talon et al; 1991) and therefore are generally
associated with the initiation of growth. The endogenous GAs found in citrus
fruits are mainly members of the 13-hydroxylation pathway [GA53, GA97,
GA44, GA17, GA19, GA20, GA29, GA1, epi-GA1, and GA8 (Talon et al;
1992)] leading to GA1, the bioactive GA (Zeevaart et al; 1993). Developing
fruits also contain at lower levels of GAs, such as GA4, GA24, GA25, and
GA9. It is generally accepted that GAs are involved in set and development
of citrus fruits. Several studies indicate that exogenous GA3 considerably
improves parthenocarpic fruit set and growth of self incompatible genotypes
such as Clementine that in the absence of cross pollination show negligible
 Fruit Growth and Development 125

parthenocarpic fruit set (Soost and Burnett, 1961). In developing fruits, the
GA1 levels are low just before and after anthesis and approximately double
at anthesis. This transitory rise in GA1 levels can be detected in seeded
genotypes as well as in seedless cultivars possessing high or normal ability
for setting (Talon et al; 1990). In seeded cultivars, GA increases at anthesis
are therefore induced by pollination whereas in parthenocarpic species the
rise is developmentally regulated. The intensity of abscission during the
initial phases of growth is also related to the phenology of flowering.
Interestingly, the presence of leaves increases GA1 levels and the chances of
setting (Lovatt et al; 1984). In seeded cultivars, pollination stimulates
hormonal synthesis and, therefore, increases GA1 levels in developing ovaries
(Ben-Cheikh et al; 1997). It has also been shown that in seeded varieties
exogenous GA arrested fruit drop of non-pollinated ovaries. Collectively,
these observations indicate that the increase in GA1 detected in mature ovaries
shortly after pollination is a signaling stimulus of the regulatory mechanism
that reactivates fruit development after anthesis.
(ii) Cytokinins : Cytokinins are also factors stimulating cell division. Higher
levels of cytokinins have also been found in developing ovaries at anthesis
(Hernández and Primo-Millo, 1990), as reported for GAs. In addition,
exogenous cytokinins have been reported to enhance parthenocarpic fruit
development and stimulate sink strength in developing fruits of certain
cultivars, although these regulators are not commercially used to improve
fruit set in citrus.
(iii) Auxins : They either delay or induce fruit abscission, and hence may operate
as growth hormones or as abscising agents. Auxins promote cell enlargement
rather than cell division. Although endogenous auxins also increase in
developing ovaries, it is well established that exogenous treatments do not
improve fruit set. Auxins are also high during the beginning of phase II, the
period of cell elongation, and it is at this moment when exogenous auxins
are effective increasing fruit size (Coggins and Hield, 1968). These
observations may suggest that auxins are related to cell enlargement, the
essential factor controlling fruit size during the phase of rapid growth. The
enlargement of the auxin treated fruits is apparently due to cell expansion
rather than to cell division. On the other hand, auxins may act either as
delaying or accelerating agents of abscission. During the initial phases of
abscission auxins operate as inhibitors, but once the process has been initiated
auxins appear to stimulate abscission. Here, auxins could operate through the
promotion of ethylene synthesis. It has also been suggested that the causal
reason of the dual effect of auxins on leaf abscission may conveniently be
explained by the auxin gradient concept: auxin coming from the leaf would
126 Fruit Tree Physiology

tend to delay abscission, whereas auxin moving down the stem might promote
abscission. In citrus, synthetic auxins increase abscission of developing fruitlets
during the cell division period. Auxin applications at the beginning of the
cell enlargement period may have minor effects preventing on fruit abscission
and result in fruit size increase. The prevention or retardation effects of
auxins on abscission can be perceived, however, at later stages. At the end
of phase II or at the onset of phase III, synthetic auxins are commercially
used to prevent or delay eventual pre harvest fruit drop (Agustí et al; 2002).
The application of either gibberellic acid or benzyladenine at flower
opening in Satsuma mandarin (Citrus unshiu Marc.), caused a transient increase
in cell division in the ovary wall, but had no significant effect on final fruit
size. Late fruit growth and final fruit size were increased by the application of
the synthetic auxin 2,4,5-trichlorophenoxyacetic acid, which had a specific
effect on the enlargement of the juice vesicles. The three growth regulators
enhanced vascularization in the pedicel, but the growth effects observed were
unrelated to their influence on the transport capacity of the phloem but caused
by their direct effects on the fruit tissues. The sensitivity of the fruit tissues to
the applied growth regulators changed markedly during early fruitlet
development, and was characterized culturing the fruit tissues in vitro
(Guardiola et al; 1993). Spring applications of GA3 and 2, 4-D alone and in
combination, were tested on Blood Red sweet orange trees at full bloom.
Fruit weight, diameter, peel thickness and peel quantity were significantly
decreased by the growth regulator treatments compared with control while
juice contents (%), pulp (%), reducing sugars, non-reducing sugars and total
sugars, seeds quantity and quality were significantly improved by GA3
treatments compared with control. TSS (%) and vitamin C contents were
increased by growth regulators treatments compared with non treated ones. In
organoleptic testing, taste, peel colour, pulp colour and appearance were also
improved by growth regulator treatments compared with control. In conclusion
mixture treatments performed best with regards to biochemical parameters
compared with control (Saleem et al; 2008).
Girdling : Fruit development, flowering and fruitlet compete strongly with
one another and with vegetative growth, for metabolites. The trees can not sustain
all developing structures under high flowering intensities, which triggers abscission
process that, in turn regulates fruit load (Augsti et al; 1982; Rivas et al; 2004)
indicating that carbohydrate status plays a pivotal role in determining the fruit set
capability of tree by modulating its abscisic acid, 1-amino-cyclopropane-1-
carboxylate and ethylene thereby adjusting the fruit load to match the carbohydrate
supply (Talon et al; 2000).
 Fruit Growth and Development 127

During the period of physiological fruit drop, the number of fruitlets remaining
correlates negatively with fruit growth rate which is attributed to the high demand
for carbohydrate by the developing organs. Translocation of carbohydrates to the
fruitlets depends on re-mobilization of reserve supply from leaves through de
novo synthesis (Iglesias et al; 2002), transport, sink strength, and cleavage into
hexoses through enzymatic activity (Quick and Schaffer, 1996).
Girdling can improve carbohydrate availability and thus increases fruit set
and yield. It affects the expression of genes related to the starch accumulation (Li
et al; 2003), partitioning of photosynthates, mineral nutrients and plant growth
regulators in the tree (Mataa et al; 1998). Girdling improves fruit set in commercial
orchards of Fortune, Clausellina and Satsuma mandarin, irrespective of
parthenocarpic fruit set. Girdling 15 days before anthesis and 35 days after anthesis
of ‘Fortune’, mandarin was most effective. It increases yield by 125 per cent . In
high bearing orchard best results were achieved by girdling 35 days after anthesis,
which increased yield by 28 per cent (Rivas et al; 2006).
Irrigation : Citrus requires lot of water. Lack of water can lead to problems
like leaf drop and fruit split, and can have a negative impact on fruit size, health
and sweetness. The exact amount of water required by citrus plants will depend
on its age and size. In some cases, some citrus can be tolerant of mildly salty
water, but salts can build up in the soil and damage the roots. Following table
shows the effect of irrigation on various fruit parameters.
Table : Effect of irrigation on fruit quality (Syvertsen and Hanlon, 2008).

Measurement Effect of proper irrigation

Juice content Increased

Soluble solids Decreased
Acid Decreased
Soluble solids to acid ratio Increased
Color No effect
Solids per box Decreased
Solids per acre Increased
Fruit size Increased
Fruit weight Increased
Green fruit Increased
Peel thickness Decreased

Mineral nutrients : Fertilization of citrus can improve both fruit size and
yields. Balanced fertilizers offer the good plant and tree growth with good flowering
and fruiting. The amount of fertilizer used will depend on the type of tree, the
128 Fruit Tree Physiology

age of the tree, and the place where the trees are grown. Because different
fertilizers need to be applied in different ways, follow the directions on the
fertilizer bag for the proper procedure for the age and size of your tree. Nitrogen
and potassium play important role in fruit development and quality. Higher doses
beyond certain limit are detrimental. Increased N and K nutrition increases peel
growth and thickness. For good size the N:K ratio of 2.4 and 3.0 is required.
Fertilizers like potassium nitrate (2%), MAP (2%), DAP (2%), Zn, Cu, Fe etc. are
commonly used to improve fruit size in mandarins.
Heat : Although long periods of hot temperatures are generally needed to
produce very sweet citrus like oranges, long periods of very dry, hot weather are
thought to contribute to problems with fruit splitting.

Grapes
Pollination, fertilization and seed development results in berry set. However, some
cultivars set by parthenocarpy. In seedless cultivars, fertilization occurs but the
embryo subsequently aborts which is termed as stenospermocarpy. The grape berry
growth follows a double sigmoid growth. Sasi-Kumar et al. (2004) studied berry
growth and development in Pusa Urvashi and observed a double sigmoid growth
pattern like those in typical seedless grape genotypes; however, the total duration
was remarkably short including lag phase. Growth duration was divided into three
stages i.e. stage I (0-42 days), stage II (43-49 days) and stage III (49-70 days).
Veraison stage was identified at 8th week after anthesis and berry ripening was
started from 9th day after veraison. Vines were trunk girdled at fruit set stage alone
or in combination with three concentrations of GA3 (20, 30 and 40 ppm) once and
twice after a week of first application to improve berry quality. The different
combinations of girdling and GA3 showed that trunk girdling at fruit set prior to
application of 40 ppm GA3 at full bloom hastened the berry ripening by 5 days.
Berry quality improved without affecting the yield i.e. by loosening the bunch with
uniform, large and elongated berries having better TSS/ acidity ratio.

Factors Affecting Growth and Development

1. Girdling : Girdling is widely used in viticulture to increase berry size, enhance
colour and advance ripening, although the quality of the fruits depends on
the cultivars and time of girdling. Girdling prior to fruit set improves berry
set (Jackson, 1985). In seeded varieties, girdling after fruit set has no effect
or increased berry size and decreased the sugars content while for the seedless
cultivars, girdling after fruit set has a positive effect on berry size and yield
(Dokoozlian et al; 1995). However, girdling at the beginning of the ripening
phase has been reported to enhance fruit colour of red cv. Crimson Seedless,
 Fruit Growth and Development 129

Fig. 2 : Diagram showing relative size and color of berries at 10-day intervals after
flowering, passing through major developmental events (rounded boxes).
Also shown are the periods when compounds accumulate, the levels of juice
brix, and an indication of the rate of inflow of xylem and phloem vascular
saps into the berry. Illustration by Jordan Koutroumanidis, Winetitles.
(Coombe and McCarthy, 2000).

and advance fruit ripening for white and red varieties. Double girdling, girdling
after fruit set + girdling at beginning of the ripening (veraison) + ethephon
(250 ppm) for red cv. Gulabi gave the best grape quality with lowest number
of green berries, and advance ripening by about 10 days compared with
control (Reddy and Prakash, 1989). Covering vines after training with a film
of polyethylen sheets advances bud break and fruit ripening by 6 and 11
days, respectively in table grape vineyards. Practices like covering with mesh
to prevent hail and wind damage advances bud break slightly, enhances
cluster length, berry weight and delays fruit ripening (Liuni, 1994).
Girdling after fruit set increased both berry weight and yield in Italia
grapes significantly. While girdling at the beginning of the ripening increased
soluble solids, maturity index and berry colour, decreased titratable acidity
and advanced fruit ripening by 5 days. Increased berry weight, yield and best
results as regards soluble solids, maturity index, berry colour, and earliness
130 Fruit Tree Physiology

were produced by double girdling. Mesh covering delayed ripening by 9

days, decreased soluble solids, maturity index, berry colour although yield
were higher than that of control because bunches were bigger and had more
berries and the delay in fruit ripening is probably associated with change in
micro climate (Carreno et al; 1998).
2. GA3 : To stimulate parthenocarpic fruit development in grapes and other
fruits GA3 is mostly used. GA3 at 100 ppm applied by dipping of flower bud
clusters of grapes nearly 10 days before and again 10 days after anthesis
resulted in 100 per cent seedless fruit. Single and repeated GA3 application
in Swenson Red seeded grape induced seedless fruit development. Pre and
post anthesis application resulted in most of seedless berry development and
there was no difference in per cent of seedlessness, number of berries per
cluster dipped or sprayed with 0.3 mM GA3 (Fellman et al; 1991).
3. Cluster dipping : Effects of different cluster dipping treatments (1% CaSO4,
1% K2SO4, 5%) MgSO4 and 100 ppm NAA) once for 30 seconds either at
5 days before (stage I) or at veraison (stage II) were studied by Sharma
et al. (2004). They observed decreased percentage of water berry in a bunch
at harvest over the control, with no significant differences when applied at
stage I or II on Perlette vines. K2SO4 resulted in maximum reduction in
water berry development (77.77%), followed by CaSO4 (66.31%). NAA
produced maximum bunch weight (491.9g), followed by K2SO4 (422.5 g)
and MgSO4 (420.2 g), applied at stage I. Significant reduction in bunch
weight was observed in all treatments applied at stage II, but values were
still higher when compared with control. All treatments produced an increase
in TSS and decrease in acidity.
4. Crop regulation : Pruning is the cheapest and easiest way of crop regulation.
Heavy crop load impair the quality and delay ripening, therefore balanced
pruning is considered essential. Sharma and Jindal (1981) reported that
thinning reduced uneven ripening and improved colour development.
5. Partial defoliation : Partial defoliation stimulates the development of fine
and extension roots, which may increase the absorptive capacity of the root
system, canopy density leading to increased light penetration, fruit exposure,
and photosynthetic activity of mature and old leaves. Although, partially
defoliated vines had much less leaf area per gram fresh berry mass at ripeness,
yield/ vine increased considerably with defoliation at pea size and veraison
stages. The total soluble sugars content was unaffected, but the titratable
acidity was increased and the pH of the must decreased with partial defoliation.
Increased anthocyanins, phenolics, colour density, cultivar character intensity,
and overall wine quality were obtained in partially defoliated vines (Hunter
et al; 1995).
 Fruit Growth and Development 131

6. Irrigation : The use of drip irrigation with longer duration and less frequent
application on fine-textured soils favoured water distribution in the soil,
resulting in a better status of the plant, larger root development, and greater
weight of cluster and pruning materials. At the same time it increased berry
size and soluble solids at harvest (Selles-Van-Sch et al; 2004).
7. Effect of BRs and CPPU : BRs and CPPU @ 0.2 ppm and 2 ppm, respectively
twice at 7+15 days after fruit set are used to improve the quality of berry
w.r.t size and TSS required for export (Bhat et al; 2004).

Apple
Due to lack of cultivars suitable for cross pollination fruit set of apple may be
affected. Weather conditions during blooming season, such as temperature rainfall
and humidity may also produce an unsatisfactory fruit set. Frost at bloom time
can prevent pollination by killing the style. High relative humidity (>50%) in
March, delay flowering without affecting fruit set and yield. Very low temperature
(<4.5°C) reduced fruit set by interfering pollen germination and bee activity.
Spray of aminothoxyvinylglycine (AVG 200 ppm) applied alone or with GA +
BA (50 ppm) inhibits endogenous ethylene production and increase fruit set in
treated flowers (Greene, 1980).
There is a continued enlargement of receptacle during the development of
fruit. A rapid phase of cell division occurs in first few weeks after pollination,
which cease abruptly 30-40 days after full bloom in Cox’s Orange Pippin. The
subsequent fruit growth occurs mainly due to cell expansion. The fruit growth
pattern follows a smooth sigmoid curve (Mitra, 1991).

Factors Affecting Fruit Growth and Development:

1. Flower type and position in cluster : The apical or king (K) flower in the
bud of the apple develops first followed by the lateral (L) flower which
develops in sequence beginning at the base of the spur. K flower is also first
to open and cell expansion starts earlier and ultimately results in larger fruit
(Faust, 1989). Cluster position also affects fruit cellular density (air space),
K fruits have larger cells and lower specific gravity than side bloom fruits
(Ferree et al; 2001).
2. Pre bloom factors and fruit size :
● Post-harvest defoliation : Small fruit size in the following year.
● Spur size and position : Larger spurs bear larger fruits; King bloom of
apple produces largest fruit.
132 Fruit Tree Physiology

● Age of bearing wood: Larger fruit on 2 year-old spurs than 1 year-old

spurs.
● Pre-blossom temperature: Low temperatures, smaller fruit (shorter growing
season, greater fruit set and competition or reduced pre bloom cell division.
3. Crop load and Cell number : During early stages of fruit development the
cell number in both heavy and light crop show slight or no variation, however,
as the fruit growth advances the cell number in light crop increases as
compared to heavy crop. This could be the reason for larger fruit size of light
crop.
Heavy crop Light crop
CELL NUMBER (x1000)

1600
1400
1200
1000
800
600
400
200
0
April 16 July 2 Aug 20 Oct 13

(Bergh, 1985)
4. Post-bloom factors and fruit size :
Seeds : Fruit size dependent for first 7 weeks; aborted seeds alter fruit shape.

(Williams, 1979)

Light and carbohydrates : Controls supply of CHOs resulting in competition

between fruit and shoot growth and between fruit. This is the most important
source of within tree variation in fruit growth.
 Fruit Growth and Development 133

5. Temperature : When the temperature are low the fruit growth is slow, however,
as the temperature increases the growth rate also increase because every
100C rise in temperature doubls the chemical reactions.
Fruit growth rate (mm/day)
Delicious Golden Delicious Fuji
1.0
0.8
0.6
0.4
0.2
0.0
13/3 16/6 19/9 22/12
Maximum/minimum temperatures (°C)
(Warington et al; 1999)
6. Water stress : Water stress, particularly during peak water demand period
(mid june to mid August) had a considerable effect on fruit growth. During
this period both cell division and cell elongation may be slowed down due
to shortage of water.
Apple (England)
120
100
Fruit wt (g)

Dip-irrigated
80
60
40
Non-irrigated
Rain (mm)

20
0 20
10
0
18 25 2 9 16 23 30 6 13 20 27 3 10 17
June July August September
(Goode et al; 1978)
7. Fruit thinning : Thining of fruits at peanut stage had opposite effect in
enhancing the fruit size as it reduces the competition for water and nutrients
between the sinks (fruits). Late thining may produce non significant effect.
Light & Fruit Growth ‘Cox’s Orange Pippin’ Apple

200
Mean fruit weight (g)

No thining
160 Alternate clusters removed
3 of 4 clusters removed
120 All clusters on alternate
branches removed

80
0.10 0.20 0.30 0.40 0.50
Light interception per fruit (Palmer, 1979)
134 Fruit Tree Physiology

8. Fruit shape : During early fruit development stages the L/ D ratio is more
than one while in latter stages it is less than one and is also a varietal
character. In more elongated varieties it is more than one. It also depends on
various climatic factors particularly temperature and sunshine hours.

Stem & calyx ends

change from
convex to concave

(Westwood and Blaney, 1999)

9. Growth regulators : Pre bloom applications of GA3 at 10 ppm + BA at 5

ppm + NAA at 5 ppm gave the best quality fruits (in terms of firmness, total
soluble solids and total sugars), and the most efficient in marginal apple
growing areas of Himachal Pradesh (Sharma and Ananda, 2004).
● 50 percent GA4+7 + 50 percent 6-benzyladenine (BA): Promalin stimulates
growth of apical portion of apple fruit, increased length/diameter ratio.
● 5 percent GA4+7 + 95 percent BA: Accel, a post bloom thinner that
sometimes increases fruit growth (size) of apple.
● Gibberellic acid (GA3): Pro Gibb delays maturity and increases fruit
size, soluble solids and firmness of cherry fruit.
CPPU and BA : Crop load reduction with commercial thinners is used to
increase fruit size. Although, total yield is reduced, the remaining fruits are
of higher value because of their increased size. Bezyladenine (BA) is used
to thin apples and it increases fruit size (Moran and Southwick, 2000) by
stimulating additional cell divisions within the fruit (Wismer et al; 1995).
CPPU [N(2-chloro-4-pyridyl); N-phenylurea] at 10 ppm and BA
 Fruit Growth and Development 135

(Benzyladenine) at 50 ppm resulted in significant increase (>50%) in fruit

size of Royal Gala cv. of apple when applied two weeks after full bloom due
to simulation of fruit cortex cell division (Wismer et al; 1995). BA promotes
cell division in tissues possibly by increasing zeatin ribozide levels in the
fruits (Yuan and Green, 2000). Both cytokinins increased fruit size without
affecting fruit shape and seed number, and with no reduction in the return
bloom or yield of the following year. Use of 4-6 ppm CPPU in McIntosh
apple (at fruitlet size between 6-10 mm), increases the fruit size without
causing fruit asymmetry and reduction of return bloom (Green, 2001).
Promalin : Promalin (GA+BA) sprayed at 50 per cent full king bloom and
repeated at full bloom in single and split doses to Starking Delicious apple
increased the TSS, total sugars, soluble proteins and anthocyanin and decreased
acidity. In areas exposed to warm weather during flowering and development
promalin treatments improved the quantitative as well as qualitative parameters
(Jindal et al; 2004).
10. Summer pruning : Summer pruning (removing shoots and leaves during the
growing season) has been traditionally used in apple to improve canopy light
environment thus enhances fruit colour as well as reduce tree size. However,
undesirable reduction in fruit size often have been reported as a result of
summer pruning. In addition, changes in growth and development of different
parts of tree have also been associated with summer pruning. The varied
results of summer pruning indicate that many factors such as varieties, orchard
management and environment are likely to be involved. Some studies have
noted the importance of canopy carbohydrate supply and the impact due to
loss of leaf area by foliar pests or summer pruning on growth and fruit
quality. Canopy gas exchanges and carbohydrate supply decreases in
proportion to summer pruning intensity (Li et al; 2003).
In addition, the root system of mature trees are believed to be very
sensitive to limitation in assimilate supply especially due to heavy crop
levels. Li et al. (2003) suggested that within commercial cropping ranges,
light and moderate summer pruning had slight influences on fruit size and
fresh weight. Summer pruning did not affect fruit colour, soluble solids
content, starch, fruit firmness and internal breakdown after storage.

Fruit Maturation
● Fruit growth rate slows.
● Pigment synthesis and degradation leads to color development.
● Waxy cuticle formation retards dehydration.
● Starch → sugars conversion.
136 Fruit Tree Physiology

● Respiration declines (pre climacteric).

● Flavor volatile synthesis and degradation.
● Cell wall degradation → reduced firmness.
● Pre mature fruit abscission.

Pear and Stone Fruits

Cross-pollination is required in most of the commercial cultivars of Japanese pear
(Pyrus pyrifolia Nakai), to set fruits because they exhibit gametophytic self
incompatibility. To improve fruit set and produce acceptable commercial fruit
size and perfectly round shape, artificial pollination using compatible pollen is
common practice in the pre dominant Japanese pear. Cultivars like Nijisseiki,
Kousui and Housui do not have parthenocarpic ability and are self incompatible.
Growth and development : Generally fruit set is not a problem in commercial
pear cultivars. Fruits of Patharnakh pear take 150 days to mature and physiological
development involves measuring changes in size, specific gravity (SG), skin colour,
pigment, TSS, phenolic contents, and fruit firmness. Fruit growth shows 3 phases,
active, very active and very slow, as measured by size (length and diameter) and
weight. Fruit firmness decreases during the final 50 days to harvest. TSS shows a
continuous increase during development (from 7.80 to 13.53%); this increase is
most rapid between 15 and 30 days, and again between 75 and 90 days. Reducing
and total sugars both increase continuously until harvest and the increases are
rapid during 120 to 150 days. Total pigment concentration also shows a continuous
increase (from 10.6 to 129.4 mg/ 100 g). Starch increases up to 105 days (3.76%)
then decreases until harvest (1.60%). Total phenolics increased up to 30 days
(0.278%) then gradually declined (0.021%). Fruit firmness decreased between 105
and 150 days from 24.56 to 16.48 lb/ inch2 (Mann and Singh, 1990).

Factors Affecting Fruit Growth and Development:

1. Polyamines : Pre pollination application of 1.0 mM putrescine treatment
increases fruit set in the late hand pollinated flowers of Housui Japanese pear
and pollen germination after in vitro pollination is higher in the styles of
flowers sprayed with putresine (Franco-More et al; 2005).
2. Defoliation and ethephon : Defoliation before vegetative maturity causes
reduction in peach flower and fruit number. In several peach cultivars all the
ethephon treatments had detrimental effect on flower density, fruit set and
yield in spite of bloom delay. The ethephon treated prune trees yielded more
than the untreated tree as a result of frost avoidance i.e by delayed bloom
(Miller et al; 1990).
 Fruit Growth and Development 137

3. Water stress / deficit irrigation : Fruit size is a major criterion of stone fruit
quality. The two important orchard practices that affect fruit size are fruit
thinning and irrigation. It is important to optimize crop level and water
availability, in order to maximize the number of large fruits. The response of
deciduous tree to the deficit irrigation depends on stage of fruit growth at which
it is applied. During stages I and II of fruit growth of stone fruits the deficit
irrigation did not affect yield, but deficit irrigation at stage – III decreases fruit
size and changes many quality attributes (Li et al; 1989). The crop water
consumption at stage III is much higher than that of stage II and reaches 120
per cent. However, fruit size did not respond to irrigation above 100 per cent or
0.92 of potential evapo-transpiration (Naor et al; 2004) at stage III. Irrigation
levels affects soil water availability and, consequently plant water status, shoot
growth, stomatal conductance, assimilation rate and fruit size (Berman and
Dejong, 2003). The most sensitive period of water stress in stone fruit is the
stage III of fruit growth. In plum, fruit yield and size increases with increasing
irrigation rate. Deficit assimilates availability is apparent in the low irrigation
treatment at high crop levels. Relative fruit growth rate decreases drastically
during an extreme warm dry period, when temperature exceeds 40°C. Increase in
irrigation level decreases the soluble solids content in the fruit juice (Naor et al;
2004), but fruit firmness and total acids are not affected by irrigation level.
4. Calcium : When Redhaven peach and Stanley prune trees budded on dwarfing
Prunus besseyi root stocks were grown and fruited in greenhouse sand culture,
peach fruits were smaller, greener and firmer and had less soluble solids and
red blush at 2 and 90 ppm Ca than at higher concentrations, and had poorer
flavour at 2 ppm. Optimum peach fruit quality was obtained when the Ca
supply was 180 to 270 ppm. Prunes were little affected by Ca treatments,
except that at 2 ppm, most fruits dropped after fruit set and the few that
remained were smaller and misshapen (Abdalla and Childers, 1973).

Plum
All prunus species follow the double sigmoid growth curve. During the subsequent
development of the fruit the outer wall of the ovary gives rise to three distinct
layers of the pit. Two hypodermal layers in the ovary produce the skin of the
fruit, which consist of cuticle, the epidermis and a few layers of collenchymatous
cells. The flesh contains most of the vascular bundle of the fruit, which are
embedded in the lignified tissue of the pit. The fresh fruit weight and size increases
through out the growing season, while the sugar content increases gradually and
rapidly during ripening. Dry matter content increases gradually, being sharper at
ripening. The organic acid increases initially but decreases during fruit development
(Josan and Chohan, 1982).
138 Fruit Tree Physiology

Factors Affecting Fruit Growth and Development:

1. Climatic factors : Climatic factors like late spring frosts, precipitation, rainy
days and low noon temperatures produce most negative effects on percentage
fruit set and yield. Unfavourable meteorological conditions during flowering
produce low yields and the greatest reduction in yield has been found to be
due to late spring frosts (Grabowsky and Zielenkiewicz, 2000).
2. Chemicals : Dormex spray resulted in earlier flowering by 2-5 days in
apricots and in plums by 4-8 days. Application of 1 per cent and 2 per cent
hydrogen cyanamide (Dormex, 49%) gave better flower and vegetative bud
break ratio and fruit set in dormant trees of Papaz plum and Tokaloglu and
Karacabey apricot cultivars. The highest bud break and fruit set ratios were
87.67 and 85.33 per cent in Papaz plum treated with 2 per cent hydrogen
cyanamide, respectively. Promalin applications also caused better flower and
vegetative bud break and fruit set. Chemically treated trees could be harvested
2-9 days earlier than control trees. All the chemical applications increased
fruit weight in both plum and apricot (Son and Kuden, 2005). Spraying with
PCPA (4-CPA) at 30 x 10-6 M at the stage of 75 per cent flower fall increased
the fruit set. Application of mixture of 30x10-6 M PCPA + 2000 x 10-6 M
borax + 2000 x 10-6 M potassium dihydrogen phosphate also produced good
effect (Wu et al; 1993).
3. Water stress : Growth of shoots, flowers and fruits in deciduous trees appears
at different times and therefore the timing of regulated deficit irrigation can
influence its effects. The stage II of fruit growth has been identified, as an
appropriate period at which fruit growth is minimal and therefore not affected
by drought, while shoot growth can be reduced in stone fruits (Li et al;
1989). In early cultivars, this period is very short, the main shoot growth
occurs after harvest, and thus may be ideal time for droughting (Johnson and
Handley, 2000).
Water deficit studies in Black Gold Japanese plum were investigated by
Intrigliolo and Castel (2005) during phenological stages II and III of fruit
growth, or after harvest, replacing 33 or 66 per cent of tree evapo-transpiration
(ET), or during both periods at 66 per cent of ET. They reported that, water
deficit during fruit growth reduced average fruit weight, with a negative
correlation between fruit size and integrated mid day stem water potential.
In contrast, drought after harvest did not affect flowering, fruit set, fruit
growth or yield, in the short term, but drought may reduce productivity, in
the long term, as a consequence of the cumulative effects of water deficit on
tree growth.
 Fruit Growth and Development 139

4. Nutrients : The tree performance (branch elongation, flowering and fruit

setting intensities) and fruit size has been found to be increased by titanium
sprays in combinations with other commercial compounds containing Ca2+
and Mg2+. This improvement due to calcium absorption as a consequence of
the beneficial effect of titanium is on the absorption, translocation and
assimilation processes (Alcaraz et al; 2003).

Strawberry
Maximum fruits in strawberry develop from tertiary flower than from primary
and secondary flowers.
Thermoperiodic influences : Plant growth, development and composition of
fruits are affected by genetic factors. However, environmental factors such as
water availability, day and night temperature and day light intensity also influence
the fruit production. The effects of temperature as well as its interaction with
other environmental factors, like photoperiod often vary with cultivars and species
e.g exposure to high temperature (35°C) results in reduced plant growth and
lower yield (Hellman and Trvis, 1988). The influences of day/ night temperature
combination on plant growth and fruit quality was studied by Shiow et al. (2000)
in cv. Earliglow and Kent of strawberry. The optimum day/ night temperature for
fruits was found to be 18/ 12°C. The fruit surface and flesh colour become darker
(L-value decreased) and greater pigment intensity (chrome value increased) as the
day/ night temperature increased. Darkest, reddest and with greatest fruit surface
and flesh pigment intensity strawberries were obtained at 32/ 22°C day/ night
temperature. Two anthocyanidin glycosides i.e pelargonidan-3 glucoside and
cyanidine-3- glucoside are almost exclusively responsible for red colour of
strawberries (Timberlake and Bride, 1982). High temperature (25-30°C) increased
fruit soluble solids, titratable acidity and ascorbic acid content as well as soluble
solids/ titratable acid ratio. Plants grown at 18/ 12°C had high ascorbic acid
content, probably due to decreased metabolism at lower temperature. High
irradiative conditions favour ascorbic acid content, while wet, cloudy weather
results in low values (Hansen and Waldo, 1944). Low content of organic acids
(mainly citric acid) in strawberry grown at high temperatures may result from
increased respiration. The total organic acid level is positively correlated with
titratable acidity. Temperature also greatly affected soluble carbohydrates viz.
fructose, glucose, myoinositol and sucrose which are major sugars, comprising
>65 per cent of the total soluble solid content in strawberry fruit.
Magnetic field effects : Studies have shown that MF had a positive effect on
flower number, yield and earliness (Danilov et al; 1994). The plants of short day
strawberry Camarosa treated with magnetic flux density field recorded increased
140 Fruit Tree Physiology

fruit yield per plant at low strengths than control and with high MF. Increasing
MF strength upto 0.096 Tesla increased fruit yield per plant but higher than this
MF value reduced fruit yield. All the MF strength increased average fruit weight
compared to control and the largest fruits (8.92) were obtained at 0.096 Tesla.
These effect can be explained by the effect of MF on plant metabolism such as
photosynthesis, hormones and enzymes and changes in endogenous solute specially
carbohydrates, growth regulators and enzymes. Plant nutrient uptake from soil is
also affected by MF. The application of low MF strength may be of benefit for
fruit yield and plant growth for greenhouse strawberry production.

Loquat
Loquat trees flower profusely in compressed panicles, and although fruit set
percentage is very low, fruit size is commonly very small. In fruit crops, fruit size
is inversely related with number of fruits produced per tree. Increase in average
fruit size can be obtained by thinning the fruit manually or chemically or by
applying synthetic auxins at the onset of cell enlargement (Agusti et al; 2005).
The competition for carbohydrates among surviving fruits is minimized by manual
thinning and also increases final fruit size and the percentage of purple spotted
fruits (Gariglio et al; 2003). Enhanced flesh cell expansion, increasing capacity
of fruits to grow is achieved by chemical thinning (Agusti et al; 2005).
Ringing (removing 1-20 mm strip of bark around a limb) obstruct phloem
transport and improve the availability of metabolites for developing fruit above
the ring. It damages the tree and branches may take several months to heal. Scoring
is a method in which a simple cut (about 1 mm wide) is made on each branch
around its circumference, without removing bark and without affecting xylem
tissue. The scoring causes phloem discontinuity but heals quickly (takes 2 weeks).
Branch scoring after fruit set increases fruit size, grows faster and reaches
commercial colour and flesh softness earlier than fruit from no scored trees. Scoring
is effective as ringing at improving fruit development (Agusti et al; 2005).

CONCLUSION
1. Most fruits follow single sigmoid, some double while some others triple
sigmoid (Kiwi) growth curve.
2. Cell division, cell enlargement and maturation are important steps of fruit
growth and development.
3. Various internal factors like hormones, cell size, cell number, seeds,
photoassimilates etc. and external factors including temperature, light,
chemicals, cultural practices etc. influence fruit growth and development.
 Fruit Growth and Development 141

4. Groth regulators play an impottant role in development of fruits.

5. Various physico-chemical changes like increase in size, weight, volume,
change in colour, TSS, acidity, reducing sugars etc. occur during fruit growth
and development.

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 6

UNFRUITFULNESS IN FRUIT CROPS –

CAUSES AND CONTROL MEASURES

U nfruitfulness is a major problem in many fruit crops and their varieties

which results a huge loss to growers and making fruit cultivation a less
profitable. Fruitfulness refers to the state of plant when it is not only capable of
flowering and fruit setting but also takes these fruits to maturity. The inability to
do so is known as unfruitfulness or barrenness. Thus, a fruit plant, which is not
productive, is unfruitful. In spite of adequate flowering, low fruit yields in orchards
have been experienced because of low initial fruit set and subsequently higher
fruitlet abscission. In an orchard, all the fruit trees do not bear equally or regularly
and sometimes fail to flower and fruit under similar conditions where another
fruit tree bears heavily. This failure to fruit may be attributed to unfruitfulness.
Thus, unfruitfulness is one of the serious problems of fruit crops and its
causes need to be understood properly for effective control and obtaining
economically acceptable production level.

CAUSES OF UNFRUITFULNESS
Unfruitfulness can be due to lack of balance between vegetative growth and
fruiting, lack of flowering and poor fruit set as the result of unfavourable
environment. It can also be due to heavy cropping, leading to inhibition of fruit
bud production and poor crop in the following year. The season of flowering and
fruiting can also be modified to adapt crops to local environments for better
performance.
Sterility also leads to unfruitfulness due to impotence, incompatibility or the
abortion of embryo. The causes of unfruitfulness can be broadly grouped into two
categories:
148 Fruit Tree Physiology

1. External factors
2. Internal factors
(A) Internal factors (B) External factors

I. Evolutionary II. Genetic III. Physiological 1. Temperature.

tendencies factors factors 2. Pruning and
1. Imperfect 8. Sterlity or 10. Slow growth of grafting
flowers unfruitfulness pollen tube 3. Season
2 Structural due to hybridity 11. Premature or 4. Age and vigour
peculiarities e.g 9. Incompatibility delayed of plant
heterostyly pollination
(i) Interfertility 5. Nutrient supply
3. Dichogamy 12. Nutritive
(ii) Reciprocal 6. Locality
(i) Protogyny conditions
crosses 7. Light
(ii) Protoandry within the plant
4. Stigmatic (i) Pollen 8. Disturbed water
receptivity sensitivity relations
5. Abortion of (ii) Fruit setting 9. Rains at bloom
pistils or ovules and their 10. Pests and
6. Non viable position diseases
pollen/ 13. Poor pollen 11. Wind
Impotency of germination 12. Spraying
pollen 14. Position of 13. Root stock
7. Sterlity flower

External Factors
The various external factors affecting fruitfulness are as:
1. Temperature : Among the environmental factors, temperature has a great
importance. It affects flowering and fruit set in several ways. It is common
knowledge that a period of cool yet frostless weather is conducive to better
blossoming, fertilization and fruit set. The abscission of flower bud, fruit,
etc. is a function of temperature. Temperature is quite important for pollen
germination in plum, cherry, apple, pear, etc. where 4°C or lower completely
checks the same.
The pollination and fertilization of almonds are negatively affected by
the low temperatures. The low temperature during blooming period can prevent
the germination of pollen on stigma to prevent the development of pollen
tubes in the style. Cuevas et al. (1994) has reported total inhibition of fruit
set in olive at 30°C and 25°C. At 20°C pollen tube growth was slow resulting
in delayed or reduced fertilization.
 Unfruitfulness in Fruit Crops – Causes and Control Measures 149

Temperature affects the rate of pollen tube growth of Bartlett pear taking
12 days at 5°C and 2 days at 15°C.
1. Winter thaws followed by cold weather can kill flower buds. Frost during
blossoming can damage flowers, and cool temperature can decrease the
viability of pollen.
2. Lower temperature exerts an important effect on the fertilization
mechanisms of Tokaloglu apricots by stimulating the growth rate of
pollen tube even in the cross pollinated combinations (Guldau and Askin,
1991).
3. Excess heat can greatly reduce avocado fruit set. Hot, drying winds
during the blooming period can desiccate the tender stigmas or styles,
sharp environmental variations reduce pollen viability. Finally heat can
reduce fruitfulness by injuring the tree (Bergh, 1976). Flower and fruitlet
drop in Satsuma mandarin exceeds in May-June when temperature was
35-37°C.
4. At too higher temperature (40°C) or too low temperature, the bee activity
is hindered and thereby low fruit set in cross pollinated crops.
5. Sex reversal takes place in papaya at low temperature i.e male plants
bear fruits sometimes in cool climate.
6. Ovule senescence become faster in Italian than in Brooks cultivar of
plum with increase in temperature. Only one ovule per flower remains
viable by 8 DAFB for Italian, whereas for ‘Brooks’ cultivar at 15oC,
higher temperature decreases ovule longevity (Moreno et al; 1992).
7. In mango higher temperature and dry weather is associated with increased
production of pollen per anther leading to better fruitset. Panicles emerging
early in the season (December-January) in north India are usually
unproductive and become malformed i.e. low temperature favours
unfruitfulness.
8. A rise in diurinal air temperature under green house reduced the production
of pollen grains in peach cv. Granda ( Nava et al; 2009)
Flower bud formation is negatively correlated with temperature in apple.
Increase in temperature results increase in sucrose, glucose, fructose but
decrease in malic acid.
2. Humidity : Low atmospheric humidity in avocado causes drying of stigmatic
secretions so that pollen does not germinate. Rain is an important factor that
indirectly affects fruit setting by disturbing the process of pollination,
germination of pollen grain and even staminal fertilization. The poor
germination of pollen in almonds is attributed to damp weather during fruit
150 Fruit Tree Physiology

set (Klugness et al; 1983). Wet and humid weather favors anthracnose and
poor fruit set in mango. Relative humidity on its own is an important factor
in bee activity. Low temperature and high humidity have the double effect
of reducing bee activity and slowing the release of pollen (Goodman, 1994).
3. Light : Poorly shaded plants hardly differentiate into flower buds. There is
poor fruitset, yield and colour development in overlapping tree canopies.
Light effects fruitfulness indirectly by its effect on photosynthesis. Light is
pre-requisite for photosynthesis and low light intensity or its duration reduces
the carbohydrates reserves in the trees. In addition to this poor light conditions
promote fruit abscission. Nuzzo et al. (1999) observed that in Prunus
armeniaca cv. Tirynthos the light had positive effect on flower bud number
per fruiting shoot and fruit set and growth rate were lower in the shaded zone
of the canopy. Pollinator activity in peach is positively correlated with the
light intensity (Choi, 1987) which affects the fruit set.
4. Wind : Wind controls pollination either by promoting wind pollination or by
checking insect pollination. It is desirable in wind pollinated fruit trees like
hazelnut and walnut but if its speed is too high, it is harmful because it
results in low fruit set on exposed sides and heavy crop on other sides.
Excessively speedy winds cause ovary abortion (Mastrofillipe, 1966) and
also make the stigma dry. Wind pollinated crops include: walnut, pecanut,
coconut, hazelnut etc. However, most of the fruit crops are insect pollinated,
the wind hinders pollination in them.
5. Rain at bloom : Rains at the time of blooming period cause unfruitfulness
by washing away pollen grains, inhibit pollinators and cause spread of diseases
and insect and pest. Every fruit species need a specific time period without
rains at the time of blooming for successful pollination.
6. Tree age and vigour : This factor is in majority of cases linked intimately
to the effect of nutritive status of the plant or its limb, locality and the
season, having direct effect on flowering and fruit set, and thereby
unfruitfulness. Excessive vegetative growth reduces fruitfulness of trees, on
the other hand, optimum vegetative growth to support the crop load is very
much essential, otherwise leads to fruit drop either at early or later stages e.g
as in strawberry, walnut, etc.
7. Soil moisture status : Soil moisture status at the time of flowering and fruit
set is also an important consideration for the fruitfulness of tree. Pollen
abortion is induced by moisture stress in pecan, whereas drought or stress
just before harvest results in reduced fruit set (Polla et al; 1993). Thakur
et al. (1993) observed higher soil moisture and fruit set and yield following
mulching with black polythene in apple. Chandel and Singh (1992) observed
 Unfruitfulness in Fruit Crops – Causes and Control Measures 151

higher fruit set and yield of mango when irrigation was applied at 20 or 40
per cent soil moisture deficit. In general, there is low fruit set if the moisture
during flowering and fruit set is either excessively high or there is water
stress. Water stress in fruit plants is associated with the decrease in nucleic
acid and protein contents of the fruit and leads to formation of abscission
layer (Goncharova et al; 1985).
8. Nutritional status of the soil : Higher nitrogen levels have also been reported
to increase fruit set of Anjou and Bosc pears, while moderate levels of N has
resulted in maximum fruit set of Bartlett pears (Westwood and Bjornstad,
1974). Sufficient amount of phosphorus during fruit setting increase the fruit
set and yield in apple, but excess of N before flowering promote another
vegetative flush and reduce fruit set (Okomoto et al; 1982). Awasthi and
Singh (1990) reported that fertilizer application at appropriate stages
significantly increases fruit set, yield and quality. The potassium content less
than 0.8 per cent was associated with excessive fruit drop in Valencia and
Pineapple oranges.
9. Grafting : Grafting (Union of root stock and scion wood) is most important
factor for producing healthy, productive and cross compatible cultivars. If
wrong types of cultivars are grafted with each other it causes unfruitfulness.
It is mostly happened when grafting is done without knowing their
compatibility status.
It is better to graft onto dwarfing and semi dwarfing root stocks than on
vigorous root stock, and better or spur type cultivars than standard types in
Red Declicious apple (Hull, 1978). Fruit set was higher in Starking Delicious
apple grafted on M-9 root stock (Tareen et al; 2003).
10. Pruning : Pruning modifies hormonal levels thereby affecting fruitfulness.
The influence of pruning varies with amount, season and kind of pruning.
Too early or too late pruning in deciduous cultivars also cause unfruitfulness.
Grochowska et al. (1977) reported that hormonal levels are modified by
pruning. The pruning increases fruit set as a direct result of growth hormones.
Incorrect pruning can change the natural habit of tree and can stimulate
excessive growth. Both will delay flower and fruit production. The most
characteristic symptom of apricot bare twig and unfruitfulness besides the
lack of fruit is the change in the habit. In grapevine, it has been observed that
cane-pruned vines produce berries and fruits better than the severe spur-
pruned grapes. Pear requires heavy pruning for heavy fruit-set. A better fruit
setting of apples on trees pruned moderately or heavily has been observed
than on trees pruned lightly or left unpruned.
152 Fruit Tree Physiology

11. Locality : Fruit setting on trees of the same variety is often much better in one
locality than in another. It is known fact that various factors, like soil,
temperature, humidity, light etc, that constitute the locality influence fruit
setting in various fruit crops. It is only the environmental complex, which
influences the fruit - set in a locality. Some examples of the effect of locality
on fruit set are as under:
The Jonathan apple is almost sterile in Victoria (Australia), although it is
fertile in USA. The Bosc pear is self fertile in many localities, but becomes
partially fertile in New York (USA) and develops parthenocarpic fruits
regularly in South Africa. The commercial mango varieties of south India
(e.g Neelum and Baneshan) usually don’t perform better under northern
Indian conditions.
12. Girdling : Improved fruit retention by girdling was found associated with
higher GA level and lower ABA level as well as with the higher level of
carbohydrates in the fruit on gridled trees. An experiment with labeled 14C
showed that down stream translocation of carbohydrates was almost or
completely blocked by spiral or closed girdling respectively causing the
inhibition of root growth in litchi cvs. Nuomice and Guiwei (Huang, 2002).
13. End- season fertility : End season fertility of normally self-sterile plants is
rather common in some fruit plants. Some grape varieties (e.g., Ideal) proved
to be self-sterile early in the season, but self-sterile late in the season. In
mango, the early flushes of flowering (December – January) are rather
unproductive than the late flushing (March) because of defective pollen and
higher proportion of male flowers in the early flushes. Indeed, end-season
sterility is quite expected in plants producing indeterminate type of
inflorescence, which is very common in strawberry. A striking example of
seasonal influence on fruit setting and unfruitfulness occurs in figs of San
Padro class. In this class, the early crop (breba) set fruits freely without
pollination and produce seedless fruits. However, the summer (late) crop
won’t set freely without caprification (pollination by wasp). This example is
similar to strawberry as it is an instance of early season fertility rather than
late season fertility which is the characteristic feature of almost all the varieties
of strawberry and San Padro figs.
14. Chemicals and pesticides : The use of pesticides can kill bees therefore
reducing pollination. Some pesticides can also be toxic to delicate flowers
causing abortion and loss of fruits. Stigmatic surface may be injured due to
pesticide sprays and can result in an inhibition of pollen germination and
tube growth (Wetzstein, 1990). Reduced pollen viability, impaired pollen
release and death of pollen has been observed in apple due to spraying of
 Unfruitfulness in Fruit Crops – Causes and Control Measures 153

fungicides like fungicides containing captan, dinocarp, sulphur and triforine

on dehisced anthers (Ruth et al; 1978).
15. Pests and diseases : Sap sucking insects such as aphids, scale and mites, plus
the larger insects, caterpillars and grasshoppers all cause major damage to
flowers and fruits.
Diseases can be particularly deliberating and in some cases will kill trees
quickly. Prolonged wet weather can encourage diseases such as anthrocanose,
phytophthora, black spot, mildew and sooty mould, all of which adversely
affect fruit set or maturation. Among diseases, fire blight is recognized as
one of the most serious diseases, which limits fruit setting in pears
considerably. In apple and pear, scab is responsible for maximum flower and
fruit drop, brown rot attacks the blossoms of almost all the stone fruits,
powdery mildew on developing fruits of mango, ber, downy and powdery
mildew on grape berries, causing excessive drop of affected flowers and
developing fruits after fruit setting.
The losses caused by the attack of insect pests and diseases vary greatly
with the locality, variety and seasonal variation. For instance, mango
malformation causes severe losses under north Indian conditions, but mango
orchards in south India are virtually free from this malady. Downy mildew
causes serious loss to grape berries in south, but rarely in north India.
16. Root stocks : Root stocks are known as backbone of fruit industry. Quince
is used to induce precocity and fruitfulness in pear. Malling series (M9, M27
etc) is used to induce precosity in apple.
17. Interstocks : Interstosk pieces of varied lengths have been found to induce
precosity in apple e.g interstem pieces of clonal root stocks (M9, MM106 and
MM111) in Ambri apple have reduced the gestation period and induced early
flowering.

Internal Factors
There are a number of internal factors which are associated with unfruitfulness.
They are further categorized into 3 major categories:
1. Evolutionary tendencies
2. Genetic factors
3. Physiological factors
1. Evolutionary tendencies : Due to evolutionary tendencies, the cross
fertilization is must for maintaining the vigour of the species. In these species
self fertilization is difficult or impossible. These factors helpful in maintenance
154 Fruit Tree Physiology

of these species, may, in cultivation, limit its usefulness and range. The
evolutionary factors leading to unfruitfulness are:-
Imperfect/ defective flowers : A perfect flower possesses both male (stamens)
and female (pistil) parts, whereas an imperfect flower may be either staminate
(functionally male, having only stamens) or pistillate (functionally female,
having only a pistil or pistils). The style is very short or lacking in some
pistillate flowers, like in pecan, but the stigma and ovary are always present
if the pistil is functional. If staminate and pistillate flowers are borne on the
same plant but in different locations, the species is termed monoecious. If
staminate and pistillate flowers occur only on different plants, the species is
termed dioecious. One can see the ramifications for pollination and orchard
design. A dioecious species such as pistachio or kiwi must be planted in an
orchard with male plants near females for pollination. Aside from pollination,
the male plants are useless since they do not possess ovaries that will ripen
into fruit.
In case of cultivars of oranges, grapefruit and tangerines the pistillate
flowers give rise to fruits parthenocarpically. Strawberry produces both
pistillate and perfect flowers. Walnut, pecan, chestnut, banana and litchi are
monoecious. In monoecious fruit plants in general, there is no or very little
problem of pollination, fruit setting and fruitfulness. A number of sex forms
have been reported in papaya. The evolution of unisexual flowers cuts down
chances of self fertilization completely. Of the unisexual plants the pure
staminate flowering ones are essentially barren. The sex distribution in papaya
is very unusual and even variable between different flowers of a plant. Earlier,
about 13 classes of flower types are recognised in papaya, depending on the
combination or separation of stamens and pistils in the flower clusters, corolla
and fruit. Presently only 8 classes are recognized and are as follows:
1. Pistillate flowering plants.
2. Staminate flowering plants.
3. Plants producing both staminate and perfect flowers.
4. Plants producing both staminate and perfect flowers, but with sterile
pollen, more often called as pseudo - hermaphrodite plants.
5. Plants producing staminate and perfect flowers, but neither pollen nor
pistils are fertile; more often called as sterile hermaphrodite plants.
6. Plants producing staminate, pistillate, and perfect flowers.
7. Plants producing staminate and perfect flowers.
8. Plants producing staminate and pistillate flowers.
 Unfruitfulness in Fruit Crops – Causes and Control Measures 155

Out of above 8 classes, types 2 and 5 are necessarily unfruitful. Type 5

is fruitful apparently because of incompatibility but is non-functional. Type
1 and 4 are self-unfruitful and the other types are self fruitful. In general,
type 2 plants are more common to occur and consequently, it is customary
to retain at least 10 per cent staminate or male plants in an orchard so as to
have good fruitset because they provide pollen to pistillate flowers and thus
are equally important.
Structural peculiarities : The presence of short styles with long filaments
(Thrum) e.g sapota and pomegranate or long style and short filaments (Pin
type) e.g almond and carambola is dimorphism, a type of heterostyly (Salaria,
1999). In carambola with thrum selfing, the pollen tubes are arrested at
stigma style interface while in pin type selfing; the pollen tube may penetrate
longer (Wong et al; 1994). Arora (1993) observed heterostyly in sapota and
cross pollination gave higher fruit set in cvs. Kalipatti, Cricket Ball and
Calcutta Round relative to self pollination. Singh et al. (2006) observed three
types of flowers, namely thrum, homostylous and pin in pomegranate cvs.
Ganesh-1 and Kandhari. In some apricot cultivars the stigma is located above
the superior plane of the anthers (greater than 3 mm) and as such, prevents
self pollination (Ruiz and Egea, 2008). However, heterostyly is a relatively
less important factor in dioecious plants. In tri-stylic conditions, flowers with
long, medium and short styles are produced which leads to unfruitfulness
due to different level of stigma and the style. Heterostyly is relatively less
important factor in dioecious plants.

Fig. : Tristyly flowers a) with long style b) with medium style c) with short style
(Singh, 2008).

Dichogamy : When stigmatic receptivity period does not coincide with pollen
viability in monoecious plant is known as dichogamy. In dichogamy, self
pollination is prevented in perfect flowered plant, due to maturity of two sex
156 Fruit Tree Physiology

organs at different times. If the stamens ripe before the stigmas become
receptive the flowers are known as protoandrous and if stigmas become
receptive before the stamens produce viable pollens it is known as
protogynous. Dichogamy has been reported in hermaphrodite (avocado,
mango, ber, annona), monoecious (cashew, pecan, walnut, chestnut) and
dioecious (persimmon, fig, pistachio) species. In walnut occurrence of both
protoandrous and protogynous cultivars prevents self pollination which
warrants cross pollination. Kumar et al. (2005) suggested that walnut
plantations should include either homogenous cultivars or both protoandrous
and protogynous to achieve adequate levels of pollination and fruit set. In
pecan, the dichogamy may be complete or incomplete and the special type
may be protoandry or protogyny. Certain varieties like Moore, Alley, Texas,
Prolific and San Saba have proved to be reliable in the production of pollen
at an early date.
Stigma Inferior Ovary
Style Stigma
Petal Petal
Anther Anther Rudimentary
Style
Stamen

Filament

Sepal Sepal
Ovary Ovary
Hypanthium Hypanthium
Ovules Ovules
Peduncle or pedicel Peduncle or
pedicel

Petal Anther Stigma

Petal Rudimentary
Stamen StyleAnther
Filament
Sepal
Sepal
Peduncle or pedicel Ovary Hypanthium
Ovules Peduncle or
Staminate Flower
pedicel
Fig. : Different forms of flowers (http//www. flowerforms.edu.in)

Spencer and Kennard (1956) reported that in mango, the duration of

stigma receptivity is only 2-4 hours, following anthesis since pollen shedding
does not start within few hours of anthesis, the periods of maximum male
and female fertility in bisexual flowers fail to coincide.
 Unfruitfulness in Fruit Crops – Causes and Control Measures 157

(i) Duodichogamy : A condition when two batches of male flowers are

temporarily separated by a batch of female flower in between. The resting
period in between avoids selfing compeletly. It is an adaptive mating
system in plants which results from competition of males to access small
number of females. The plants flower in the order male → female →
male. Examples: chestnut.
(ii) Heterodichogamy : It is a state of having two genetically reciprocal
morphs which function as female first, then as male, the other behaving
in opposite ways. Examples: walnut, hazelnut, pistachio nut.
(iii) Protogynous diuranally synchronous dichogamy (PDSD) : In this, the
flowering synchronises in such a way that each flower is female at one
time and male at another time. Diuranal synchronization is observed on
the tree and flowers behave like functional female during one part of the
day and as functional male during another part of the day. The dichogamy
is protogynous in nature and pistil matures prior to androcium. Thus
flower appears structurally bisexual but acts like unisexual. When cross
pollination does not occur, the plants remain unfruitful. Example: avacado.
The inflorescence of avocado technically has been termed as Protogynous
Diurnally Snychronous Dichogamy (PDSD) by Bergh (1969). Dichogamy
indicates that female and male parts mature at different times but
synchronization is such that the flowers, which open on a tree are female at
one time and male at the other time. The pistils mature earlier than the
stamens i.e it is protogynous. Besides the synchronization of flower opening
is diurnal because the flower is functionally male at one part of the day and
female during another part of the day. Thus, the flower is structurally bisexual,
but functionally unisexual. Each flower opens twice and closed in between
and functions first time as female and second time as male. The opening of
the flowers is so synchronized that self pollination is prevented even though
the flowers are perfect and pistils are receptive and pollens being shed
everyday through-out the flowering season. The fruit production becomes
possible only when adequate provision is made for cross pollination. Chandler
(1985) classified avocado varieties into two groups based on its unique
flowering behavior. In group A flower first open in the morning and the
second opening during the afternoon of the following day i.e the time gap
between two openings is more than 24 hours. In group B, flowers first open
in the afternoon and then again in the next day morning. Thus every morning,
A pistils can be fertilized with B pollen, while during afternoon, B pistils are
ready to receive A pollen.
The stigma is receptive only for a few (2-4 hrs) hours after anthesis,
whereas, pollen is shed for a longer period in case of mango. Since pollen
158 Fruit Tree Physiology

shedding does not start within few hours of anthesis, the periods of maximum
male and female fertility in bisexual flowers fail to coincide (Verma and
Jindal, 1997). About 50 to 2000 staminate flowers in coconut inflorescence
open each day continuously for 15 to 20 days and this period is followed by
5 to 16 days in which pistils become receptive.
Stigmatic receptivity : It is the ability of the stigma to support pollen
germination and it limits the effective pollination period (EPP). EPP is the
number of days during which the pollination is effective in producing a fruit
and is determined by the longevity of ovules minus the time lag between
pollination and fertilization. In kiwi fruits reduced stigmatic receptivity is due
to degeneration of stigma and rupture of papillar integrity (Sanzol and Herrero,
2001). Stigmatic receptivity is a limiting factor for flower receptivity in Agua
de Aranjuez pear. Maximum fruit set was observed when flowers were
pollinated at anthesis and 2 days after anthesis than 4 to 6 days after anthesis
which indicates that stigmatic receptivity is an important factor limiting pear
flower receptivity and thereby fruit production (Sanzol et al; 2003). About
80, 36 per cent and no (0 %) fruit set was obtained in kiwi when pollination
was done during the first 4, 5 and 7 days following anthesis respectively.
Thus, the EPP was limited to the first 4 days after anthesis and the stigmatic
receptive averaged 84 per cent and was nil after 7 days (Gonazel et al; 1995).
Abortive flowers or aborted pistils or ovules : Interference either in the
development of the flower or in the full development of sex elements and
their function may lead to unfruitfulness. Floral abortion is more common in
indeterminate inflorescence as compared to determinate inflorescence. Pistil
degeneration leads to unfruitfulness in certain cvs. of plum and ornamental
pomegranate while in strawberry pistil abortion is late so unfruitfulness does
not take place. Certain olive varieties have 10-60 per cent abortive embryos.
Rovira et al. (2001) observed the highest average value of pistillate
flower abortion in Spanish selections and lowest in French cvs. and Chilean
selections of walnut. California cvs presented an intermediate average pistillate
flower abortion value. In most of the Prunus spp., 2 ovules differentiate and
develop in each carpel. One of these, the secondary ovule, usually aborts
sometime after pollination leaving only the primary ovule to be fertilized. In
almond, in some cases, both ovules may be viable and double kernels are
produces and in some of the almond selections, the abortion of both primary
and secondary ovules has been reported. Pimienta and Polito (1982) concluded
that ovule abortion is accompanied by blockage in metabolite supply.
The proportion of aborted pollen grain, varied from 22.5-46.8 per cent in
cashew nut showing a steady increase with plant age, reflecting an increase
 Unfruitfulness in Fruit Crops – Causes and Control Measures 159

in genetic load with plant age (Bhattacharya, 2005). Most cvs. of Chinese
jujube (Zizyphus jujube Mill.) have very serious embryo abortion and
recommended embryo rescue to get hybrids (Liu and Qi, 2004). In self and
cross pollination study of plum cv. Nonpareil with pollinizer Marcona revealed
that though the pollen tubes in both the cases reached the ovule almost at the
same time but the selfed ovules showed abnormalities as some failure in the
polar nucleus unit and the lack of the elongation of embryo sac that could
induce ovule degeneration (Oukabli et al; 2000).

Table : Various causes of flower abortion in different fruit crops (Gardner, 1952).

S.No Fruit plant Causes of abortion

1 Apple Defective embryo, defective ovules

2 Almond Defective embryo sac, gynoecium abnormality
3 Grapes Degeneration of nucellus
4 Kiwi fruits Pollen degeneration
5 Strawberry Lower bud abortion, defective pistil
6 Sour cherry Defective embryo
7 Mandarin Abnormal pistil
8 Pecan nut Defective pistil
9 Plum Degeneration of pistil
10 Peach Degeneration of nucellus, embryo abortion
11 Olive Pistil abortion
12 Litchi Embryo abortion
160 Fruit Tree Physiology

Non viable pollen : It is due to non functional pollen or the ovule. Non
viability or impotence of pollen results in unfruitfulness. Unfruitfulness in
case of Muscadine grape is due to defective pollen. In India, various studies
on loquat pollination indicate that early flowers in number of varieties show
abnormal stamens possessing very low viable pollen. Azarenko et al. (2006)
reported that in some cvs. of hazelnut produce a high percentage of defective
pollens, because of high percentage of non viable pollen. These cultivars are
not recommended as a pollinizer. Ruiz and Egea (2008) observed that late
flowering apricot genotypes showed lower pollen viability than early flowering
genotypes and is one of the important factors attributing to lower production
of these cultivars.
Differences in the viability of the pollen and in the morphology of the
peach flowers in Forastero were observed by Radice et al. (2004) when this
cv. was grafted onto different root stocks. FJM (Ferdor-Julior), Cuaresmillo,
Mr. Sim, Mr. SM root stocks induced the greatest viability of pollen grains
on Forastero flower by 35, 34, 34 and 32 per cent, respectively.
Sterility : Some species have genes that prevent development of the pollen
or the ovule. Generally, sterility is due to failure to obtain normal development
of pollen, embryosac, embryo, and endosperm. Morphological sterility is due
to suppression of rudimentry pistil or abortion of sex organs. Pollen sterility,
called male sterility is more common when one of the sexes is inactivated,
then cross pollination has to take place. Pollen sterility is common in peach
cv. J.H. Hale and also many olive cultivars. Verma and Jindal (1997) reported
22 and 98 per cent proportion of sterile ovules in apricot and avocado,
respectivel. A high percentage of ovules in Swan Hill olive cultivar contain
poorly developed embryo sacs at anthesis and were not fertilized (Rallo,
1981).
In Z. mauritiana, male sterility has been observed. These male sterile
lines will produce good fruit if pollination occurs. Unlike male sterile cvs.
a female sterile cultivar will not set fruits even in the presence of pollinizers
unless it has tendency to develop parthenocarpically. However, total ovule
degeneration has been observed in mango (Sturrock, 1969) and almond
(Pimienta and Polito, 1983). In Moorpark and Trevatt Knit cvs. of apricot,
ovules contained all reproductive structures while multiple ovules which
were small and related in development were present in the Trevatt Blue
flowers and anthers contained degenerated microspores. This is the first
report of a simultaneous mutation in both female and male function in apricot
(Lillecrapp et al; 1999).
 Unfruitfulness in Fruit Crops – Causes and Control Measures 161

2. Genetic factors :
Unfruitfulness due to sterile hybrids : Hybridity is associated with sterility
as well as unfruitfulness. The degree of sterility increases with wider crossing.
Peach plum hybrids known as Blackman or Mule have complete sterile and
barren flowers as also in Kamdesa- a hybrid between peach and sour cherry.
Triploid apple and some pear varieties are sensitive to pollen ovule or embryo
sac abortion e.g apple cv. Baldwin and Gravenstein (Watenable et al; 1994).
The popular tangelo is a hybrid produced by crossing a grapefruit (C. paradisi)
with a tangerine (C. reticulata). They are seedless or produce seeds with
only nucellar embryos (Gardener, 1952).
Table : Some other sterile hybrids (Gardner (1952) :

S. No. Cross Hybrids

1 Troth early peach x Wild goose plum Mule

2 Peach X Sour cherry Kamdeso
3 Pear X Quince Pyronia
4 Persian walnut X Royal California walnut Royal
5 Rersian walnut X Gastern black walnut Paradox
6 Sweet orange X Trifoliate orange Citrange

Incompatibility : Incompatibility is defined as failure of viable pollen to

grow down the style of flower of the same variety (self incompatibility) or
of the different varieties (cross incompatibility). Many cross pollinating species
exhibit self incompatibility, such that fertilization by their own pollen is
disfavored or prevented through physical or biochemical factors. There are
different degrees of self-incompatibility, and many self incompatible species
will produce a few fruit even when self pollinated. Thus, a single apple tree
may have a bushel or so of fruit, since apples are not completely self-
incompatible, but the same tree may produce several bushels if cross-
pollinated. Horticulturists have coined the terms self-fruitful and self unfruitful
to describe cultivars that can set commercial crops, or cannot set commercial
crops (respectively) when self-pollinated. Thus, self fruitful and self-unfruitful
are economic or horticultural terms, whereas self-incompatible or cross-
incompatible are botanical terms.
Self incompatibility is more common in fruit crops like apple, pear, sweet
cherry, almond, avocado, fig, mango, citrus, olive etc. than cross incompatibility
(apple, pear, sweet cheery, European plums, and almond). The sexual
incompatibility is a genetic mechanism to ensure out crossing of plants, thus
bring together germ cells of potentially greater genetic diversity. Incompatibility
162 Fruit Tree Physiology

is genetically controlled character manifested by the presence of multiple

alleles at a single locus (Sedgly and Griffin 1989). In fruit production
incompatibility may create a platform to create variation leaving no scope for
inbreeding. In mango, self unfruitfulness is reported in cvs. Dashehari, Chausa
(Ram et al; 1976), Langra and Fajri Kala (Afifi et al; 2000). While evaluating
superior genotype of almond, Dhillon et al. (1984) found all of them self-
incompatible. Fruit set following open pollination ranged from 12 per cent in
JKS-168 to 28.2 per cent in JKS-55 while from cross-pollination, it ranged
from 2.6 in JKS1-68 ´ JSK-65 to 51.9 per cent in California Papershell ´ JKS-
65. Fruit set percentage was higher in cross combinations as compared to self
pollinations (Eti et al; 1994). The unfruitfulness in pears, apples, cherries,
plums, dewberries, blackberries, almonds, blueberry and most of the stone
fruits is also due to incompatibility. Most pear (Pyrus communis L.) show
gametophytic self incompatibility system which prevents the self fertilization
and thereby fruit set (Sanzol, 2007). Incompatibility was reported in loquat
varieties like Improved Golden Yellow, Pale Yellow and Golden Yellow by
Singh and Rajput (1964) who observed that the pollen tube penetrated the
stylar canal up to 1/4 to 1/3rd of its length and did not go further below, even
after 72 h of pollination. Sanzol (2007) reported that about 80 per cent of the
pollen tube reaches the base of the style and fertilized 70 per cent ovules after
cross pollination, whereas only 10 percent pollen tube reaches the ovule and
fertilized only 5 per cent of the ovule after self pollination in pear cv. Agua de
Aranjuez, indicating a high degree of incompatibility in pear. Stone fruits
have high degree of sterility e.g all the almond varieties are reported to be
self-sterile or nearly so. When Tangelo (a citrus type) is grown in isolation it
fails to bear satisfactory crop because of self-incompatibility. Self
incompatibility is also observed in commercial varieties of loquat, like Golden
Yellow, Late Yellow, Tanaka and California Advance. Similarly, north Indian
varieties of mango like Dashehari, Langra, Chausa, etc. and that of lemons
like, Pant Lemon, Kagzi Kalan etc., also show self-incompatibilty and do not
bear a satisfactory crop when grown in isolation.
Inter-fruitfulness and inter-fertility : The ability of two plants or two varieties
to set fruits and develop seeds with each other’s pollen is called as inter
fruitfulness or inter fertility. Until, recently, dioecy was considered as the
main cause for self unfruitfulness e.g in Smyrna fig, date palm etc. Inter-
sterility was also regarded as one of the factors responsible for unfruitfulness
in some grape varieties. However, now it has been proved that most fruit
varieties are inter fruitful, even though they might be self-sterile, provided
that they bear good pollen. Hence any apple variety can pollinate other
variety successfully, but they should coincide in their flowering times.
 Unfruitfulness in Fruit Crops – Causes and Control Measures 163

Inter-sterility due to incompatibility has also been reported in some

instances e.g sweet cherries (Gardener, 1952). Unfruitfulness has also been
found to be due to cross incompatibility in some cases e.g apple varieties,
like Baldwin, Rhode Island Greening, Stayman Winesap produce very little
pollen which is not sufficient for fruit setting in other varieties. In apple
some strains of self unfruitful varieties are unfruitful even when they are
inter-planted with each other or with their parent variety.
Reciprocal crossings : It has been observed in certain fruit species that if a
certain crossing proved sterile, reciprocal crosses were also sterile. However,
in some cases few crossings have been found to be fruitful, but the reciprocal
crosses are sterile e.g Tragedy plum- a variety of European type is a good
pollinizer for several varieties of Japanese type, but their reciprocal crossing
is a failure. Sweet cherries, are good pollinizers for sour cherries, but reverse
is unsuccessful. Similarly in grapes, some species of grape like V. rotundifolia
and V. mumsoniana act as good pollinizers for Vitis vinifera, V. labrusca and
V. cordifoila while the reverse crossings are always a failure.
3. Physiological factors :
Slow growth of pollen tube : Poor growth rate of the pollen tube in the style,
possibly due to hormonal, nutritional and environmental conditions results in
unfruitfulness. Besides this, fertilization should take place within a short
time failing which abscission will take place at the base of style, ovary
pedicle or peduncle and fruit setting does not take place. Kuhn (1998) reported
that the main reason for poor fruit setting in sour cherry cv. Stevnsbar was
a lack of pollen tube growth through the style. In 15-42 per cent of the hand
pollinated flowers no tubes were found at the base of the style.
Self pollination in mango cv. Taimour revealed a degree of self
incompatibility where pollen tubes showed terminal plugs and failed to reach
the end of the style (Nariman et al; 1997). Hampson et al. (1993) reported
that in incompatible pollens, pollen tubes were distorted and did not penetrate
the stigma. Pollen tubes in mandarin cv. Clementine after self pollination
stopped growth in the upper part of the style due to slow growth, which may
be the possible reason for low fruit set (Eli et al; 1992).
Premature or delayed pollination : Premature or delayed pollination leads
to unfruitfulness. Mastrofilippe (1966) reported that premature pollination
followed by germination and tube growth causes fruit drop due to toxicity
in pistil. However, in case of oranges, premature pollination did not have any
deleterious effect. Low setting due to premature pollination was noticed in
persimmon, pear, plum and peach. Similarly, if pollination is delayed, the
flower falls without setting.
164 Fruit Tree Physiology

Nutritive condition of plant : Nutrition of plant controls the percentage of

defective pistils. Defective pistils are formed especially on exhausted or
weakened plants cause by over bearing, drought and poor nutrition. Nutrition
also determines the percentage of flowers carrying for setting, for maturity
and also the pollen viability. Deficiency of essential plant nutrients at critical
stages of flowering and fruit development leads to poor fruit set, more fruit
drop and even pistil drop.
Fruit setting of flowers in different positions : Fruit borne on terminal
growth have more competition in many fruit crops and mature and set under
normal nutritional conditions but per cent of set is small. This positional
competition takes place between fruits and branch as well as between different
fruits influencing fruitfulness. The apical or king (K), flower in the bud of
apple (Malus doestica) develops first, followed by the lateral flower (L)
which develop in sequence beginning at the base of the spur. In apple, old
spurs are less likely to initiate flowers than the young ones because spurs
remain productive up to 4-5 years only (Jackson, 2003). Ferree et al. (2001)
concluded that in unthinned trees limited resources are preferentially taken
by the K fruit which reduced set and size of the nearest L fruit. Koike and
Ono (1998) reported similar results with Fuji and Tsugaru and concluded that
fruit originating from lateral flowers are useful in producing high quality
fruits.
Poor pollen germination : Poor pollen germination due to various internal
and external factors reduces fruit set.

MANAGEMENT OF UNFRUITFULNESS
1. Balancing fruiting and vegetative growth : The main techniques for
controlling vigour of fruit trees and increasing their relative fruitfulness are
use of dwarfing root stocks, compact, short-internode scions, and of trees
training and pruning system which give horizontal or wide-angled branches.
Growth retardants are also used. Hansen (1980), studying pot grown apple
trees found that more than 70 per cent of the total dry matter increment of
4-5 year old apple trees of Golden Delicious on M9 root stock was as fruits.
Spur-type apple scion varieties commonly give trees of about 2/3 to 3/4 of
the size of the conventional variety, from which they arose by chance or by
induced mutation. Mika and Piatkowski (1986) expressing cropping efficiency
in terms of fruit yield per cm2 of trunk cross-sectional area reported relative
fruitfulness of Macspur/ MM 106 to McIntosh/ MM 106 from 242 to 161.
Upright growing shoots can compete successfully with developing fruits so
that fruitlets on young wood on vertically growing branches are commonly shed.
 Unfruitfulness in Fruit Crops – Causes and Control Measures 165

Bending down of branches results in the reduction of their growth

accompanied by increased flower bud formation and fruit set. The latter
effect is associated with an extended period of flower fertility (Robbie and
Knight, 1985). The influence of pruning varies with the amount, season and
kind of pruning. The judicious and proper pruning is needed to improve the
fruit set and fruit retention on the trees. Also the removal of dead wood
results in loss of carbohydrates reserves along with pruned wood. Severe
pruning promotes too much vegetative growth and hence reduces the
productivity. Singh and Daulta (1986) obtained maximum fruit set in peach
cv. Sharbati by light pruning i.e up to 12 buds. Reduced fruit set, yield and
nutrient content of leaves in apple after severe pruning was observed by
Mika (1992).
2. Induction of flowering : Gibberellins generally inhibit flowering with the
exception of GA, and C-3 epi-GA4, which have been shown to promote
flowering in apple trees (Looney et al; 1985). In mango, the highest number
of new flushes per shoot was achieved with severe pruning and spraying of
GA3 at 100 ppm (Sheban, 2009).
3. Overcoming winter chilling : At high altitudes in tropics and sub tropical
and temperate zones, temperatures may be low enough to induce dormancy,
but not sufficiently low to satisfy chilling requirements. Foliation is delayed
and bud break may extend over longer periods, giving problems in cross-
pollination. In Brazil, a combination of 2 per cent thiourea and 10 per cent
potassium nitrate, followed by 4 per cent mineral oil and 0.12 per cent
DNOC has proved useful in apples. Breeding of low chilling apple varieties
has made it possible to grow apples in warm fruit-growing areas in Israel.
4. Control of blossom quality : In pome and stone-fruits, blossom quality, as
measured by fruit-set, when blossoms are pollinated by hand, varies
considerably from tree to tree and from one season to next. It can be improved
by cultural practices such as branch bending (Robbie and Knight, 1985) and
also by others, such as autumn fertilizer application and early fruit harvesting
(Williams et al; 1980) which may be effective by influencing assimilate supply
to developing buds. High temperatures in early spring, between the completion
of rest period and blossom opening, have led to reduced fruit set (Jackson and
Hamer, 1980), and this has been shown to be at last in part due to a shortened
duration of ovule viability. Supplementary pollination at blossom time or
blossom thinning in a heavy crop year is likely to be beneficial.
5. Control of pollination : Pollen transfer may present an application problem
in fruits which are self incompatible. Many apple varieties are at least partially
self fertile especially under warm weather, but in most cases fruit set improve
166 Fruit Tree Physiology

by the use of pollinator varieties and bees. Cherries and plums also need
cross pollination and should be interplanted with pollinator varieties.
Breeding and selection of self fertile cultivars or clones of cherry and
apple may reduce difficulty for achieving satisfactory pollination, especially
in cool marginal areas of fruit production where temperatures at blossoming
time are sub-optimal both for bee activity and for pollen tube growth (Alston
and Spiegel-Roy, 1985). Until such improved varieties become available,
provision of suitable pollinator varieties and bee hives can be of great help
in ensuring satisfactory fruit set. There is an increasing tendency for using
crab apples, as pollinators as these occupy very little space in orchards. It is
advisable to have a number of pollinator varieties with widespread flowering
dates to ensure cross-pollination.
One of the basic requirements for setting fruit is an adequate requirement
of compatible pollen. With most true fruit crops, need for cross pollination
is recognized. Pollinating insects are necessary for fruit set on many cultivars.
and most will benefit from cross pollination. Under general conditions, the
closer a tree is to a pollinizer, the better the set will be (Fell, 2005).
The population of natural pollinators has gone down due to indiscriminate
use of pesticides and deterioration of ecosystem. Managed bee pollination is
very limited and available bee hives during bloom hardly meet 2-3 per cent
of the demand. All these factors have led to poor fruit setting in Delicious
apples (Gautam et al; 2005). Sequential introduction (introducing half of the
number of the recommended number of colonies at 10% FB and half at full
bloom) of honey bees in pear increases fruit set and yield by 50-80 per cent
due to increase in bees per tree and their mobility among the rows (Stern et
al; 2004). High fruit set and higher yield (50 to 100%) were obtained in Red
delicious apple by sequential introduction of honey bees (Stern et al; 2001).
Sharma et al. (2004) reported higher yields in apple orchards having > 15
per cent pollinizers than those with insufficient ones.
Cross pollination necessitates the availability of sufficient quantity of
compatible pollen, pollinizer cultivars flowering synchronously with the main
cultivar and suitable agent for the successful and effective transfer of pollen.
Therefore, suitable pollinizer cultivars must be interplanted at the time of
orchard layout. Apple cultivars like Starkrimson, Oregon Spur, Topread, and
Redchief and mangoes like Dashehari and Chausa (Ram et al; 1976); almonds
like IXL and Nonpareil (Kester and Asay, 1975) and many cultivars of sweet
cherry (Thompson, 1966) and European plum (Childers, 1983) are cross
incompatible.
 Unfruitfulness in Fruit Crops – Causes and Control Measures 167

Efect of distance form ‘Dlogo’ on fruit produced in ‘Fiji’ Apple.

(Matsumoto et al; 2008)

Efect of distance form ‘Maypole’ on fruit produced in ‘Fiji’ Apple.

(Sapir et al; 2007)

168 Fruit Tree Physiology

(Matsumoto et al; 2008)

6. Enhancement and induction of fruit set and retention : Fruit set is

frequently sub optimal in apple, pear, plum and cherry. This can often be
increased by tipping growing shoots to reduce competition from these either
just after flowering or at any early fruitlet stage (Quinlan and Preston, 1971).
Fruit set can be aided or induced by the application of plant growth substances
at flowering or after this. Mixture of GA3 with an auxin induces fruit set of
cherries. GA4+7 is generally more effective than GA3 on apples.
Pollinizers placed in solid rows

Shaded trees represent pollinizers

Pollinizers placed within rows

 Unfruitfulness in Fruit Crops – Causes and Control Measures 169

7. Control of frost damage : Fruit buds become more sensitive to frost towards
full bloom partly due to the result of increase in water content. Even buds
at early stages can be de hardened by a period of warm weather. Breeding
for late-flowering varieties should offer effective control (Jones, 1985) which
has been practiced by Alston and Spiegel-Roy (1985). Efforts to increase
frost hardiness by chemical sprays or to delay blossoming by use of plant
growth regulators have not yet been as successful to be adopted on a large
scale. Sprinkler irrigation is commonly used to protect fruit crops from frost
damage in the temperate zone. The water releases heat as it freezes, keeping
flower bud temperatures above the killing point of about 28°F.
8. Reduction in crop load : The need to avoid over cropping is another very
common reason for manipulation of fruiting. Commercially it is frequently
desired to have smaller number of large fruits rather than a large number of
small ones. Fruit thinning also reduces incidence of biennial bearing. The
first approach to reduce number of fruits per tree is by pruning. This usually
involves selective removal of thin shoots with weak fruit buds, which will
give small fruits and have less number of spurs. However, thinning fruitlets
after blossoming provides a more flexible technique for adjusting actual fruit
load in any particular year.
Blossom thinning by DNOC is practical when there is certainly of
oversetting in the absence of thinning e.g Golden Delicious apples grown in
favourable climates and with minimal pruning. It is achieved by physical
damage to blossoms, by preventing pollen germination and by inactivating
pollen-tubes growing down the styles. Post blossom thinning of apples is
usually achieved by the use of NAA or Carbaryl (sevin). Childers (1983)
suggested chemical application between 10 and 25 days after full bloom.
NAA application imposes a physiological stress on the tree which causes
shedding of least vigorously growing fruitlets. The growth of the remaining
fruit may also be checked, their ultimate size may also be less as compared
to hand-thining (Weaver, 1972). Stone fruits do not respond to NAA or
Carbaryl (sevin) but can be thinned using DNOC on blossom or 3-CPA on
fruitlets (Webster, 1980). Thinning fruitlets early in the season does not
necessarily decrease yield but improves fruit quality and increases yield in
the following year.
9. Hybridization : Improved technique of hybridization in mango will help to
evolve a larger number of hybrids for screening for desirable characters.
Embryological studies have shown that in mango pollen tubes grow down
the style and effect fertilization but the development of zygote is blocked due
to saprophytic type of incompatibility. Large scale hybridization may offer
a solution to the biennial bearing problem (Mukhjere et al; 1968).
170 Fruit Tree Physiology

10. Irrigation : Maintenance of proper soil moisture at the time of flowering

and fruit set is very important. Excessive soil moisture or water stress during
these processes results in flower and fruit drop. It has been reported that the
irrigation was stopped at the end of September and trees were stressed for
6 weeks until beginning of rains in November-December. The trees set fruit
and doubled the yield of litchi whereas irrigation at 20 or 40 per cent soil
moisture depletion gave maximum fruit set and yield in mango trees (Chandel
and Singh, 1992).
11. Nutrition : Proper and balanced supply of nutrients is always desirable for
realizing optimum fruit production. Generally, it is advocated that application
of fertilizers should be made a few days before emergence of blossoms as
it favour flowering and fruit set e.g nitrogen application after terminal bud
formation led to the development of flowers with enhanced embryo sac
longevity (Jackson, 2003). Application of boric acid (25 – 200 mgL–1 of boric
acid) increased the pollen tube growth in pear and the values were significantly
higher at higher concentration (Lee et al; 2009). Foliar application of H3Bo3
+ Paras (0.1 % + 0.6 ml/L) in walnut cv. Local selection resulted in highest
fruit set (32.50), fruit retention (41.35) and nut yield (4.02 kg/ tree) (Tomar
and Singh, 2007). However, excess use of manures and fertilizers may produce
abnormal flowers. Foliar applications of boron before full bloom or after
harvest in Conference pear increased fruit set and fruit yield (Pawel and
Marzena, 2003). Zn foliar sprays applied before anthesis is the most beneficial
in terms of fruit yield in citrus and grapes (Swietlik, 2002). Higher levels of
micro nutrients (0.8% zinc and 3.4% boron) showed marked effect on fruit
retention in litchi cv. Purbi. Zn at 200 ppm produced the highest fruit set
(13.33 fruit/ panicle) as reported by Sharma et al. (2005). A significant
increase in fruit set was recorded by Sharma et al. (2003) with the increasing
nitrogen levels.
Various nutrients such as nitrogen, phosphorus, calcium, magnesium, zinc,
potassium and boron have a specific role in maintaining the fruitful behaviour
of the plants. The balance application (through soil and foliar application)
and availability of these nutrients is essential for obtaining good fruit set and
yield. Fruit retention was significantly improved by the application of ZnSO4
in Kinnow (Daulta et al; 1986) and sweet lime (Arora and Yamdagni, 1986).
12. Use of growth regulators : The unfruitful behavior of several fruit plants
can be overcome by the use of plant growth regulators which may be due to
decreased fruit set and abscission at various developmental stages. Some of
the recent findings on the use of plant growth regulators to overcome
unfruitfulness in fruit trees are given in the following table.
 Unfruitfulness in Fruit Crops – Causes and Control Measures 171

Table : Commercial utilization (Bhat, 2009).

Fruit Crop Growth regulators Response

Apple Paclobutrazol 1000ppm Improve fruit set

NAA or Carbaryl (at full bloom Decrease fruit set, increase yield
stage)
GA3+NAA (At petal fall) Increase initiation and final set
Daminozide @ 2000ppm Increase flower buds
Ethephon (for thining) Decrease fruit set, increase size of fruit
Litchi TIBA, KNO3 Increase pollen fertility
NAA, 200 ppm Increase size, quality and retain bloom
Baoguoji (to prevent fruit drop) Increase fruit set, increase photosynthetic
rate
Grapes GA3 Increase fruit set
CCC@ 2000 mg/l Increased fruit set, inhibited shoot growth
Mango Pacloburazol Improve fruit set and retention
CPPU (10 ppm) 14 days after Increase fruit retention, yield and quality
bloom
Alar, TIBA, CCC Profuse flowering
Citrus GA3 Increase fruit retention
2,4-D @ 10 ppm Increase fruit retention
Paclobutrazol @ 2.5g/l Increase fruit set
Pear GA3 @ 50ppm Increase fruit set and retention,
parthenocarpy
1-MCP 0.75 /l during Increase fruit set
prepollination
GA3 10 g/ ha Increase fruit set
Chlormiquat @ 500ppm Role in flowering and fruit set
Peach Dormex, Paclobutrazol, KNO3 Reduce fruit set`
CCC 1000 ppm (mid bloom Improved fruit set in LeConte
period)
Alar @ 500ppm Increase fruit set and fruit size

CONCLUSION
Unfruitfulness is one of the serious and complex problems of fruit crops and
occurs due to various internal and external factors. The basic step to address the
problem is to select the crop or variety based on climatic and edaphic factors,
selection of high yielding and regular varieties, proper nutrition, planning and
planting of pollinizers, introduction of pollinators, rejuvenation of old orchards,
crop regulation measures, etc. Genetic and molecular approaches can be an
excellent tool in overcoming this problem.
172 Fruit Tree Physiology

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Further Suggested Readings

Bose TK, Mitra SK and Sanyal D (2001) Fruits: Tropical and sub tropical, Vol.1, Naya
Udyog, Kolkata, India.
Chadha KL and Awasthi RP (2005) The apple. Malhotra Pub House, New Delhi.
Singh A (1996) Physiology of fruit production. Kalyani publishers, Ludhiana, India.
Westwood MN (1993) Temperate zone pomology. WH Freeman and Co. San Fransisco,
USA.
 7

ALTERNATE BEARING IN FRUIT

PLANTS – CAUSES AND MANAGEMENT

A lternate bearing, the term is used to describe a fruit tree(s), orchard(s) or

branch(s) that produce a heavy crop during one year and a light or no crop
in the next year. It refers to the cyclical changes in cropping possessing two
phases “on” and “off” year.” “On” year with excessive fruit set, too little fruit
drop and too large crop and “off” year caused by lack of flowers, a poor fruit set
or excessive drop. Thus, alternate bearing is a phenomenon wherein the cropping
does not follow a systematic pattern. Sometimes, an “on” year is followed by
more than one “off” years and vice versa.

Years
Fig. : Schematic differences in cropping of a regular bearing and alternate bearing
cultivar under same environmental triggers (Monselise and Goldschmidt, 1982).

ALTERNATE BEARING INDEX (ABI)

The alternate bearing index (I) is a measure of a cultivars tendency to produce
alternating high and low yields. Ranges are from 0 to 1, with 0 = no alternation
and 1= complete alternation. I values are test cultivars ranged from a low of 0.19
180 Fruit Tree Physiology

to a high of 0.90. Generally, lower value is better, because this means the cultivar
tends to produce moderate yields yearly and quality and production is often
superior in these cultivars. The cultivars with I values above 0.65 are much less
likely to be accepted by the industry. This is especially true for cultivars with
large nuts because it is difficult to fill large nuts when the crop is excessive.
However, if methods are used to reduce crop load in years with an excessive
crop, the usefulness of some cultivars with high I values may be increased.

1 a2 a1 a3 a2 an an 1
I ...
n 1 a2 a1 a3 a2 an an 1

(Pearce and Dobersek-Urbanc, 1967)

Where n is the number of years and a1, a2, an-1 and an are yield of
corresponding years.
AB1 = (yield, year 1 - yield (years 2)/ yield, year 1 + yield, year 2) in kg/
tree for two consecutive (on/off) years. It can be expressed as a percentage by
multiplying with 100.
Table : Alternate bearing index (ABI) of some popular pecan cultivars.

Cultivar I Value Cultivar I Value

Caddo 0.32 Kiowa 0.65

Cape Fear 0.62 Money Maker 0.68
Desirable 0.40 Oconee 0.37
Elliot 0.68 Pawnee 0.61
Forkert 0.53 Sioux 0.64
Gloria Grande 0.19 Stuart 0.47
Kanza 0.72 Sumner 0.56

(Sparks, 1974)

Theories of Alternate Bearing

Two main theories have been proposed to explain why alternate bearing occurs
in fruit plants.
1. Starch depletion theory resulting from a heavy “on” crop implies that flowering
and fruit set for the “off” year takes place in trees with greatly reduced
reserves of carbohydrate energy e.g avocado trees (Whiley et al; 1996).
Scholefield et al. (1985) first established the close relationship between starch
reserves and yield in avocado.
 Alternate Bearing in Fruit Plants – Causes and Management 181

2. Nutrient withdrawal (primarily carbohydrates) hypothesis by a heavy crop.

The seeds inhibit flowering by exporting hormones, especially gibberellins
(GA’s) which have an anti flowering effect. GA applications inhibit flowering
in many tree crops, and GA concentrations in seeds are high. However, an
alternative explanation that seeds compete with shoot apices for flowering
hormones is equally possible (Dennis and Neilsen, 1999). In avocado, the
inhibitory effect of heavy fruiting on flower bud initiation on a particular
branch, as compared to a poorly fruiting branch can be observed.
AB can be understood as a hierarchy of causative factors which may be
controllable and uncontrollable and gradually getting closer to the proximate
and ultimate factor(s) (Fig.). The ultimate ones may reside in AB gene(s)
which are yet to be identified. However, a better understanding of tree growth
habit and phenological cycles will help to a great extent in AB management.

Fig. : Hierarchy of causative factors affecting alternate bearing

Factors responsible for alternate bearing

behaviour in fruit crops
Broadly divided into two main groups:
A. Endogenous, and
B. Exogenous factors
I. Environmental factors
II. Edaphic factors
III. Biotic factors (pests and diseases)
182 Fruit Tree Physiology

A. Endogenous factors :
1. Genotype differences : Alternate bearing tendency is controlled
genetically. It varies among different families, genera species and from
cultivar to cultivar e.g in apple, generally the shy bearers are regular
bearer, whereas heavy bearers are alternate bearer. On the basis of bearing
tendency apple varieties have been divided into 3 groups (Jonkers, 1979).
Regular bearers : Early Worecestor, Galla Beauty, Golden Delicious,
Golster, Jonagold, Rome Beauty.
Moderate bearers : Delicious group, Golden spur, Granny Smith,
Jonagold, McIntosh, Northern spy, Red Gold and Rhode Island Green.
Distinctly alternate : Baldwin, Benoni, Boskoop, Cox Orange Pippin,
Laxton’s Superb, Miller Seedling, Wagner, Wealthy and York Imperial.
In mango, there are two distinct groups of varieties with regard to
flowering behaviour. Varieties grown in north India like Langra (strongly
alternate bearer), Dashehari (moderately regular), Chausa and Bombay
Green flower in alternate years while those of south India like Totapari
Red Small, Neelam and Banglora flower and fruit every year ( show
distinct regular behaviour).
In citrus, most popular cultivars of sweet orange and grapefruit are
usually regular bearers. However, mandarin and its hybrids like Wilking,
Kinnow, Murcott and Michal are strongly alternate bearer. Similarly,
Satsuma a world famous mandarin of Japan and Nagpur santra of India
also show strong tendency towards alternate bearing. Other fruit crops,
like olive, avocado, walnut, pecan nut, pear etc., also have regular and
alternate bearing cultivars. The reasons for such differences in cultivars
with regard to bearing behaviour are not fully understood yet.
Regular bearing cultivars, when occasionally thrown out of the balance
by external causes, will rapidly regain their balance. Alternate bearing
cultivars on the other hand have more unstable habit and when thrown out
of the balance, continue cycling for many years until new environmental
events corrects their behaviour (Monselise and Goldschmit, 1989).
2. Competition between vegetative and reproductive sinks : In general,
fruiting has a strong effect on assimilate partitioning and tends to
antagonize vegetative shoot and root growth which are weaker sinks,
especially roots (Cannel, 1985., Wolstenholme and Whiley, 1990). The
order of priority among sinks is typically seeds > fleshy fruit parts =
shoot apices and leaves > cambium > roots storage. Seeds in the
developing fruits are the powerful sinks, which favour strong mobilization
 Alternate Bearing in Fruit Plants – Causes and Management 183

of photosynthetic products and hormones from the leaves. However,

there are instances, when vegetative flush may be a better sink than the
fruits. It is particularly true with mango, avocado and olives. A delicate
balance between vegetative and reproductive branches is needed for
regular cropping. However, in mango this hypothesis was rejected because
regular bearing cultivars (e.g Amrapali) produce flowers on vegetative
shoots of any age, but in alternate bearing cultivars (e.g Langra) vegetative
shoots of any age may not flower at all. Some mandarin hybrids like
Kinnow, Wilking ete. produce little or no vegetative growth in summer
during “on” year, which affects the following season’s flowering partly,
though, flowering usually takes place in spring season’s flush and rarely
on summer flush. Similar is the case with avocado and alternate bearing
mango cultivars, which rarely produce subsequent growth flush in “on”
year to flower in the following year.
3. Carbohydrate reserves (carbon: nitrogen ratio) : Annual patterns of
carbohydrate vary among species and genotypes based on growing
conditions, growth characteristics, crop load and other factors. Temperate
deciduous trees rely on reserve carbohydrates (i.e stored carbohydrates
that accumulate during periods when source supply exceeds sink demand)
for early season development because of the absence of a current supply
of photosynthates. Levels of stored carbohydrates have been suggested
by several workers to be a limiting factor in flower formation in “off”
years of alternate bearing cultivars of fruit crops (Mika, 1992). Fruit are
dependent on both current photosynthates and reserve carbohydrates as
evidenced by an accumulation of reserves after the spring flush followed
by a depletion of reserves during fruit development. The concentration of
starch in the trunk and large branches (10 cm in dia) is correlated with
biennial bearing in avocado. In litchi, the small and medium branches
(1-5 cm in dia) contain about half of starch present in spring before anthesis
and reduced during shoot and fruit development. During ‘on’ years root
carbohydrate concentrations are relatively low and the effects of girdling
are minimal during “off” years, there is a dramatic increase in root starch
and soluble sugar concentrations in non girdled control trees.
Studies on the C:N ratio indicated that higher starch reserve, total
carbohydrates and C:N ratio favour flower bud formation in most mango
cultivars, except Baramasi, and other regular bearing cultivars. Although,
nitrogen and carbon reserves play important role in flower bud initiation,
but these don’t form the primary cause of alternate bearing in fruit plants.
However, accumulation of these compounds may create favourable
conditions for the synthesis and action of the substances responsible for
184 Fruit Tree Physiology

flowering. The developing fruits draw large amount of reserve metabolites

from vegetative organs (leaves) during “on” year which contributes
towards alternate bearing.

Soluble sugars like glucose, fructose and manitol were found to

increase during “on” year, however, their concentrations were very low
in ‘off’ years in olive, indicating that carbohydrates role in alternate
bearing. They may possibly play a dual role i.e. i) reduce transport speed
of manitol from leaves to fruits in ‘‘off’’ year and cause its accumulation
in leaves ii) prevent transformation of glucose to fructose and manitol
during ‘‘off’’ year ( Nejad and Niroomand, 2007).
4. Hormonal imbalance : Higher level of auxin-like substances, an inhibitor
(similar to ABA) and lower levels of gibberellins like substances
(especially GA3) are vital for a floriferous shoot in mango. An inverse
relationship between the level of endogenous inhibitors in the shoot and
vegetative growth has direct relation with flowering in mango. Thus, an
optimal balance of endogenous levels of auxin, inhibitors (ABA) and
gibberellins must be there for mango tree to flower in an “on” year.
Concentrations of these regulators are in reverse order, with gibberellins
at a much higher concentration in “off” year or in vegetative shoot
(Chen, 1990). Al-shdiefat and Qrunfleh (2008) while investigating the
endogenous growth hormones during “on” and “off” years in olive
suggested a complementary relationship between endogenous plant
hormones, vegetative and reproductive growth.
Auxin : It plays an important role in induction of flowering in mango.
Shoots of Dashehari were found to contain high auxin during ‘on” year
than during “off” year.
 Alternate Bearing in Fruit Plants – Causes and Management 185

Gibberellins : It was found in higher concentration during “off” year

than in “on” year (Pal and Ram, 1978).
Cytokinin : They are directly involved in flower bud induction e.g Zeatin
riboside and zeatin was higher during ‘‘on’’ year in Dashehari mango
shoots (Agarwal et al, 1980).
ABA : In mango higher ABA levels were found in “on” year than in
“off” year.
Polyamines : Polyamines have an important function in reproductive
organ development serve either as nitrogenous sources or as signal
molecules regulating the fruit let abscission. Polyamine concentration in
leaves in “on” year trees was lower than that in “off” year’s trees (non
fruiting ones). The reduction of polyamines in leaves led to senescence
with subsequent enhancement of the biosynthesis of either ABA and/ or
ethylene and bud abscission.
In nutshell, higher levels of auxin like substances, ABA and cytokinin
and lower levels of gibbrellins are vital for floriferous shoot formation.
5. Flower inhibition by growing fruits : Flower bud initiation process for
the following year is strongly inhibited in most deciduous fruits, because
it occurs during the initial stages of fruit development in the preceeding
year (Buban and Faust, 1982). Seeds within the developing fruits exert
strong inhibitory influence on flower bud production (e.g, apples) or it
may cause differentiating flower buds to drop pre maturly (e.g pistachio
nut). The developing seeds increase the number of persisting fruits and
reduce the self thinning capacity. This is due to increased production of
different growth regulators and strong sink activity. The physiological
processes underlying these effects indicate that auxins produced by the
seeds within a fruit, moves into the fruiting spur and this movement is
stronger in alternate bearers (e.g Laxton’s Superb) than in regular bearers
(e.g Cox’ Orange Pippin) (Hoad, 1978). Besides, it has also been
postulated that auxins maintained in low concentration in spurs, which
initiate flowering process in an “off” year mainly by a greater translocation
of phloridzin from leaves, which enhances, IAA oxidase activity when it
is degraded to phoretic acid. Gibberellins produced by seeds also inhibit
flower initiation. Like auxins, they diffuse in large proportions from the
fruits of alternate bearing cultivars than those in regular bearer ones. It
has been found that the diffusion of gibberellins is early in alternate
bearing cultivars (about 2 weeks after bloom) while it is late in regular
bearing cultivars (about 5 weeks after bloom) and hence unable to inhibit
flower bud initiation in them (Ebert and Bangerth, 1981). Not only the
supply of hormones from the developing seeds of the fruit, but there are
186 Fruit Tree Physiology

other ways also by which fruits inhibit the flower initiation process in
other ways also.
Developing fruits and seeds within fruits draw some important
metabolites which are necessary for flower bud initiation as developing
fruits are powerful sinks for photosynthates. Hence, it can be summarized
that when there is considerable crop of seeded fruits, different hormonal
and nutritional factors together depress the flower formation, thereby
initiating the alternate bearing process.
6. Pollination : Self incompatible forms require cross pollination for
satisfactory crop. However, excessive pollination may lead to alternation
in bearing as self pollinated apple cultivars tend to show extreme
alternation compared to the self sterile ones (Williams, 1973). But in
citrus, the alternating mandarin cultivars may be both self sterile and self
fertile e.g Satsuma mandarin of Japan and Washington Navel sweet orange
of the USA, although self sterile, but exhibit strong alternate bearing
behaviour. Improper pollinizers and pollinators may cause poor yields in
fruit crops like apple, annona, mango, avocado and various self
incompatible mandarins, lemons and even some stone fruits. In an orchard
of Delicious apples for effective and fruitful yield, it should have at least
33 per cent plants of a pollinizing variety (Tydernan’s Early Worcester,
Golden Delicious, Granny Smith, Manachurian, Golden Hornet etc.),
more specifically a combination of different pollinating varieties and
2-4 bee hives/ ha.
In avocado, pollination plays a vital role and is considered to be the
most limiting factor for fruit set. The colour and size of flower in avocado
is not attractive enough to attract bees. Moreover, due to complexity of
problems like protogynous diuranally synchronous dichogamy (PDSD) -
opening of flowers at different times of the day and non synchronization
of opening of male and female flowers, pollination is a major problem
in fruit-set and yield (Bergh, 1975).
7. Effects of seeds on fruit drop : Developing seeds are the major source
of auxin production which inhibit fruit drop in seedless cultivars as in
citrus. However, if the concentration of auxins falls below a certain
critical limit, the fruits even of the seeded cultivars may exhibit heavy
drop. The peel of the fruit plays a major role in the production of auxins
in seedless cultivars e.g citrus. Regular bearing cultivars maintain fruit
load year after year through self thinning capacity. But alternate bearers
are unable to thin out fruits in “on” year and thereby produce little or no
crop in the following year. For example in regular bearing oranges,
grapefruits or plums, the percentage of flower bud production is not
 Alternate Bearing in Fruit Plants – Causes and Management 187

adversely affected as long as fruit production does not reach a minimum

level, depending on the tree vigour, as determined by size and genetic
potential. Regular bearing cultivars have also tendency to control fruit
drop and produce moderate amount of fruits year after year.
8. Effect of leaves : Leaves contribute in two ways, i.e. nutritional and
hormonal to the bearing behaviour of plant and both of these factors
either individually or in combination, play important role in vegetative
or reproductive growth of the plant. Both of them are produced in close
proximity and translocated to different parts through the same vascular
system and it is difficult to diffentiate between these two factors. In most
of the fruit plants e.g mango, apple (Harley, 1932), citrus (Moss et al,
1972) and pistachio nut the contribution of leaves to flower bud formation
has been established. Photosynthates and different endogenous growth
regulators produced by leaves are generally regarded as a pre requisite
for flower initiation. In pistachio nut presence of appropriate number of
leaves may have antagonistic affect on flower buds and drop of growing
nuts. Role of leaves for fruit-set and development is well documented in
citrus. The fruits are borne on mixed inflorescence presumably because
of higher gibberellin’s activity. GA is very effective in fostering fruit set
and development of citrus fruits. Gibberellins produced by the leafy
inflorescence may be the cause of better fruit-set in citrus. This leafy
inflorescence provides sufficient foliar surface and photosynthetic activity
to support fruit set and further development of the fruits in early stages.
Similarly, in pecans also, leaves strongly affect fruit set, development
and return bloom.
Leaf area also plays an important role on fruit set, growth and
development in different fruit crops. It has been recommended to retain
one fruit per 25 leaves in Satsuma mandarins in Japan. Similarly, certain
ratios have been suggested for apples (40 leaves per fruit) and peaches.
It is evident that presence of certain number of leaves has a significant
influence on bearing behaviour in various fruit crops.
9. Crop overload : The most universally accepted cause of alternate bearing
in most fruit crops is the heavy crop produced during the “on” year.
General effect of over crop load is the exhaustion of plants because
fruiting is an exhausting process. The future flowering and fruiting is
affected by the number of fruits retained on the tree till harvest, though;
it may be a varietal feature. The fruit load has a direct influence on the
production of new shoots and their subsequent fruit bud differentiation
in the following season. Higher fruit number creates a cumulative sink,
which requires a continuous supply of food reserve. Both mineral and
organic nutrients may be obtained from either newly assimilated materials
188 Fruit Tree Physiology

or reserves previously accumulated in different tissues of the plants. Tree

collapse may occur due to over load during “on” year leading to depletion
of reserve food material. Because of carbohydrate starvation the trees
bearing heavily in an “on” year suffer degradation in the following years.
Starch is the most common carbohydrate reserve in the plants and is the
best indicator for the nutritional status of a plant than sugars e.g in citrus,
apple, pecans and mangoes. It has been found that starch levels are
usually higher in shoots during “off” year and low in an “on” year. Thus,
the starch needs to be replenished regularily to overcome its exhaustion
due to crop overload (Goldschmidt and Comb, 1982).
Crop over load may also alter the hormonal balance of the tree and
thus may affect the future morphogenetic and developmental events e.g
production of excessive gibberellins by citrus fruits during “on” year
usually prevents flower bud formation in the following year. Similarly,
accumulation of ABA in over loaded citrus trees may reflect an over
loaded stress. Crop load seems to be the main contributing factor for
“on” or “off” year in fruit crops and has a systematic depressing effect
on many manifestations of tree growth (Crane and Nelson, 1972).

Table : Effect of crop load on starch, arginine and proline concentrations (mg/g)
dry weight of shoot apical buds in avocado.

Tree type Starch Arginine Proline

On crop trees 45.45 21.69 13.55

Non crop trees 115.09 24.99 15.99

10. Tree age and vigour : This factor in majority of cases is linked intimately
to the effect of nutritive status of the plant or its limb, locality and the
season, having direct effect on flowering and fruit set, and thereby
unfruitfulness. Excessive vegetative growth reduce fruitfulness of trees,
on the other hand optimum vegetative growth to support the crop load
is very much essential otherwise leads to fruit drop either at early or later
stages e.g as in strawberry, walnut, etc.
11. Natural abscission of buds, flowers and fruits : Even when pollination
is successful and the trees are in good health, not all pollinated flowers
lead to effective fertilization of ovaries that will later become edible
fruits. A number of environmental and endogenous factors contribute to
reduced fruit set due to abscission of buds, flowers, and fruits. Large
fruited trees such as apple, may shed 90 per cent or more of their pollinated
 Alternate Bearing in Fruit Plants – Causes and Management 189

flowers and young fruits. However, since a mature apple tree may have
more than 100,000 flowers at full bloom, it is a benefit that not all
flowers set fruit. Small fruited species shed a lower percentage of flowers.
On many fruit and nut trees, flower buds, flowers, and immature fruits
abscise (separate) from the trees at distinct times during the year,
particularly during June drop (occurs in May in California). The fruit
that remain on the tree are said to have set. Profuse flowering decreases
the yield of avocado because of immature fruit drop after early fruit
abscission of the ‘off’ and ‘on’ crop years.
12. Nutrient status : Optimum nutrient level is pre requisite for regular
bearing. N, P and K decreases while Ca increases in “on” year. “off” year
bear reverse condition where N, P and k are sufficently present in stem
and bark whereas Ca level decreases. Nutritional deficiency caused
irregular bearing in mango where proportionately increase in N led
vegetative growth and proportionately drecrease favoured flowering. Low
K content and high Mg and Ca contents are present in “on” year trees
of Balandy mandrin as compared with “off” year (Bannisab, 2007).
13. Age and size of shoot : Physiological maturity of the shoot also plays
an important role in determining the bearing behaviour of fruit crops e.g
in mango, the earliest emerged flush (usually 3-5 flushes) which is 8-10
months old is capable of producing flower buds. However, there are
contradictory results as well. But one thing is clear that non flowering
shoots lack in some vital substance necessary for flower bud formation
e.g. apple (Jonkers 1979).
14. Fruiting habit : Terminal flower formation is regarded as a characteristic
favouring alternate bearing e.g mango and apple. Any climatic vagaries
(snowfall, hails, wind) directly destruct the inflorescence leading to
conversion of an “on” year into an “off” year.
15. Growth pattern : Tree architecture also plays an important role in
determining the alternate behaviour e.g columnar and spur type cultivars
tend to be alternate bearers as such conditions lead to a strong “on” year
ahead and subsequent year becomes “off” year. However, spreading and
terminal bearing cultivars bear crop regularly.
16. Others : Chlorogenic acids in developing fruits of olive inhibit floral
initiation next year.
17. Root stocks : Root stocks play important role in the process of alternate
bearing. For example root stock affects regularity in bearing in apple and
citrus. Weak or dwarfing root stocks in apple reduce biennial problem in
apple like the malling series (M9, M27, MM106 etc). In Mediterranean
region Sour orange (citrus root stock) is one of the major causes of
190 Fruit Tree Physiology

alternation in almost all the mandarin cultivars. In avocado 10 root stocks

were used to observe their effect on alternate bearing behaviour and it
was found that trees showed AB on all root stocks (RS) particularly on
Topa Topa and Toro Canyon and lowest ABI on G755A, B and C (Whiley,
2009).
18. Branch autonomy : Orchards or single plants in an orchard and even
individual branch in a single plant behaves as autonomous entities with
regard to alternation. Autonomy of branches increases with the increase
in age in many fruit crops. When a particular branch or part of a branch
does not flower in one year, it produces heavy crop in the next year.
Autonomy is also observed among cultivars and in these cultivars one
branch or its part may or may not be fully autonomous. In apple and
plums, it has been found that lack of fruits on a particular branch than
the rest of the plant may lead to an increase in the flower production on
a bearing branch, and in general, there is reciprocal influence between
the bearing and non bearing parts of a plant.

(Monselise and Goldschmidt, 1982)

B. Exogenous factors:
(i) Environmental factors : Different fruit crops require different climatic
requirements for their proper growth and fruiting. However, any adverse
climatic condition may trigger or initiate the phenonmenon of alternate
bearing. But the conditions, which don’t act as trigger in a particular
zone can become trigger in another zone or for different trees in the same
zone. Among the different environmental factors, climatic and edaphic
stresses are most important ones leading to alternation. In addition, indirect
effects of environment (incidence of insect-pests and diseases) can also
lead to alternate bearing.
 Alternate Bearing in Fruit Plants – Causes and Management 191

Vrious environmental factors included are:

1. Temperature
2. Light
3. Relative humidity
4. Rains
5. Snow
6. Spring frost
7. Hails
8. Water stress
1. Temperature : Temperature has a great importance as it affects
flowering and fruit set in several ways. A period of cool yet frostless
weather is conducive to better blossoming, fertilization and fruit set.
The abscission of flower bud, fruit, etc. is a function of temperature.
Winter thaws followed by cold weather can kill flower buds. Frost
during blossoming can damage flowers, and cool temperature can
decrease the viability of pollen. Temperature is quite important for
pollen germination in plum, cherry, apple, pear, etc. where 4°C or
lower completely checks the same. Hot, drying winds during the
blooming period can desiccate the tender stigmas or styles, sharp
environmental variations reduce pollen viability and reduce avocado
fruit set. The pollination and fertilization of almonds are negatively
affected by the low temperatures. The low temperature during
blooming period can prevent the germination of pollen on stigma to
prevent the development of pollen tubes in the style. Under favourable
conditions, the fruit-set may be high, but high temperature and dry
hot winds during post set period may force the fruits to drop off,
which again help to trigger alternation in fruit plants.
Hot and dry winds during the blooming period can desicate the
tender stigmas or styles, sharp environmental variations reduce pollen
viability. Finally heat can reduce fruitfulness by injuring the tree.
Flower and fruit let drop in Satsuma mandarin exceeds in May-June
when temperature was 35-37°C. High or low temperature may also
interfere with the pollen carrying insects, particularly bees, flies and
wasps, as they don’t take flight under very low or high temperature,
which results in poor pollination and thereby in poor fruit set.
2. Light : Adequate sunlight is important for flower differentiation and
fruit yield. Good exposure to sunlight is one of the most important
factors influencing the formation of flower buds on fruit trees.
192 Fruit Tree Physiology

Through the process of photosynthesis, the leaves on the tree produce

sugars used for vegetative growth, flower bud formation, and fruit
development. If sunlight is limited, the tree will continue to grow,
but there will not be sufficient sugars for the formation of flower
buds. Under such conditions, young trees will be slow coming into
bearing, the blossoms will be sparse, and the fruit that forms will be
poorly colored (Nuzzo et al, 1999). Dwarf trees also will grow larger
than anticipated. For these reasons, fruit trees should be planted in
locations where they will receive maximum sunlight. Avoid shaded
sites near buildings or other trees. Light is pre requisite for
photosynthesis and low light intensity or its duration reduces the
carbohydrates reserves in the trees. In addition to this, poor light
conditions promote fruit abscission and had positive effect on flower
bud number per fruiting shoot and fruit set and growth rate were
lower in the shaded zone of the canopy. Pollinator activity is positively
correlated with the light intensity.
3. Humidity : Poor fruit set and excessive drop of the fruits in different
fruit crops may occur due to extremely low or high air humidity e.g
olive (Gardner, 1966), oranges, grapefruits and most of the sub-
tropical and temperate fruit crops. Drying or desiccation of stigmatic
fluid occurs due to low and high humidity which may affects fruit
set in olive and avocado due to poor pollen germination. Under low
or very high atmospheric humidity activity of bees and other pollen
carrying insects is hindered, which also results in poor fruit set in
many fruit crops. There is increased tendency towards vegetative
growth in mango and hence poor fruit set under humid conditions
(Addicot, 1973).
In avocado, low atmospheric humidity causes drying of stigmatic
secretions so that pollen does not germinate. Rain is an important
factor that indirectly affects fruit setting by disturbing the process of
pollination, germination of pollen grain and even staminal fertilization.
The poor germination of pollen in almonds was attributed to damp
weather during fruit set. Wet and humid weather favors anthracnose
and poor fruit set in mango. Relative humidity on its own is an
important factor in bee activity. Low temperature and high humidity
have the double effect of reducing bee activity and slowing the
release of pollen.
4. Rains : Erratic rainfall or drought during flowering, may convert
“on” year into an “off” year directly or indirectly. Heavy rains may
wash off the pollen grains and hence interfering with the pollination
and reducing fruitset. Water stress during flowering, fruit-set and
 Alternate Bearing in Fruit Plants – Causes and Management 193

fruit growth and developement is equally harmful. In mango, rains

during blooming period reduce the crop by creating favourable
conditions for the diseases like powdery mildew and anthracnose.
Besides rains during flowering interfere with bee activity and reduce
the fruit set. In cherry, rains cause cracking of fruits during May-
June and sometimes lead to low or no yields in the following season.
In almonds, rains during flowering may wash off pollen and even
low temperatures lead to freezing of rain water and thereby freezing
injury.
5. Snow : Snowfall during winter has tremendous effect on fruitfulness
of apple as it helps in completion of chilling requirements and ensures
better flower bud development in spring. But snowfall during pre
bloom or blooming period has negative effect on fruit set and may
lead to an alternate year of an otherwise “on” year.
6. Spring frost : Frost during blooming period leads to death of flowering
buds and flowers which are very delicate and sensitive to very low
temperatures and thereby leading to poor fruitset and alternation e.g
almond is highly susceptible to spring frost.
7. Hails : Hails at the time of flowering or fruiting lead to heavy flower
and fruit drop and also directly hit the buds and other sensitive plant
parts thereby rendering them to bear low or no crop during on years.
8. Water stress : Water stress in fruit plants is associated with the decrease
in nucleic acid and protein content of the fruit and formation of
abscission layer. Severe water stress (-1.5 to -2 MPa) in Satsuma
mandarin during autumn causes heavy leaf fall, and reduces the
percentage of flowering nodes. It may be due to increased levels of
either GA1 or GA3 during this severe water stress (Cemeich, 2005).
(ii) Edaphic factors : The structure, texture, porosity, aeration, moisture
contents and presence of salts may have some influence on alternate
bearing behaviour of fruit crops. Soil conditions, which are detrimental
to root activity, reduce plant health, causing them to produce low yields.
For example, problematic soils like salt affected or acid soils may cause
scorching of leaves and premature leaf fall, which reduces the
photosynthetic area and reserve food. During growth or flowering soil
moisture stress leads to leaf and fruit abscission and production of sterile
flowers in many fruit crops. Drought favours excessive drop of
reproductive organs, leaves and developing fruits and thus may have an
effect on alternation (Hartman and Pentsos, 1961).
194 Fruit Tree Physiology

(iii) Biotic factor : Mites in apple have been found to cause alternation for
a couple of years because they suck the sap from leaves and branches
and lead to tree exhausation of essential nutrients etc. Similarly, hoppers
in mango and aphids in pecan lead to alternate bearing. Disease like
premature leaf fall and scab in apple, powdery mildew in mango etc.
may convert an “on” year into an “off” year

Diagramatic presentation of various factors resulting in alternate bearing

(Lavee, 2007)

MEASURES TO OVERCOME BIENNIAL BEARING

Several strategies have been suggested from time to time to control or reduce the
problem of alternate bearing in different fruit crops. Some of these strategies can
be used, but all of them may not be suitable for a particular fruit crop at a given
locality and thus these may be tried with some suitable modifications.
1. Proper orchard management: A proper cultural schedule is very important
for maintaining fruit trees in healthy and disease free conditions and thereby
obtaining better plant performance. Various cultural practices such as regular
ploughing, judicious manuring, fertilization, weed management and assured
irrigation after fruit set and ringing/ girdling of branches etc. have been
recommended to overcome the problem of alternate bearing in different fruit
 Alternate Bearing in Fruit Plants – Causes and Management 195

crops. Proper maintenance and upkeep of the orchard may help in reducing
the erratic bearing, but can’t induce regularity in bearing among biennial or
alternate bearing cultivars. For obtaining optimal crop and regularity in
fruiting, various management practices including, fruit thinning, fruit abcission
control, plant nutrition and judicious pruning can be adopted.
2. Regulation of flowering :
Use of chemicals : Use of chemicals for the control of flower formation is
one of the most desirable controlling mechanisms of alteration in fruit trees.
Example
1. Use of GA3 on Australian oranges e.g autumn application of GA is
recommended for eliminating excessive flowering in winters, which
usually takes place between the ‘off’ and “on” years. Moderate crop in
the expected “on” year can be obtained in the Valencia oranges in Australia
by GA application. Similar practices can be followed in mandarins also.
2. Retardants like daminozide and cycocel in citrus induce flowering in
“off” year. With the application of cycocel (CCC), “off” season flowering
can be induced in lemons during autumn, so as to obtain fruits during
peak summer months that can fetch higher market prices.
3. Morphactins, ethephon, NH3 ions, cytokinins, daminozide and maleic
hydrazide (MH) can be used to increase flowering intensity, whereas
bromacil could be used to reduce it. Daminozide induces flowering in
meadow apple orchards. Cycocel and alar each at 5000 ppm are used to
induce early and intensive flowering in mango. Ethrel (2-Chloroethane-
phosphonic acid) is used to induce flowering in mango during “off”
years.
4. Auxin-61 and KNO3 have also been reported to induce “off” season
flowering in fruit crops (Bondad and Linsangan, 1979). In Philippines
KNO3 has been reported to be more economical and effective for inducing
regular flowering in polyembryonic cultivars (strongly alternate bearers).
However, its effectiveness has not been proved in India.
5. Dormex (3%) applied 40 days after bud break enhances flowering and
improved fruit set in mango.
6. TIBA @ 50 ppm applied during October stimulated flowering in mango
during “off’ year.
7. Chlormequat @ 2000 ppm increases average number of panicles per
shoot in mango (Chaco, 1971).
8. GA3 @ 20 ppm decreases alternate bearing in kinnow.
196 Fruit Tree Physiology

9. Trunk injection of Hass avocado trees with the auxin transport inhibitor tri
iodobenzoic acid (TIBA). TIBA alone or in combination with cytokinin
increased the total number of floral shoots produced by shoots with fruit
compared to the untreated crop. TIBA combined with compound cytokinin
increased the total number of floral shoots and the number of determinate
inflorescences (P = 0.0789) compared to untreated ones. However, TIBA
alone increased return yield more than 2.5-fold (Lovatt and Riverside, 2010).
Smudging : In Philippines, in 1923, the practice of smudging has been
reported to induce flowering in mango. The exact physiology behind smudging
to induce flowering is still a question and a point of great controversy.
However, some workers attribute this flowering to heat, whereas the others
opine that CO2 is responsible for it. In India, however, this practice has not
been found to be helpful in inducing flowering in mango.
Girdling : Horticultural practices like girdling, ringing or notching of branches
can also control flowering in fruit plants. Girdling has been used successfully
to induce flowering during “off” season in various fruit crops like citrus,
mandarins, avocados, apples etc. Only half of the branches in a particular
tree should be girdled to get regular crop in alternate bearers.
Paclobutrazol : Paclobutrazol - a growth retardant and GA antagonist has
been proved to be the most effective chemical in inducing regularity in
bearing in many fruit crops. Paclobutrazol (PP333 or Cultar) is most widely
used chemical to induce dwarfness and regularity in fruit crops like apple,
pear, olive and mango. Both soil application as well as foliar application is
effective. Soil application of Paclobutrazol @ 5 g a.i./ tree is the most desirable
dose and should be applied about 90 to 120 days before flowering to get
desirable results and its effects may last for 2 to 3 years. It is recommended
to use Paclobutrazol only once in three years. In Ambri apple (highly alternate
bearer) Paclobutrazol @ 2g/ m2 of canopy area has been found to overcome
alternate bearing.
3. Tipping growing shoots : Upright growing shoots can compete successfully
with developing fruits so that fruitlets on young wood on vertically growing
branches are commonly shed. Bending down of branches results in the
reduction of their growth accompanied by increased flower bud formation
and fruit set.
4. Proper nutrition : Various nutrients such as nitrogen, phosphorus, calcium,
magnesium, zinc, potassium and boron have a specific role in maintaining
the fruitful behaviour of the plants (Ravishankar et al, 1989). The balance
application (through soil and foliar) and availability of these nutrients is very
much essential for obtaining good fruit set and yield. Boron is essential for
 Alternate Bearing in Fruit Plants – Causes and Management 197

pollen germination, for successful growth of the pollen tube through the
stigma, style, and ovary to the ovule, and the mitotic divisions necessary to
produce the sperms (Coetzer et al; 1993). Jaganath and Lovatt (1996) tested
the effect of pre bloom foliar applications of boron and low biuret urea at the
cauliflower stage of inflorescence development on pollen tube growth, ovule
viability, and yield. They found that boron and low biuret urea increased the
number of pollen tubes reaching the ovary by 2.5 and 2.0 fold, respectively,
and increased the cumulative yield by 25 per cent and 23 per cent, respectively
in avacado.
5. Deblossoming : This practice is employed to conserve the reserves of the
shoots, which could otherwise be depleted later on during the process of fruit
development. The deblossomed plant puts on new vegetative growth, which
flowers and fruits in the following year. To regulate bearing behaviour in
mango, apple, citrus and some other fruit crops, deblossoming has been
found very useful. It can be done manually, but deblossoming with DNOC
and NAA has been very effective to overcome alternation in bearing. Ethephon
and NAA have been found to be very effective chemicals for crop regulation
in oranges and mandarins. NAA (200-300 ppm) have been found useful for
deblossoming in Kinnow during “on” year to get optimum crop in “off” year.
6. Fruit thinning : The problem of alternate bearing in many fruits, such as apple,
oranges, mandarins and olives can be reduced by thinning of fruits in “on”
year. Thinning reduces the crop load in the “on” year, so as to get some crop
every year. Thinning of fruits can be done by hand or by chemicals like sevin,
DNOC, bromacil and NAA at slightly higher concentrations. However, thinning
of fruits is not possible due to heavy fruit drop in the initial stages of fruit
growth. Moreover, even without pollination and fertilization fruits may grow
up to pea stage, but drop off afterwards. Till the end of April fruit drop continues
and panicles thinned earlier may not carry any crop at all. Moreover, food
reserve exhausation is caused by the developing fruitlets when these are still in
pea stage of their growth and the plant takes at least some time to recoup,
leading to alternate bearing in mango. From above discussion it is clear that
alternation in bearing can be reduced by thining in some fruit plants, while in
the others it should be done with great care like mango.
7. Selective pruning : Selective pruning regulates cropping in most temperate
fruit crops. Pruning is primarily done to maintain a proper physiological
balance between vegetative and reproductive growth. Pruning is also
considered as a thinning process to reduce crop load and thereby getting
regular crops in deciduous fruit crops. Mango, being an evergreen crop
(earlier pruning not recommended) responds favorably to pruning. It has
198 Fruit Tree Physiology

been used successfully to obtain regular crops from high-density orchards of

Amrapali and Dashehari mangoes. Under north Indian conditions, the best
time for pruning mango is after harvesting (July). Selective pruning is equally
useful for reducing alternation in apple, olive, grape, avocado and citrus
(Leonardi, 2009).
8. Early harvesting : Harvesting is sometimes delayed in citrus, avocados and
apples also, which affects flowering and fruiting of the trees in the following
years, and thus, results in an unusual trend of alternate bearing. Timely
harvesting or stripping trees completely of the first crop is recommended,
which would help in producing large crop of good quality in “off” year also.
9. Growing regular bearing cultivars : Growing of regular bearing cultivars
is the most suitable and desirable alternative to overcome the problem of
alternate bearing. Some regular bearing varieties/ hybrids have been developed
in different fruit crops e.g in mango varieties like Rumani, Amrapali, Mallika,
Arka Aruna, Arka Puneet, Ratna, Dashehari-51, Pusa Arunima, Pusa Surya,
Ambika etc., besides the traditional regular bearing cultivars like Bangalora,
Neelum, Totapuri Red Small. Spur type cultivars of apple like Oregon Spur,
Golden Spur, Wells spur, Red chief, Early Red One, Tydemans early worcestor
etc. which are regular bearer and produce quality fruits, have been developed
in apple. In other fruits crops also, emphasis should be given for growing
regular bearing cultivars.
10. Dwarfing root stocks : Use of dwarfing and semi dwarfing root stocks may
be encouraged for apple cultivation because dwarfing and semi-dwarfing
root stocks in apple are known to induce dwarfing and precocity in bearing
in the scion cultivar budded or grafted on them. Root stocks like M9, M27,
MM106 and MM111 induce precocity in bearing in standard and spur type
apple cultivars. Similarly, Dragon Fly should be used in citrus in many parts
of the world but in India, Troyer citrange induces regularity and precocity in
bearing in Kinnow mandarin. Thus, exploitation of potential root stocks can
help to overcome alternate bearing to a great extent. In avocado, clonal root
stocks also reduced the alternate bearing tendency to a great extent (Mickelbart
et al; 2007).
11. Gibbereliins as anti flowering agents (“on” crop year) : A 100mg/ l GA3
spray applied in early winter before an “on” bloom reduced the number of
inflorescences, increased the number of vegetative shoots, and reduced “on”
yield by 47 per cent - all desirable responses to possibly break or modify an
AB cycle in avacado (Lovatt, 2010).
 Alternate Bearing in Fruit Plants – Causes and Management 199

CONCLUSION
Alternate bearing is an economic problem for growers and the industry affecting
fruit size and net economic returns, causing a price fluctuation between “on” and
“off” years, and the loss of market share during “off” years that is not always
regained in the following “on” year. It is difficult to find a hypothesis that
explains all aspects involved in alternate bearing. Competition between developing
fruits and new vegetative shoot growth for carbohydrate and other nutrients,
especially nitrogen, might play an important role in the biennial bearing pattern
of fruit trees. Orchard management practices can be used to successfully eliminate
alternate bearing and to sustain regular and economic yields year after year.
Horticulturally, the key is to attempt the maintenance of the ideal vegetative:
reproductive balance as determined for the particular environmental conditions,
orchard intensification through closer spacing, pruning, girdling, provision for
cross pollination, choice of cultivar, root stock, and use of growth retardents like
paclobutrazol and uniconazol etc. and management philosophy to reduce the
intensity of AB. Research in specific areas is needed in order to fine tune orchard
management practices that growers can use to eliminate alternate bearing.

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Further Suggested Readings

Chadha KL and Pal RN (1993) Biennial bearing in mango. In: Advances in Horticulture
4: 2025-2046.
Rao MM (1998) Problems of alternate bearing in mango: causes and management. In:
Mango cultivation. Pp. 339-362.
Singh RN (1997) Biennial bearing in fruit trees-accent on apple and mango. Bull. 30,
IARI, New Delhi.
 8

PARTHENOCARPY–
A MECHANISM FOR SEEDLESSNESS

P arthenocarpy is the development of fruit without pollination and fertilization.

It is termed as formation of seedless fruits in the absence of pollination or any
other stimulus. The term was introduced by Noll (1902). In botany and horticulture,
parthenocarpy (literally meaning virgin fruit) is the natural or artificially induced
production of fruit without fertilization of ovules. The fruit is, therefore, seedless.
Parthenocarpy occasionally occurs as a mutation in nature, but it is usually
considered a defect, as the plant can no longer sexually reproduce, but may
propagate by asexual means. Gustafson (1942) listed the occurrence of
parthenocarpy in about fifty species including apple, pear, grape, citrus, breadfruit,
date, fig., banana and pineapple. These are usually seedless. Parthenocarpic fruits
are usually but not always seedless and seedless fruits are not always parthenocarpic
because seedlessness may be caused due to embryo abortion resulting due to
inability of the embryo to accumulate the required food reserves during its
development.
Parthenocarpy may be viewed as a phenomenon or an ultimate stage of a
sequence in which fruit development becomes progressively independent of seed
development. Some fruits are entirely dependent on their seeds for fruit
development. In strawberry, removal of the achenes (commonly called seeds) at
any stage of development leads to the cessation of fruit growth. There is no free
auxin in the receptacle tissue. Whereas Washington Naval orange and Black
Corinth grape develop even though they are not fertilized and have no seeds and,
thus, they are completely independent of them.
Many seedless horticultural varieties of fruits are exclusively parthenocarpic.
Many seeded fruits are not necessarily capable of parthenocarpy and even some
single seeded fruits can be parthenocarpically set. The seeds from parthenocarpic
fruits are not fully developed and they do not germinate.
 Parthenocarpy – A Mechanism for Seedlessness 205

Types of parthenocarpy
Parthenocarpy is broadly of two types
1. Natural or genetic.
2. Artificial or induced.
1. Natural or genetic parthenocarpy: It has been documented in various crops
like grape, tomato, mandarins, banana etc. It is further classified into following
classes:
● Obligatory parthenocarpy: Parthenocarpy due to prevention of pollination.
It always results in seedless fruits.
● Facultative parthenocarpy: It occurs due to conditions adverse to
pollination or fertilization or resuls from genetic sterility due to continous
vegetative propagation e.g tomato, banana, pineapple etc.
● Vegetative or autonomic parthenocarpy: Pollination or other external
stimulation is not required to produce parthenocarpic fruit. e.g. Washington
Navel orange, Oriental persimmon, cucumber etc.
● Stimulative or aitionomic parthenocarpy: Pollination or other stimulation
is required for parthenocarpy. This is termed as stimulative parthenocarpy
e.g banana because it is a triploid (it is the result of a diploid and a
tetraploid parent) and therefore cannot produce seeds e.g. Black Corinth
grape, Black berries, pear etc.
● Stenospermocarpy: Pollination and fertilization occur but the embryo
gets aborted i.e embryo abortion is the cause of seedlessness e.g grapes.
2. Artificial or induced: Induced parthenocarpy is the production of seedless
fruits by treatment of the flower with materials such as dead pollen (mentor),
pollen extract, chemicals or growth substances. Plant hormones like auxins,
gibberellins and cytokinins when sprayed on flowers always stimulate the
development of parthenocarpic fruits. The auxin required for the development
of fruit, normally supplied by fertilized ovules, may also be occasionally
raised above the threshold level required for fruit development by high
temperature, frost, insect attack, bark ringing and by mechanical irritation to
stigma and style. Osborne and Went (1953) were able to induce parthenocarpy
with low temperature and high light intensities in tomato. Nitsch (1952)
induced parthenocarpy in cucumbers at short photoperiod and with low night
temperatures. Frost and low temperature also result in production of
parthenocarpic fruits e.g apple and pear.
Insect attack produces small seedless fruits in some apple varieties. By bark
ringing, parthenocarpic fruits have been produced in gooseberry, grapes and apples.
Cutting of style from base before the pollen tube reaches the ovary induces
seedlessness in several citrus fruits. Chemicals can be used for providing the
206 Fruit Tree Physiology

stimulus for the development of mature fruits. Auxins like indole acetic acid,
indole propionic acid, indole butyric acid and phenoxy acetic acid have been used
to induce parthenocarpy in fruit crops.
The induction of parthenocarpy, however, results in several changes in the
physico-chemical characteristics in the fruits e.g
1. Size: In mango seedless fruits had smaller size than seeded fruits, but had
good quality and matured earlier than seeded fruits. In grape also the size of
seedless berries was smaller than seeded ones of the same variety.
2. Form: Parthenocarpic fruits (induced by GA) in apple were usually more
elongated than normal seeded fruits.
3. Composition and quality: In most grape varieties, seedless fruits are sweeter
than seeded fruits of the same variety.
4. Maturity: Seedless fruits ripen later than seeded fruits which may be attributed
to ethylene produced by seeds.

Importance/ advantages of parthenocarpy

1. It is a desirable trait in edible fruits with hard seeds such as pineapple,
banana, orange and grapefruit.
2. Desirable in fruit crops that may be difficult to pollinate or fertilize, such as
tomato and summer squash.
3. Parthenocarpy increases fruit production in dioecious species, such as
persimmon, because staminate trees do not need to be planted to provide
pollen.
4. Parthenocarpy also increases the novelty of fruits like cherry, grapes,
strawberry – they are easier to eat with less waste.
5. Improves taste and palatability.
6. Improves processing quality and ultimately increases the profitability in many
fruit crops.
7. Increases shelf life due to reduced ethylene generated by seeds.
8. Increases labour efficiency in food processing industry.
9. Fruit set and production is less affected by environmental factors which are
adverse for pollination and fertilization.
10. In parthenocarpic plants, fruit set and growth often starts before anthesis and
it may allow early fruit production and harvest.
11. Parthenocarpy is a valuable trait to be used to improve both fruit quality and
productivity in horticultural species grown for the value of their fruit.
 Parthenocarpy – A Mechanism for Seedlessness 207

Undesirable effects:
1. Parthencarpy is undesirable in nut crops, such as pistachio, walnut, almond,
where the seed is the edible part.
2. Parthenocarpy occasionally occurs as a mutation in nature and is considered
as a defect because the plant can no longer sexually reproduce.

Origin of parthenocarpy
1. Due to absence of pollination or fertilization.
2. Due to male sterlity.
3. Degradation of pollen mother cell.
4. Degradation of fertilized ovules.
5. Due to chromosomal irregularities during meiosis leading to triploidy
(seedless) e.g Tahiti lime, Orobalanco.
6. Due to self incompatibility e.g citrus.
7. Due to embryo abortion e.g grape.

PHYSIOLOGICAL BASIS OF PARTHENOCARPY

The physiological basis of parthenocarpy is still not fully understood. It occurs
most commonly among fruits with large number of ovules, suggesting that ovule
may provide some of chemical constituents which stimulate fruit set and fruit
growth. The ovules are sites of auxin synthesis in some fruits (Nitsch, 1952) and
the fact that auxin application can bring about fruit set enhances the possibility
that auxin are critical in fruit set. However, many fruits do not respond to auxins,
and so, there must be further component of fruit set stimulus in plant. It has been
observed that the factors favouring fruit set come from the roots of fruit trees may
be interpreted and imply a role of gibberellins and cytokinins. A relationship
between auxin content and the natural inclination towards seedlessness was
established by Gustafson (1942) while examining the ovaries of seeded and seedless
species and found that the seedless varieties have appreciably higher auxin content
than the seeded varieties in citrus fruits and grapes and parthenocarpy may represent
a state of auxin autotrophy in the ovary (Luckwill, 1957). Crane (1965) was able
to induce parthenocarpy with auxin, gibberellin or cytokinin and suggested their
role in triggering or mobilization of activities in the fruit.
To obtain parthenocarpic fruits is a physiological challenge. Fruit development
comprises early development and maturation (Fig.). Early fruit development in
most of the plants involves three phases: (i) fruit setting (ii) cell division and
(iii) cell expansion. The ovary takes the decision to either abort or to go further in
fruit development during first phase. During cell division phase growth of fruit
takes place, however, the increase in fruit size is low because the dividing cells are
small and tightly compressed and the number of cells decides the final fruit size.
The third and last phase begins after cell division ceases, the fruit grows by the
208 Fruit Tree Physiology

increase in cell volume, until it reaches its final size. Cell expansion commonly
increases fruit size by 100-fold, and this makes the greatest contribution to the
final size of the fruit. At the end of early development, a green fruit is obtained,
which has the size of a mature fruit, and the maturation phase occurs from this
point onwards. During early fruit development, many pathways of communication
between the sporophyte and the gametophyte are established. The decision of
whether or not to set fruit is dependent on the successful completion of pollination
and fertilization. The pollen produces gibberellins and exogenous gibberellins
application can induce an increase in auxin content in the ovary of an unpollinated
flower, and therefore trigger fruit setting in the absence of fertilization and the
developing embryo controls the rate of cell division in the surrounding fruit tissue
(Gillaspy et al; 1993). The number of developing seeds influences the final size
and weight of a fruit (Nitsch, 1970) as they promote cell expansion by the production
of auxin and other unknown molecules within the fruit. Therefore, a developing
seed has a very important role to play in the early development of fruit and hence
producing parthenocarpic fruits is a complicated phenonmenon. But seedlessness
is not uncommon because seedless crop plants have existed for many centuries.
Development of parthenocarpic fruit: Three aspects are important in

Fig. : Schematic representation of the links existing between fruit and seed
development: (a) different steps in fruit development, (b) different steps in
seed development, and (c) examples of seedless fruit. Arrows drawn
between (a) and (b) indicate the positive effect on fruit development of
events preceding, or linked to, seed development. Arrows between (b) and
(c) indicate the points in seed formation that might be deficient in seedless
varieties (Varoquaux et al; 2000).
 Parthenocarpy – A Mechanism for Seedlessness 209

development of parthenocarpic fruit. These are:

I. Development of the ovary wall: While comparing fruit development in
pollinated and parthenocarpic American holly fruits, Gardner and Marth (1939)
observed the parthenocarpic fruit set by indole acetic acid, could not be
distinguished from pollinated fruit at the time of maturity and ripening on
the basis of external appearance. The whole pistillate structure was found
similar in both pollinated and parthenocarpic fruits through microscopic
examination during early development. However, there was lack of
disorganized cell growth such as has been observed after applying indole
acetic acid to bean plants (Kraus, 1936). No unregulated cell division, tumerous
out growths occurs in holly pistils when sprayed with aqueous solutions of
this growth substance. There were no differences in color, texture, taste, and
chemical make up of the carpel wall between seedless fruit set by growth
substance and that of pollinated fruit.
2. Seed coat development: In most fruits set by growth substances, seed coat
development is much more variable while carpel or ovary wall development
is normal or nearly so. The plants in which fruits have been set by growth
substances, in 80 per cent or more the seed coat do not develop. It is not
known why seed coats develop in some fruits and not in others. There is
considerable variability among closely related plants. Wong (1939) reported
that Early Prolific Straight neck contained empty seed coat while Dark Green
Zucchini had no seed coat at all. Wong (1939) has reported stimulated set of
a perfectly seedless water melon, while Howlett and Marth (1946) stated that
hollow seeds almost as large as seeds would normally be present in all
parthenocarpic fruits he examined.
3. Embryo development: No embryos have been observed in fruits set by growth
substances and it is not surprising that embryos do not develop, since in the
absence of pollen, the egg is not fertilized. However, among plants, many
instances of embryo development in the absence of fertilization of egg are
known intimation of embryos by growth substances in the absence of
pollination has not been possible. From horticultural point of view, it is
evident that in those crops where the seed (embryo) is used no yield benefit
can come from using growth substances as a substitute for pollination e.g
growth abscission in dry cherries, though it is not clear that abscession is a
primary cause of poor fruit set or not.
Factors affecting parthenocarpy
The factors which influence or favor the parthenocarpy in fruit crops are:
1. Environmental factors (high or low temperature).
2. Growth regulators.
210 Fruit Tree Physiology

3. Genetic disorder (chromosome imbalance).

4. Self-incompatibility.
5. Genetic factors like gene controlling meiosis.
1. Environmental factors: Environmental factors such as light, temperature,
relative humidity affect pollen maturation and fertilization. Unfavourable
environmental conditions have a negative impact on fertilization and fruit
development. However, parthencarpic plants can overcome this problem by
allowing seedless fruit development also under these unfavourable
environmental conditions. Parthenocarpy can be exploited for increasing winter
and early production of fruits (Acciarri et al; 2002) ensuring consumers to
find fresh horticultural products year round. The difficulty of simulating the
environment that allows maximum expression of parthenocarpic traits is a
major problem with facultative parthenocarpy (a genetically controlled
condition causing development of seeded and/ or seedless fruits on the same
plants that is influenced by environmental conditions). The degree of
facultative parthenocarpy is influenced by all abiotic factors that affect the
pollination, fecundity and fruit development processes and the maximum
temperature i.e > 300C reduce pollen viability in most of the crops, thereby
increasing the degree of facultative parthenocarpy. The facultative character
of parthenocarpy allows reproduction by seed of interesting material and the
development of hybrids. In several facultative parthenocarpic tomato
(Lycopersicon esculentum Mill.) lines, environmental factors affect expression
of parthenocarpy e.g Oregon T5-4-a parthenocarpic line sets parthenocarpic
fruit only when temperatures is below 18°C. The Italian Shapat tomato
produces seeded or parthenocarpic fruit depending on the temperature and
pollination method. Parthenocarpy is useful in breeding of tomato so as to
adapt to a wide range of temperatures which are normally unfavourable for
fruit setting. The greatest degree of parthenocarpy in tomato cv. Oregon
Pride, leading to seedlessness has been found to occur above a daily mean
temperature of 18-21°C, with a very low number of seeds produced.
2. Growth regulators: Auxins, gibberellins and cytokinins or mixtures of these
hormones have been found to be effective in inducing fruit development in
the absence of fertilization in several crop species (Tiziana Pandolfini, 2009).
The process of fruit development is controlled by many biochemical and
genetic factors. Many studies have shown that phytohormones play important
role in fruit set regulation at the transcriptomic level, growth and maturation
in a complex way.
The growth regulators such as synthetic auxin can be used to induce
parthenocarpy. However, this increases the production costs and also causes
other fruit defects. The drawback associated with these effects can be overcome
 Parthenocarpy – A Mechanism for Seedlessness 211

through exploitation of genetic parthenocarpy trait that is being currently

introgressed into several crops e.g banana, (Musa accuminata Colla (AA) X
Musa balbisiana Colla (BB), grapes, watermelon and cucumber. Genetic
parthenocarpy can also be artificially established with exogenous genes,
controlled by ovary or ovule specific promoters that derive the over
sensitization to, or accumulation of auxin in carpel tissue before anthesis
(Carmi, 2003).
In grapes, to obtain well developed berries, GA is applied. It causes
cellular expansion in mesocarp tissue thus increases berry size and reduces
the seed traces. The optimum concentration and timing of GA application is
cultivar dependent.
In cucumber, tomato and peas, the parthenocarpic fruit set is associated
with the endogenous auxin and gibberellin in the ovary. Exogenous application
of auxin, GAs and cytokinins has been shown to induce parthenocarpy with
various plant species responding differently to auxin and GAs. High levels
of endogenous auxin and GAs occur naturally in parthenocarpic citrus and
tomato cultivars. GA causes cellular expansion with restricted cellular division,
and auxin causes cell division. Application of sprays of a-naphthalene acetic
acid to pineapple plants after normal differentiation of floral primordia had
completely influenced the growth of fruit, peduncle, and axillary branches.
Relatively high concentrations of the growth-regulating substances caused a
marked increase in size and weight of ripe fruit, although the number of
fruitlets was not influenced by any of the treatments. This increase in growth
of the fruit was accompanied by the development of large peduncles from
which the fruit was separated with some difficulty. The highest concentration
of naphthalene acetic acid also delayed ripening, and, particularly when
applied in the center of the plant, partially inhibited growth of slips and
suckers. Relatively low concentrations applied at the same time caused some
stimulation of the growth of dormant slip buds, resulting in a larger number
of slips per plant. The ovaries of parthenocarpic fruit have higher content of
auxin than corresponding non-parthenocarpic cultivars. Perhaps this is
responsible for their development into fruits without the necessary pollination
and fertilization. The ovules of seeded fruits hardly produce auxin before
fertilization. Hence the auxin content of the ovary do not reach to optimum
level. Therefore it does not develop into fruits. It is logical to think that the
ovary should develop if this natural defeciency is made up with external
application. In many plants a single application of low auxin will suffice to
induce ovary to develop into fruit. It is rather doubtful if this external auxin
is stable and sufficient for the development of fruits lasting over a few weeks
to few months depending upon the species. It may be recalled that in the
normally fertilized ovary, auxin production in developing seeds continue for
212 Fruit Tree Physiology

almost the entire period of fruit development. So, it can be concluded that
wherever the single application of auxin induce parthenocarpy, growth of
some tissues of unfertilized ovules such as the nucellus or integuments are
stimulated and these tissues probably continue to produce auxin. However,
native auxin so produced by the ovular tissues can not be the same as that
of synthetic auxin, which was initially employed for inducing parthenocarpy.
Now it is known that auxin alone can not induce parthenocarpy, therefore it
is natural that the ovaries of only such plants which are deficient in any such
hormone will respond to external application. Depending upon the fruit crop
and cultivar the combination of one or more hormones are responsible for
inducing parthenocarpy.
Auxins: Auxin induces parthenocarpy and the stimulus leading to fruit set
may not only be from the pollen but also from the ovary. Pollination stimulates
auxin synthesis in the ovary and unpollinated ovaries have only small amount
of auxin. Maximum auxin production is at the tip first followed by style and
finally in ovary having the maximum content. High auxin content associated
with natural parthenocarpic fruit development is reported from some citrus
varieties and other horticultural species. Synthetic auxin like growth substances
are used for the production of parthenocarpy fruits artificially. Every kind of
tissue (except the embryo which may be empty or filled with undifferentiated
cells) irrespective of origin or composition develops in parthenocarpic fruits
in a way exactly similar to the development of several fruits following
pollination viz. Anana sativus (swallon receptables), Ficus carica, Prunus,
Malus etc. (receptacles). Following fertlization that provides required
stimulation to the ovary and the surrounding tissues to continue growth, cell
enlargement and cell differentiation lead to maturity of the fruit. Auxin
regulates almost all developmental processes in plants including fruit
development. Since natural and artificial auxins supplied exogenously to
unpollinated flowers induce fruit growth in horticulttural plants thus replacing
the signals provided by pollination and fertilization (Nitsch, 1952).
Gibberellins: GA1 or GA3 induce fruit set in several horticultural species
(Gillapsy, 1993., Dorcey et al; 2009). In pollinated ovaries increased levels
of gibberellin together with an increased expression of GA biosynthetic genes
have been observed (Serrani et al; 2007). They act as potent fruit setting
agents. Gibberellins induce parthenocarpy and sustain fruit growth to maturity
in apple (Molesini et al; 2009). This pronounced fruit-setting activity and the
established observation that asymmetric growth of apple fruit is related to
incomplete seed development, has encouraged investigations on the
relationship of endogenous GA in endosperm and it was found that immature
apple seed contain GA4 and GA7. Role of gibberellins in growth and
development of parthenocarpic fruit is not clear, however, GA4 and GA7 are
 Parthenocarpy – A Mechanism for Seedlessness 213

more active than GA7 in inducing parthenocarpy and have been found in
seeds of immature apple fruit (Wang et al; 2009). It is not known to what
extent these gibberellins occur in other fruit tissue and whether or not they
participate in fruit growth.
Gibberellin has been found to induce parthenocarpy in many fruit species.
Crane et al.(1964) demonstrated the first success of gibberellin induced
parthenocarpy in Prunus species and found that application of potassium salt
of GA3 in aqueous solution induced parthenocarpic fruit more effectively in
peach than in almond and apricot, but not in cherry and plum.
Effects of applied gibberellins (GAs), GA1, GA3, GA4 and GA7 with a
cytokinin, N-(2-chloro-4-pyridyl)-N-phenylurea (CPPU) and indole-3-acetic
acid (IAA) were studied by Caixi et al. (2008) on fruit set, parthenogenesis
induction and fruit expansion of a number of Rosaceae species like Japanese
pear cv. Akibae (self-compatible) and cv. Iwate Yamanashi (a seedless cultivar).
Pyrus communis, Chaenomeles sinensis, Cydonia oblonga, and Malus pumila
were also investigated. It was observed that GA4, GA7 and CPPU are very
effective in inducing parthenocarpic fruit growth, whereas GA1, GA3 and
IAA have no ability to induce parthenogenesis in Japanese pear. GA4 and
GA7 induced parthenocarpic fruits were smaller in size, higher in flesh
hardiness and showed advanced fruit ripening in comparison to pollinated
fruit and to parthenocarpic fruit induced by CPPU. Increased pedicel length,
fruit shape index and a slight protrusion of the calyx end were found in the
fruits induced by GA4 and GA7. Parthenogenesis was also induced by CPPU,
GA4 and GA7 either alone or in combination with uniconazole in three other
Rosaceae species, although final fruit set was extremely low. GA1 was
essentially inactive in promoting fruit expansion unlike the other bioactive
GAs, and effectiveness in promoting fruit cell expansion varied as GA4 and
GA7>GA3>GA1.
Cytokinins: Cytokinins promote fruit growth in many fruit crops. Application
of cytokinins to flowers before fertilization triggers the onset of fruit growth
in some species. Cytokinins, auxins and GAs accumulate after fertilization.
Since, cytokinins regulate cell division, they are associated with the first
phase of fruit growth which is characterized by a marked increase in cells
number. A correlation has been observed between cytokinins levels and cell
division activities in many horticulture fruits (Srivastava and Honda, 2005).
Brassinosteroids: Brassinosteroids (BRs) are steroid hormones that play a
prominent role in many plant developmental processes including fruit growth
and developement. BRs have promoting effects on plant growth and show
synergism with auxins. Fu et al. (2008) observed parthenocarpic growth in
cucumber accompanied by active cell division by BR application, whereas
the application of an inihibitor of BR biosynthesis blocked fruit growth in
214 Fruit Tree Physiology

a parthenocarpic cultivar and attributed the promoting effect on cucumber

fruit growth to the induction of cell cycle related genes by BRs. Pandolfini
et al. (2009) produced seedless fruits in cucumber by BR application and
these were similar to fruits obtained from pollinated flowers.
Phenylurea (CPPU): The effects of naphthaleneacetic acid (NAA), gibberellic
acid (GA 3 ), N-(2-chloro-4-pyridyl)-N2 -phenylurea (CPPU), 6-
benzylaminopurine (BA), N, N2 -diphenylurea (DPU) and N-(4-pyridyl)-N2
-phenylurea (4-PU) on fruit set in unpollinated and pollinated ovaries of
Chinese white flowered gourd (Lagenaria leucantha) showed that only CPPU
induced parthenocarpy and all the parthenocarpic fruits developed to normal
size. The parthenocarpy percentage induced by CPPU was as high as 100 per
cent within the tested concentration (10–100 mg/l) and was not significantly
altered by application time from two days before anthesis to two days after
anthesis. CPPU also increased fruit set and fruit growth of pollinated ovaries.
Treating all ovaries on a plant or alternate ovaries increased fruit yield by
91.3 per cent and 83.3 per cent, respectively, but reduced fruit size
(Yu, 1999).
3. Genetic disorder (chromosome imbalance): Plant species which are triploids
cannot successfully produce seeds and they are genetically sterile e.g banana
is parthenocarpic because it is a sterile triploid (two sets of chromosomes
from one parent and one set from the other) instead of the normal diploid
(one set of chromosomes from each parent). Pollination occurs but fertilization
does not take place. The tiny black dots inside the banana are traces of the
unfertilized ovules. In the case of seedless watermelon, triploidy is induced
through genetic manipulation. Triploid watermelons (with three haploid sets
of chromosomes) are unable to produce viable gametes during meiosis, and
melons are seedless. They are produced by crossing a tetraploid (4n) seed
parent with a diploid (2n) pollen parent bearing haploid (n) sperm. The
haploid (n) sperm from a polle grain from the male flower of the 2n parent
fertilizes the diploid (2n) egg inside the ovule of a female flower on the 4n
parent. The resulting 3n zygote develops into a 3n embryo inside a seed.
Planting this seed produces 3n seedless watermelons. Co.I, a triploid banana
was developed by multiple crossing at TNAU, Coimbatore from Laden (AAB)
× M. balbisiana (BB) → F1 (AB) × Kadali (AA) → Co.I (AAB).
Coupling of parthenocarpy with male sterllity is another approach for
producing seedlessness. Sterillty can be induced by increasing ploidy level
or cytoplasmic manipulations e.g citrus. Ploidy manipulation using crosses
between diploid and tetraploid yield many suitable triploids including
Oroblanco and Melogold (C. grandis X C. paradisi) and Winola (Spiegel
Roy and Vardi, 1992).
 Parthenocarpy – A Mechanism for Seedlessness 215

4. Self incompatibility: Self incompatibility impedes seed formation; several

cultivars grown in single plantings produce seedless fruit because of self
incompatibility. However, mixed cultivation of self incompatibility with male
sterile ones yields seeded fruit unless they exhibit female sterility e.g barberry
(Ebadi et al; 2010). Gametophytic self-incompatibility (SI) has been observed
to produce seedlessness in mandarin cv. Wuzishatangju by blocking
fertilization in the ovary by Ye et al. (2009).

GENETICS OF PARTHENOCARPY
High levels of auxins and gibberellins are present in the ovaries of parthenocarpic
fruits, and it has been proposed that genes for parthenocarpy might affect hormone
production, transport and/or metabolism in order to promote ovary growth
precociously. Thus, pollination and fertilization are no longer needed for their
growth (Muller, 1998). Several parthenocarpic mutants like pat mutant of the
tomato have been studied and it was found that these mutants produce
parthenocarpic fruits, but the genes involved show some pleiotropic effects, such
as male and female sterility, as a result of some floral developmental aberrations.
Some deficiencies in cell elongation in different organs (short anthers, smaller-
sized seeds and fruits, undersized integuments) in thgese mutants suggests that
the pat gene product interact with gibberellin metabolism. The application of
gibberellin to flowers causes the restoration of a wild type anther phenotype, but
does not restore female fertility. Another indication of the link between genes
involved in parthenocarpy and gibberellin metabolism is found in the
parthenocarpic mutant of Arabidopsis thaliana called SPINDLY (SPY), whose
gene product is anticipated to participate in the regulation of the gibberellins
signal transduction pathway (Jacobsen et al; 1993., Vivian et al; 1999).
The genetics of stenospermocarpy have been well studied in grapevines in
which seedlessness is controlled by three complementary recessive genes: a1, a2
and a3, independently inherited and regulated by a dominant gene I 25. The
importance of the contribution of the gene I to grape stenospermocarpy was
confirmed by RAPD markers. Phytohormones like auxins and GA plays an
important role in parthenocarpic fruit development. Increased level of these
hormones in ovary and ovule can substitute for pollination and can trigger fruit
development and this has been used for development of parthenocarpy through
genetic engineering (Goetz et al; 2006). Yin (2006) obtained seedless fruits by
elevating the auxin level in ovules of transgenic brinjal, tobacco and cucumber
plants through expression of iaaH gene from Pseudomonas syringae under control
of ovule specific promoter gene DefH9 from Antirrhinum majus. In future, cloning
of the corresponding genes might open up new avenues for manipulating
parthenocarpy. In addition, the study of parthenocarpic mutants in A. thaliana is
expected to contribute additional genes.
216 Fruit Tree Physiology

Production of parthenocarpic fruits via transgenics: Recombinant DNA

technology can be used to induce parthenocarpy in fruits. Parthenocarpy is
positively correlated with the level of auxin in the ovary, and that the exogenous
application of auxin (Homan, 1964), gibberellins, cytokinins (Choudhury and
Phatak, 1965) and auxin transport inhibitors (Robinson, 1971) to flowers helps to
induce parthenocarpy. The application of these hormones causes an increase in
the auxin content of the ovary in cucumber (Kim, 1992). High content of auxin
has been discovered in parthenocarpic ovaries of the tomato. Thus, it is theoretically
possible to induce parthenocarpy in transgenic plants expressing an auxin or
cytokinin biosynthetic gene in the ovary, between anthesis and early fruit
development. Various transgenic strategies can be employed to produce
parthenocarpic fruits as under:
1. Agrobacterium RolB gene, interfering with plant auxin production.
2. Fusion between pDef H9, an Antirrhinum majus promoter that is specific to
the ovary, and the RolB gene (to obtain parthenocarpy) or a cytotoxic gene
(to obtain female sterility).
3. Promoters specific to the ovary or developing fruit like pGH3, pAGL and
pPLE36 encoding an isopentenyl transferase (cytokinin biosynthesis) or a
tryptophan oxygenase (auxin precursor biosynthesis).
Thus parthenocarpy can be induced by expressing a gene encoding a step in
the auxin biosynthetic pathway in the ovary. However, the flowers must be
emasculated in order to produce seedless fruits. This system must be coupled with
male sterility for ease of use. Further, this system must be introduced into F1
hybrid seed production strategy in order to obtain viable hybrid F1 seeds that will
germinate and give rise to seedless plants.

Preventing the development of F2 seeds

Production of cytotoxic effect by combining two independently harmless genes:
In this two individually harmless genes when combined, are cytotoxic in tissues.
An endogenous molecule is converted into a non toxic molecule by the product
produced by first gene, which in turn can be converted into a cytotoxic molecule
by the product of the second gene e.g overproduction of auxin in the seed coat
using IamS gene, whose product converts endogenous tryptophan to indole
acetamine, and the IamH gene, whose product converts indole acetamine to indole
acetic acid (Fig.). IamS and IamH are obtained from Agrobacterium tumefaciens.
F1 seeds produced through this strategy are shown in Fig. F1 seeds obtained are
viable because only one transgene (that of the mother plant) is present in the seed
coat and plants grow normally and carry both transgenes. Selfing or crossing F1
leads to the expression of both genes in the seed coat which in turn is destroyed
by the over expression of auxin ultimately causing seed abortion.
 Parthenocarpy – A Mechanism for Seedlessness 217

trends in Biotechnology
Fig. : (a) Production of a cytotoxic compound following the introduction of two
independent genes encoding Iam synthase and Iam hydrolase into plants.
(b) Production of seedless fruits in the F2 generation using the Iam synthase
and Iam hydrolase system. The seed coat is a maternal tissue; the genotype
of maternal tissue is thus different from the other parts of the seed.
Abbreviations: Ho- homozygous; He- heterozygous; Pseed coat- seed-coat-
specific promoter (Varoquaux et al; 2000).

Recombination of DNA at specific sites: In this strategy site specific

recombination system from bacteriophage P1 is used which consists of a
recombinase (Cre) and recombination sites (loxP). Recombination between lox
sites occurs on supercoiled, nicked, circular or linear DNA in the presence of Cre
(Abremski, 1983). The Cre-lox system is efficient in many organisms particularly
in plants e.g Cre-lox systems can be used to make a site-specific insertion, as a
218 Fruit Tree Physiology

meganuclease, to induce translocation or to activate genes (Odell, 1994). Gene

activation by this system is described in Fig. Production of seedless fruits through
activation of genes by this system is shown in Fig.

Fig. : (a) Gene activation using the Cre-lox system. The lox-lox DNA fragment
prevents barnase expression. To activate the barnase gene, the Cre
recombinase excises the lox-lox DNA fragment. (b) Activation of a cytotoxic
gene, such as barnase, inducing seedlessness in the F2 generation by
specific expression in the seed coat (maternal tissue). Abbreviations: Ho-
homozygous; He- heterozygous; Pseed coat- seed-coat-specific promoter
(Varoquaux et al; 2000).
Table : Examples of transgenes inducing parthenocarpy

Crop Gene Genetic modification Function Fruit phenotype Reference

Tomato, DeH9-iaaM Ovule specific transgene Auxin synthesis Obligate and Mezzetti et al;
tobacco, expression facultative parthenocarpy, 2005
cucumber, early fruit growth, normal size
brinjal and shape.
Tomato rolB Ovule/ fruit specific Auxin synthesis Obligate and facultative Carmi et al;
transgene expression. parthenocarpy, early fruit growth, 2003
normal size and shape, similar
in weight.
SIIAA9 Antisence down regulation Auxin signalling Parthenocarpy, early fruit growth, Wang et al;
of constitutive promoter normal size and shape. 2005
SlARF7 RNAi-mediated silencing, Auxin signalling Parthenocarpy, seedless, De Jong et al;
constitutive promoter. altered shape 2009
SlAUCSIA RNAi mediated silencing, Auxin response Obligate and facultative Molesini et al;
phloem-specific. parthenocarpy, seedless, 2009
reduced size.
SiChs RNAi mediated silencing, Auxin transport Parthenocarpy, seedless, Schijilen et al;
constitutive promoter. altered size, shape and colour. 2007
SlDELLA Antisence down regulation, Giberellin signaling. Facultative parthenocarpy,
constitutive promoter. seedless, reduced size, altered Marti C et al;
morphology. 2008
SlTPRI Over expression. Ethylene signaling. Parthenocarpy, seedless, reduced Lin et al; 2008
size, altered morphology.
Arbidopsis, AtARF8 Expression of mutant Auxin signalling Parthenocarpy, pseudo embroys, Goetz et al;
tomato AtAFR8-4gene. size and shape similar to weight. 2007
Parthenocarpy – A Mechanism for Seedlessness 219
220 Fruit Tree Physiology

Biotechnological approaches to induce parthenocarpy:

1. Due to induction of male sterility.
2. Seed coat destruction by suicide gene expression.
3. Induction of female sterility- destruction of ovules or stigma by suicide
genes.
4. Increased expression or sensitivity to GA in ovary.

Causes and induction of parthenocarpy in fruit crops

1. Guava: e.g Allahabad Seedless
a. Due to triploidy.
b. Failure of fertilization of well differentiated ovules.
c. Degeneration of fertilized ovules.
2. Grape: e.g Thomposon Seedless, Flame Seedless, Crimson Seedless.
Due to embryo abortion (stenospermocarpy).
3. Embryo rescue technique is used to produce seedless grapes. In this technique
the tiny embroys (immature) are removed and grown artificially through
tissue culture.
4. Citrus:
a. Due to lack of pollination.
b. Self incompatibility (mandarins).
c. Cytoplasmic male sterlity (Satsuma).
d. Chromosomal irregularities (Tahiti lime, Oroblanco)
5. Apple - Lack of pollination.
Induction of parthenocarpy in fruits: Application of synthetic auxin can
bring about the development of an unpollinated ovary into a fruit (Gustafson,
1936). He found that application of synthetic auxin directly stimulate the growth
of ovules which are necessary in the development of parthenocarpic fruits. Auxin
sprays usually cause seed abortion in apple and pear.
Auxins and Gibberellins have been successfully used in citrus, grapes and
pineapple to induce parthenocarpy at low concentrations.
Apple: Mixture of IBA and NAA (5 ppm each) applied as a water spray to
young fruitlets damaged by frost produced 100 per cent parthenocarpic fruits in
Miller’s seedling (Smarbrick; 1934). Parthenocarpic fruits of cv.Cox’s Orange
Pippin were obtained by Goldwin and Sehwabe (1975) through spraying a mixture
 Parthenocarpy – A Mechanism for Seedlessness 221

of GA3 4 DPU (diphenyl urea) + Noxa ( b- naphthoxy acetic acid) @ 300 + 300
+ 40 mg/l. Fortes and Petri (1983) obtained prthenocarpic fruits in Golden delicious
by using wye mixture (NOXA 500 + GA3 200 + DPU 300 ppm) irrespective of
time of application (petal fall, full bloom or 50% bloom).
The effectiveness of gibberllins GA3 and GA4 in inducing and sustaining
parthenocarpic growth of apple fruit demonstrates the importance of growth factors
other than auxins in fruit development. In several cultivars of Malus sylvestris,
the effect of GA3 and GA4, Chloro 4 fluorophenoxy acetic acid and l-naphthyl-
N-methyl carbamate on the induction of fruit growth, in the absence of pollination
and fertilization, were determined (Martin, 1963). Bangerth and Sohroder (1994)
reported that CPPU and gibberllins (GA3 and GA7) had a positive synergistic
effect on parthenocarpic fruit production in apple cv. Golden Delicious and
Jonagold. An increased parthenocarpic fruit induction and retention was obtained
by spraying of CPPU in combination with GA3 in apple cvs Ohrin and Fuji
(Watanabe et al; 2008).
Cherries: Application of gibberllin in conjuction with 2,4-dichlorophenoxy-
acetyl methionine resulted in parthenocarpic development of fruits of sweet cherry.
When both compounds were applied on fruits of the self incompatible Bing
cherry, a parthenocarpic set of 27-39 per cent was obtained (Rebeiz and Crane,
1961). GA3 200 to 250 ppm in combination with auxins such as 2, 4-D, 4 CPA,
2,4, 5-T, NAA and pichloram to Sour cherry resulted in 20-21 per cent
parthenocarpic fruit set. Application of GA3 at 200 ppm on Early Rivers produced
54-79 per cent seedless fruits. Treatments of NAA @ 100 ppm + GA3 @ 5-25
ppm in sweet cherry increased parthenocarpic fruit set. Bukovae et al. (1985)
demonstrated that an N substituted phathalimide induced parthenocarpic fruit
development in sour cherry and that this biological activity can be enhanced
markedly with NAA.
Citrus: There are also several classes of seedlessness in commercially available
citrus fruit ranging from self incompatible cultivars such as Clementine mandarin
that show a lower ability to set fruit in the absence of cross pollination to the
truely seedless Satsuma mandarins and Navel sweet oranges, that generally set a
normal crop of parthenocarpic fruit, due to high masculine and feminine gametic
sterility (Frost and Soost, 1968). In these varieties, parthenocarpic fruit develop
without formation of seeds and therefore all pollination, fertilization or seed
requirements for fruit growth activation have clearly been substituted by
endogenous signals. Self incompatible cultivars show a low degree of
parthenocarpy and therefore can be considered to possess facultative parthenocarpy
meaning that seedless fruit form only when fertilization does not occur. Elimination
222 Fruit Tree Physiology

of seed formation is a valuable trait for many citrus cultivars, especially the
mandarin varieties and seedy lemon varieties. Application of GA on the flowers
of grape fruit (Marsh Seeded, Excelsior and Duncan) and mandarin (Kaula, Lahore
Local and Nagpuri) at different concentrations produced parthenocarpic fruits
ranging from 40-49 per cent. At higher GA concentrations (500-1000 ppm) the
fruits were oblong almost pear shaped and had a very rough and thick skin.
Whereas, the appearance, colour and texture of fruits were not affected at lower
concentrations of GA (25-100 ppm) (Randhawa et al; 1964). The changes of
endogenous growth hormones in parthenocarpic ovaries of citrus were studied by
Talon et al. (1999) and found that seedless fruits of Satsuma and Clementine
contain higher amounts of GA.
Exogenous cytokinins have been reported to enhance parthenocarpic fruit
development and stimulate sink strength in developing fruits of certain cultivars
(Hernandez and Primo Millo, 1990). Parthenocarpic Satsuma was found to have
higher amounts of gibberllins than pollinated fruitlets after fruit set in citrus. The
addition of 25 mg L–1 of CuSO4·5H2O applied to Clementine mandarin flowers, at
the pre-anthesis stage and 2 h prior to pollination with Fortune mandarin pollen,
arrested pollen tube growth in the upper region of the style, and the average number
of seeds was reduced by 96 per cent per fruit. It also significantly reduced the
average number of seeds per fruit by 55–81 per cent and significantly increased the
percentage of seedless fruits, without reducing fruit yield when applied at full bloom
to Afourer tangor trees under cross pollination conditions (Mesejo et al; 2006).
Polyamines play important role in fruit development in several plants and three
genes encoding aminopropyl transferases have been isolated from citrus viz. CcSPDS,
CcSPM1 and CcACL5. The expression of these genes in C. clementina is not restricted
to ovaries and fruits, but it is also detectable throughout the plant. More importantly,
gibberellin-induced parthenocarpic fruit set caused a decrease in CcSPDS expression
in ovaries, paralleled by a decrease in spermidine, while the expression of CcSPM1
and CcACL5 was basically unaffected, resulting in the maintenance of spermine
concentration during early fruit development. In addition, the variation in putrescine
content was paralleled by changes in the expression of one of the two putative
CcODC paralogs (Trenor et al; 2010). Parthenocarpy can also be induced through
various biotechnological approaches (Dandekar, 2005) such as:
Through ovule specific regulation of auxin synthesis: The expression of an
auxin biosynthesis gene iaaM encoding a tryptophan monoxygenase from
Agrobacterium tumefaciens (At) regulated by an ovule-specific promoter, DefH9,
from Antirrhinum majus (Am) or an OS promoter from Arabidopsis thaliana
(AT) is used to induce parthenocarpy.
 Parthenocarpy – A Mechanism for Seedlessness 223

Through ovule specific modulation of auxin response: rolB gene from

Agrobacterium rhizogenes (Ar) regulated by the ovule specific promoters DefH9
or OS which results in modulation of auxin response. Expression of the DefH9-
rolB or OS-rolB specifically in the ovule results in the modulation of the sensitivity
of ovule to IAA such that the endogenous levels of IAA present in ovule will be
sufficient to induce parthenocarpy and the formation of seedless fruit.
Grapes: Application of PGRs in seeded cultivars of grapes produce seedless
fruits which often fetch higher market price. GA resulted in low proportion of
seeded berriers in grapes cv. Campbell Early, increased ratio and weight of seedless
fleshy berries of grapes cv. Delware (Kato et al; 2000). Parthenocarpy is also
induced by increasing ploidy level e.g Rosario Bianco produced parthenocarpic
fruits of about 4 g (Sarikhani and Wakana, 2005). Streptomycin (400 mg/ L)
applied to clusters six to eight days before bloom, followed by 30 mg/ L GA3
12 days after bloom, resulted in commercially acceptable clusters (476 g) and
100 percent seedless berries weighing 6.3 g in grape cv. Rubi (Pommer et al;
1996). GA (100 ppm) in combination with BA (100 or 200 ppm) and urea (0.5%)
resulted in induction and improved growth of seedless fruits in Delware grape
(Naito et al; 1974). GA application at150 or 300 ppm to Venus clusters 7 days
after flowering resulted in a reduction in the number of seed traces in mature
fruits while in Saturn, there was a great reduction in seed trace number after a
combined pre and post-flowering treatment. When sprayed with GA3 at 100, 200
or 300 mg/ litre on the leaves and fruit clusters of Muscadine grape cultivar
Triumph produced more than 20 per cent seedless berries (Lu et al; 1997).
Shiozaki et al. (1997) reported that development of seedless berries induced by
GA3 application follows a course of cell division and enlargement quite different
from that in seeded ones.
Guava: Application of 2, 4-D and NAA at 0.5 per cent produced parthenocarpic
fruits in guava. Commercial seedless varieties of guava are not entirely free from
seeds and contaion 9-20 seeds per fruit. Rao and Nagar (1971) reduced total
number of seeds in guava fruit when 10,000 ppm of potassium salts of GA was
applied to the emasculated flowers. According to Teotia et al; (1961) seedless
fruit produced by the application of 10,000 ppm GA to the emasculated flower
buds produced parthenocarpic fruits to a greater degree when dipped in solution
of 250 ppm GA as compared to the control. However, at higher concentration, the
parthenocarpic setting was increased.
Date palm: Parthenocarpic fruits were produced in cv. Barhee by the treatments
of different pollen grains boiling extracts or 500mg/ L glutamine by Abd El-
Zaher (2008). Application of three GA sprays, once at spathe opening and then
224 Fruit Tree Physiology

2 sprays (25-100 ppm) at 4-week interval followed by 500 ppm etherel spray
resulted in seedless fruits of normal size (Abdu Allal et al; 1982). Similarly,
Abou-Aziz et al. (1982) produced seedless berries by GA application at 50 or 100
ppm in dates. Tafazoli (1991) produced seedless dates by spraying unpollinated
clusters with GA3 (50 or 100 ppm) and beta naphthoxy acetic acid (NOA @ 50
or 100 ppm). However, the fruit number was reduced in the absence of pollination,
therefore, it was not considered to be economically feasible due to low yields.
Application of IAA, BA + GA and GA at 10 ppm on opened flower clusters of
3 cultivars (Sembelbel, Tallis and Aduri) induced the formation of seedless fruits
and the cultivar Sembel Bel and Tallis responded best with all growth regulators
(Hodairi et al; 1992). Spraying twice with 50 ppm GA + 10 ppm 2,4, 5-T
produced 70.20 percent seedless fruits in the Nebut Seif cv. whereas, the single
spray treatment with 50 ppm IAA produced 79.17 per cent seedless fruits in the
Sakaie cv (Shaheen et al; 1988).
Fig: The figs commonly grown in India are parthenocarpic in nature and do
not need any cross-pollination with fig (Capri fig), which is a very common
practice in other countries. It has been suggested that parthenocarpy is favoured
or inhibited in a given type by climatic conditions of the area in which they are
grown. Crane and Blandeau (1949) reported IBA (200-2670 ppm), NAA (25-250
ppm), 2, 4, 5-T (10-100 ppm) and 4-CPA (40-80 ppm) the most effective
compounds in inducing seedlessness in fig. Both auxin and GA can induce
parthenocarpic set in Calimyrna fig was reported by Crane (1965).
Litchi: The fluctuations of endogenous growth regulators level in the litchi
cultivars Hexiachuan and Huaizhi revealed that levels of GA and CK in
parthenocarpic fruits of Hexiachuan were higher than in cv. Huaizhi before and
after anthesis. The increase in IAA levels and the continuous decrease in ABA
levels during anthesis could help to explain parthenocarpy in cv. Hexiachuan.
Analysis of the growth regulators balance (ABA/ IAA + Gas + CK) showed that
Hexiachuan had a higher level of growth promoters than Huaizhi, which does not
exhibit natural parthenocarpy.
Loquat: Application of GA3 250 ppm after the emergence of floral buds or
NAA @ 20 ppm during full bloom was reported by Goubran and El-Zefturai
(1986) to produce seedless fruits. GA3+CPPU were found to be more effective
for seedless fruit production in the triploid loquat without decreasing the fruit
quality by (Yahata et al; 2006). The triploid seedless fruits were produced by
dipping flower and fruit clusters in an aqueous solution of 200 ppm GA3+20 ppm
CPPU twice, at flowering time (the first treatment) and between 27 to 58 days
after the first treatment (the second treatment). The parthenocarpic fruits had a
 Parthenocarpy – A Mechanism for Seedlessness 225

longer shape and higher fruit shape index (fruit length/ fruit width) than diploid
fruits with seeds (Yahata et al; 2006). Parthenocarpic fruit production was induced
by treating the trees three times with 100 mg/l of GA3 combined with ringining
which improved the fruit size and produced 26 kg, on average, of seedless fruits
of suitable commercial quality (Mesejo et al; 2010). Zhang et al. (1998) treated
the nine year old trees of loquat cultivars with different concentrations of CPPU
(Forchlorfenuron) and GA. Spraying GA before flowering induced the formation
of many parthenocarpic fruits. Spraying CPPU in mid February or GA after
flowering also induced parthenocarpy.
Mango: Parthenocarpy has been reported in some south Indian varieties like
Neelum, Banglora and Banganapalli (Venkataratuam, 1949). He removed the
staminate flowers and 20 to 30 bixexual flowers were retained in an inflorescence
after emasculating the single stamens. The emasculated flowers were sprayed
with 10 ppm IBA. The treated fruits looked like normal ones with well developed
mesocarp and endocarp but embryos were completely inhibited resulting in
parthenocarpic fruit development. Fully developed parthenocarpic fruits of mango
by spraying 250 ppm BA at the anthesis and latter by spraying a combination of
10 ppm NAA and 250 ppm GA3 were obtained and the fruits were small in size
but superior in quality to normal seeded fruits. Exposure to low temperatures (20/
10 °C day/ night) 3 days after hand pollination significantly increased the
percentage of stenospermocarpic fruit in which embryos were aborted at some
stage during early fruit development. Significant differences between cultivars
were observed in the percentage of seedless fruits produced, with 21 per cent for
Nam Dok Mai, 11 per cent for Kensington and 3 per cent for Irwin (Sukhvibul
et al; 2005). Shaban and Ibrahim (2009) observed that normal fruits contained
higher concentrations of endogenous hormones like gibberellins and cytokinins
and lower concentrations of auxins and abscissic acid compared to the
stenospermocarpic fruits resulting in slower growth rate and small size of seedless
fruits than the seeded ones, and most of these fruits dropped and fail to reach full
size. Parthenocarpic mango hybrid-117 was obtained by intensive backcrossing
between hybrid variety Ratna (Neelum x Alphonso) and Alphonso. The non
viable cotyledon free stone occupied only 3.1 per cent of the total fruit weight
and had a small degenerated ovule (1.12 g) inside the very soft endocarp (Gunjate
and Burondkar, 1993).
Peach: Spraying of GA at 250, 500 and 1000 ppm to emasculated J.H. Hale
peach blossoms promoted parthenocarpic fruit sets from 53.8 to 69.3 per cent.
The sets were significantly greater than that resulting from pollination. The
parthenocarpic fruits were smaller in diameter than pollinated ones but were
226 Fruit Tree Physiology

longer, contained longer pits, the flesh at stem and blossom ends were thicker,
and they matured a week or more earlier. The growth patterns of parthenocarpic
fruits were similar to that of pollinated fruits with seeds. It was concluded that
growth of the peach fruit is not controlled by auxins originating in the seeds
(Crane et al; 1961). Crane et al. (1960) induced parthenocarpy with GA treatment
in Prunus. Aqueous solutions of 50, 250 and 500 ppm GA applied as sprays to
almond, apricot, cherry, plum and peach. None of the treatments were effective
in promoting parthenocarpy in cherry and plum. GA at 500 ppm twice
brought about parthenocarpic sets of 11.8 per cent in the almond and 15.4 per
cent in the apricot. Parthenocarpy in the peach was induced by all concentrations
of GA.
Pear: Temperature during bloom is thought to be primary cause of
parthenocarpic fruit set. A temperature of 600F (15.60C) for about 72 hrs or more
for about 10 days period during bloom induce parthenocarpic set in pears (Elkins,
2003). The most effective compound is the sodium salt of alpha–(2-napthoxy)-
propionic acid applied at 100 ppm to promote parthenocarpy in pear (Osborne,
1949). A single spraying of emasculated flowers caused rapid initial growth, but
this ceased and the young fruits fell after about 4 weeks. Regular sprays of the
same compound at 3 days interval prevented this and in the variety Pitmaston
Duchess resulted in a 40 per cent set of fully grown parthenocarpic fruit. This
suggests that pear needs a sustained supply of hormone for full parthenocarpic
development. A single spraying of the variety Bartlett at pink bud stage with 100
ppm solutions of 2, 4, 5 – trichlorophenoxy propionic acid increased set of fruit
which were smaller and more apple shaped than normal, while in other varierties
(D. Anjou, Hardy and Winter Nelis), similar treatment caused a reduction in set,
illustrating the wide varietal differences in response to be expected from this fruit
was reported by Griggs and Iwakiri (1961). Improved fruit set with sprays of
GA3 or other GA’s stimulates parthenocarpic fruit development in pear cultivars.
The concentration of gibberllin in sprays found to be effective differs greatly with
the cultivars but in many instances the use of sprays is commercially worthwhile,
especially after frost damage. GA3 at 100 ppm to Bartlett pears resulted in
parthenocarpic fruit set but the fruits were elongated. Application of GA3 at 10,
20 or 40 ppm to Williams resulted in very high number of parthenocarpic fruits,
but these fruits were deformed (Bangerth et al; 1976).
Application of GA7 and CPPU are very effective in inducing parthenocarpic
fruit growth, whereas GA 1 , GA 3 and IAA, have no ability to induce
parthenogenesis in Japanese pear. GA4 and GA7 induced parthenocarpic fruit
tended to be smaller in size, higher in flesh hardness, and showed advanced fruit
 Parthenocarpy – A Mechanism for Seedlessness 227

ripening in comparison to pollinated fruit and to parthenocarpic fruit induced by

CPPU. GA4 and GA7 induced parthenocarpic fruit also had an increased pedicel
length and fruit shape index and also showed a slight protrusion of the calyx end
(Zhang et al; 2008). However, now it is well known that GA treatment enhanced
the proportion of parthenocarpic fruits in pear especially in cross pollinated tress
and interestingly GA4+7 showed more pronounced effect on self pollinated trees
and GA3 on cross pollinated trees. Application of exogenous GA4+7 is most
effective GA combination in inducing parthenocarpic fruits. GA4+7 (500 ppm) in
combination with or without other plant growth substances applied on the
emasculated and decapitated flowers at full bloom and 3 weeks after resulted in
parthenocarpic fruit roduction in Japenese pear (Yuda et al; 1983). Marcucci and
Visser (1983) observed that the parthenocarpic fruits had smaller core than the
normal fruits with abortive seeds and fruit diameter decreased with core diameter
due to which parthenocarpic pear fruits were more slender than normal fruits.

CONCLUSION
Parthenocarpy or seedlessness occurs both due to natural as well as artificial
means. Parthenocarpic fruits are usually but not always seedless and seedless
fruits are not always parthenocarpic because seedlessness may be caused due to
embryo abortion resulting due to inability of the embryo to accumulate the required
food reserves during its development. The physiological basis of parthenocarpy
is still not fully understood. It occurs most commonly among fruits with large
number of ovules, suggesting that ovule may provide some of chemical constituents
which stimulate fruit set and fruit growth. Several methods are employed for
inducing parthenocarpy like change in ploidy level, use of growth regulators, and
mutation etc. but due to one or the other reasons success is still limited. However,
transgenic technology has opened new windows in this aspect too and many
genes have been isolated that impart seedlessness in various crops like tomato
and arabidopsis. Through biotechnological methods seedless fruits are obtained
via i) male sterility, ii) seedcoat destruction to obtain stenospermocarpy through
expression of targeted suicide genes during early seed coat development iii)
female sterility and iv) increased expression or increased sensitivity to GA in the
ovary or ovule. Several factors like environmental, growth regulators particularly
GA, genetic disorders and self incompatibility have been observed to influence
the development of parthenocarpic fruits.
228 Fruit Tree Physiology

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 9

GRAFT INCOMPATIBILITY –
CAUSES AND REMEDIAL MEASURES

T he ability or success of two different plants grafted together to produce a

successfull union is called graft compatibility while the opposite condition
i.e inability or failure to produce successfull graft is called graft incompatibility.
The ability of two different plants to produce a compatible graft is due to their
natural relationship. There is no clear cut distinction between a compatible and
an incompatible graft union. The stock and scion of related species unite readily
and form a compatible graft union and grow as a composite plant while unrelated
species fail to do so. However, sometimes unrelated stock and scion initially
show compatibility but develop incompatibility later and die eventually. There
are also instances that lie in between these two extremes of compatibility and
incompatibility systems. Thus plants show wide variation in their ability to produce
a successful graft union. For instance some pear cultivars form compatible while
others form incompatible graft union on quince and may die soon. However,
reverse combination i.e quince (scion) on pear (stock) is always unsuccessfull.
Bartlett pear on quince root stock is incompatible and breaks off easily at union.
When Bartlett is grafted on oriental sand pear (stock), the union is physically
strong but fruits develop black end at calyx. Various structural events leading to
healing of grafts in plants are as (Pinna and Era, 2005):
1. Cambial regions of both stock and scion are in close proximity in order to
interconnect through the callus bridge due to formation of new parenchymatous
cells proliferating from both stock and scion producing the callus tissue that
soon intermingle and interlock, filling up the spaces between the two
components connecting the scion and the root stock.
 Graft incompatibility – Causes and Remedial Measures 235

2. Differentiation of new cambial cells from callus, forming a continuous cambial

connection between root stock and scion. Furthermore, prior to the binding
of vascular cambium across the callus bridge, initial xylem and phloem may
be differentiated. The wound-repair xylem is generally the first differentiated
tissue to bridge the graft union, followed by wound-repair phloem.
3. In the last step of the graft establishment process, the newly formed cambial
layer in the callus bridge begins typical cambial activity forming new vascular
tissues. Production of new xylem and phloem thus permits the vascular
connection between the scion and root stock.

Visual symptoms of incompatibility

● Failure of successful graft or bud union in high percentage.
● Low percent of scions that grow away (very low percentage of graft
success).
● Premature death.
● Declines in vegetative growth due to degeneration of tissues at graft union
and premature defoliation.
● Lack of overall growth of tree.
● Dieback of shoots, sometimes pale yellow colour.
● Marked difference in the vigor of the stock and scion.
● Deficiency symptoms or nutritional disorders.
● Stunted growth of plants.
● Excessive swelling at graft union.
● Breaking off of stock and scion at the graft union.
● Marked differences in growth rate of scion and stock.
● Development of over growth at, above or below the graft union resulting in
the formation of shoulder or inverted shoulder.
● Excessive suckering of root stock.
● Mechanical weakness between stock and scion with little pressure required
to break them away.
● A marked difference in bark patteren and structure of scion or root stock may
give an indication of incompatibility.
236 Fruit Tree Physiology

Types or classification of incompatibility

Based on length of time, incompatibility is of following types:
1. Immediate graft incompatibility: It arises when there is little or no attempt
by the scion and root stock to unite. Even though if union occurs it deteriorates
quickly.
2. Partially delayed graft incompatibility: When the union fails after a period
of 4-6 months to 3-5 years. It mostly occurs during the production stage in
nursery e.g Pyrus callyriana grafted on quince A.
3. Fully delayed graft incompatibility: In this, growth is normal till 15-20 years
e.g Bartlett pear on quince. There is development of shoulder (over growth of
root stock) or inverted shoulder (over growth of scion). However, over growth
of scion and stock always does not mean that the union is incompatible.
Another classification of graft incompatibility is as:
1) Localized incompatibility: Localized graft incompatibility is defined as
incompatibility that manifests when there is direct tissue contact between the
graft partners. Symptoms of localized incompatibility include cell necrosis
and vascular discontinuity at the graft interface, resulting in defective shoot-
root relationships and, in extreme cases, in union breakage. Important orchard
crops showing localized incompatibility include pear/ quince, apricot/ peach
and apricot/ plum combinations. The compatibility reactions apparently depend
upon the actual contact between the stock and scion. The union is often weak
resulting in the discontinuity of cambium and vascular tissues of stock scion.
There is poor translocation of the metabolites across the graft union and
masses of undifferentiated parenchymatous tissues or bark tissues or both are
commonly found at the graft union which may disrupt the normal vascular
connection between the stock and scion. Due to this poor translocation across
the defective graft union root starvation may occur. In extreme cases distortion
of vascular tissues takes place. It can be overcome by inserting a mutually
compatible interstock between the incompatible tissues e.g Bartlett pear on
quince root stock. Old Home is used as interstock to make the combination
a successful one (Macdonald, 1987).
2) Translocated incompatibility: This type of incompatibility is due to some
labile influence which can move across the graft union. It cannot be overcome
by insertion of mutually compatible interstock. There is formation of brown
line and necrotic area in the bark due to degeneration of phloem. Movement
of carbohydrates through the graft union is restricted, with the accumulation
above and reduction below e.g Hale's early peach grafted on Myrobalan B
plum root stock. They form weak union and there is distortion of the tissues
 Graft incompatibility – Causes and Remedial Measures 237

due to accumulation of starch at the base of the peach. The incompatibility

still exists even if a mutually compatible interstock Brompton plum is inserted
between these two, with the accumulation of starch at the base of Brompton
root stock. At cotyledon stage if the same combinations are tried, these are
highly compatible and no symptoms appear upto 13 years indicating that
some factors responsible for incompatibility reactions are not present at
juvenile or nursery stage of plants. Another example of translocated
incompatibility is Non Pareil almond on Mariana 2624 which shows complete
phloem breakdown, although the xylem tissue connections are normal.
However, another cv. Texas (almond) on Mariana 2624 is highly compatible.
Insertion of Texas as an interstock between Non Pareil and Mariana 2624,
bark disintegration occurs, resulting in incompatible graft union. The
compatible reactions changes with time also. For example, the pear cv. Bristol
Cross when grafted on quince makes good growth till 25-30 years but requires
interstock Beurre Hardy later to form a acceptable union.
3) Virus induced incompatibility: In some parts of world when Sweet Orange
is budded on Sour Orange, it shows incompatibility while in others it is
compatible and it was found to be due to viruses and was believed to be due
to production of some substance by the scion which is toxic to the stock.
Such changes may occur due to mutation or latent viruses.

Causes of incompatibility
The exact causes of the mechanism of graft incompatibility are not yet fully
understood inspite of many attempts made by researcher's throughtout the world.
They suggested that the possible causes include structural, physiological,
biochemical, pathogens, insects or combination of all these factors. However,
Pina and Era (2005) have summed up different reasons that influence on graft
success such as inherent system of cellular incompatibility, formation of
plasmodesmata, vascular tissue connections, and the presence of growth regulators
and peroxidases. In addition, phloem mobile proteins have been reported that
cross the graft interface when graft bridging is established and it is functional.
Brief discription of various causes of incompatibility are discussed as under:
1. Structural or anatomical aspects: The anatomical incompatibility may be
due to undifferentiated callus tissues e.g in apple and apricot (Herreo, 1955).
Due to differences in vascular tissue development and also due to excessive
growth of paranchymatous tissue between stock and scion necrotic layer is
formed. These disturbances cause structural abnormalities and disrupt vascular
continuity. The abnormalities may be found due to the presence of
parenchymatous cells at graft union, preventing the formation of vascular
238 Fruit Tree Physiology

continuity between the stock and scions although stock and scion may not
differ structurally. Sometimes, a graft layer develops at the joint and may
result in graft failure. Due to the development of some whorls or loops
distortion of the vascular tissues between stock and scions also takes place,
which restrict the movement of essential nutrients and water across the graft
union, resulting in the poor growth or failure of a graft union. Micro
spectrographic examination of cell walls of compatible graft union showed
higher concentration of lignin at the graft union and the cell wall of adjoining
cells were lacking in lignin, in incompatible unions. Thus any reaction
inhibiting the formation of lignin and development of middle lamella between
the stock and scion results in weak union. Thus, lignification is an important
event leading to compatible combinations. The incompatibility reactions in
plum, pear and peach are primarily due to structural abnormalities. The
union that occurs is mechanically weak e.g apricot on plum grafts (Errea
et al; 1994). In apple grafts the incompatibility is due to vascular discontinuity
leading to xylem interruption, by parenchyma tissues leading to death of
budded scions due to disruption of normal xylem functioning (Warmund
et al; 1993). In pear quince graft union there is accumulation of brown
pigment in cells near the contact zone due to hydrolysis of arbutin (a glycoside
in pear) and oxidation of hydroquinone. The oxidation products inhibit cellular
lignifications in the union. Reduction of these oxidation products occurs in
compatible cvs. whereas incompatible ones are not capable to do so. The
various structural and anatomical aspects which play important role in graft
incompatibility are:
i) Sieve tubes: Reduced number of well organized sieve tubes play an
important role in determining whether a union is compitable or not. In
cherry prunus incompatible grafts the number of well differentiated phloem
sieve tube is much lower at and below the union. There is a very low
degree of differentiation of phloem below the union resulting in greater
autolysis of cells (graft rejection and discontinuity at the normal vascular
connections between stock and scion i.e. localized incompatibility) and
may be due to lack of hormones, carbohydrates and other factors- the
size of sieve tubes depends upon auxin, cytokinins and sucrose
(Barckhansen, 1978). Lignin like deposits on the slime plugs near the
sieve tubes are also present in incompatible grafts whose amount increases
and tree growth gets reduced resulting in the formation of a necrotic line.
Activation of phenolic pathways like that of chlorogenic acid has been
observed in such grafts and these compounds get accumulated and are
used as sucrose by lignin like deposits. Further the lignin like deposits
get attached to cell wall as insoluble esters of ferulic and coumaric acid.
 Graft incompatibility – Causes and Remedial Measures 239

ii) Cessation of cambial activity: Cessation of cambial activity occurs earlier

in root stock of incompatible (Peach/ P18), than the compatible grafts
(peach/ P2032) because of seasonal inhibition of cambial activity at the
graft union. It has been observed in case of the localized graft
incompatibility e.g in pear/ quince (Mong and Carde, 1988). Auxin,
gibberllic acid, cytokinins, ABA and carbohydrates directly or indirectly
affect the cambial activity. Modification of the process of cambium
differentiation has been observed with fewer phloem sieve elements were
differentiating in the root stock of the incompatible than the compatible
grafts. Growth substances, sucrose or both hormones and sucrose have
an effect on the vascular differentation. There is more accumulation of
starch above the graft union due to the slowing down of carbohydrate
transport in heterograft in sieve tubes of both compatible and incompatible
grafts. The main factor of graft incompatibility was the bark discontinuity
which is partly expressed by the presence of periderm rings jointly formed
by the scion and stock (more frequent). In incompatible grafts, necrotic
areas associated with cork like cells are surrounded by phellogen, forming
a barrier between scion and root stock. The interface in the bark appeared
as a sharp line delimiting the tissues of partner, where it formed a gradually
multilayered transition zone in incompatible grafts of pear/ quince. Early
tissue differentiation occurred from the callus cells by the increase in
cells with banded secondary thickenings but delay in tissue differentiation
and vessel reconnection was evident in pear cv. Bosc/ EMqunice-C
incompatible combination.
Vascular discontinuity occurs with xylem interrupted by parenchyma
tissue in incompatible apple grafts leading to death of the budded scion
due to disruption of normal xylem functioning (Warmunt, 1993). Excess
growth of parenchymatous tissues and distortion of xylem elements
inhibits the movement of water from stock to scion in mango grafts. In
such cases, the initial disturbances occurred at outer phloem, when the
cambium zone was eventually affected and marked structural abnormalities
developed in the phloem and xylem tissues resulting in disrupted vascular
continuity across the bud union. Lignified parenchyma at the interface
and cambium continuity is always observed in compatible graft union
while unlignified parenchyma cells at the interface and cambium
discontinuity are the crucial symptoms of graft incompatibility. Phloem
degeneration of scion due to excessive callus proliferation resulting in a
compressive effect to the phloem sieve tubes and xylem trachied at the
graft union has been observed in Jhar ber. Necrotic layer formation at the
stock and scion interface prevents the cambium bridge formation which
240 Fruit Tree Physiology

further hinders the formation of vascular continuity between the

stock and scion also leads to graft incompatibility (Hartmann and Kester,
1989).
iii) Role of Plasmodesmata: Plasmodesmata play important role in the
mechanism of cellular communications (Schulz, 1999). When callus
cells come into contact, the cell walls undergo dissolution, holes in the
cell wall appear, plasmalema contact and lead to formation of
plasmodesmata. Plasmodesmata simplistically connect the grafting
partner’s and permits mutual metabolic interaction in the interfacing
cells through leaching or diffusion of some compounds from both sides
of the union to the other diffusing into the cell walls in order to achieve
a firm cohesion between root stock and scion. Research on the mechanism
of plasmodesmata formation has shown prominent differences in the
development of interspecific plasmodesmata between graft partners
suggesting that cell recongnition and functional coordination may be
involved in graft formation, with special attention placed on the
establishment and modification of primary plasmodesmatas as in the
novo formation or modification of secondary plasmodesmata. An exact
cooperation of both cell partners is required so as to form continuous
secondary plasmodesmata, since insufficient coordination between
adjacent cells leads to the formation of mismatching, half plasmodesmata
through the cell wall only in one cell partner. In graft interface of
incompatible heterografts, discontinuous half plasmodesmata have been
observed in the graft unions between different types of cells, using species
specific cell markers in order to identify the graft interface and may
contribute to the graft failure due to misalignment of the graft partners.
Recent studies on physiological control of plasmadesmata through caged
probes indicate the possible existence of plasmodesmatal connectivity
between invitro callus cells in P. munsoniana and presence of non
functional plasmodesmata in incompatible combinations (Martens et al;
2004).
iv) Vascular connections in the graft process: The basic requirement for
successful graft is the formation of vascular connections and is the last
step in the process of grafting (Wang and Kolman, 1996). The main
reason for incompatibility in woody plants may be that the new vascular
connections are not well differentiated or weakly established (Errea et al;
1994). An abnormal process of neocambium differentiation leading to
cambial involution and lack of differentiation into new vascular elements,
has been found in these cases e.g pear and quince grafts and apricot on
prunus grafts (Errea et al; 1994). The formation of functional connection
 Graft incompatibility – Causes and Remedial Measures 241

is essential for successful grafts in herbaceous plants and in woody plants.

Incompatible grafts can grow for several years without any external
indication of incompatibility, denoting the presence of functional vascular
connections in incompatible grafts (Hartmann et al; 1997). However
studies on phloem regeneration across the graft interface through C14 -
labeling have revealed that assimilated transport occurs from the apical
callus of the scion to the basal callus of the stock which indicates a close
relationship between transport and phloem restitution in compatible grafts
while as in incompatible grafts no C14 transport to the stock occurs and
there was no correlation between phloem regeneration and assimilation
transport across the graft interface. However, in apricot on plum grafts,
the formation of new vascular connections occurs in both compatible and
incompatible combinations. The transport of disodium fluorescein across
the graft union confirms the communication and functionality of these
connections since fluorescence can be seen in both partners of the graft.
In these combinations, the difference between compatible and incompatible
grafts lies in the presence of a portion of the callus tissue in incompatible
grafts that cannot differentiate into cambium and vascular tissue, resulting
in the existence of wide areas at the union similar to undifferentiated
callus cells. This lack of cambial activity in some areas of the graft union
could affect the activity of the new xylem and phloem formed, causing
discontinuities in the cambium and the formation of a parenchymatous
line interrupting the vascular connection producing a mechanically weak
union (Hartman et al; 2002).
Studies on long distance translocation of proteins and mRNA via
phloem have shown that the P-proteins (PP1, PP2) are synthesized in
companion cells of differentiating sieve element companion complex
within the phloem transport and are translocated into sieve elements
when these are established and functional. The expression of PP1 and
PP2 is related to different stages of phloem differentiation and they are
phloem mobile. The conservation of the pattern of PP2-gene expression
in the sieve element companion cell complex suggests that it plays an
important role in the development and function of the vascular system.
Although, it is also known that the majority of the phloem proteins not
only perform sieve tube maintenance, but also can be linked to direct and
indirect stress and defence responses.
Growth regulators like auxins, GA, ABA also affect the stock and
scion relationship. Auxin is an important substance involved in the
development of compatible union, which is released from vascular strands
of the stock and scion and induces the differentiation of vascular tissues,
242 Fruit Tree Physiology

functioning as morphogenic substances (Matteson et al; 2003). Its

translocation from the root system has been related to graft incompatibility
in apples, since a supra and basipetal movement of auxin can organize
the morphogenetic pattern of the entire plant body even accelerating the
formation of a successful graft. Other compounds like polyphenols also
play a prominent role in graft formation by influencing lignification
processes and by their protein precipitating feature. The stress situations
can lead to both the accumulation of flavanols and their degradation by
oxidases which can bring about marked effects on the growth and
metabolism of tissues such as inhibition of the lignin pathway. Synthesis
of flavanones can determine degree of compatibility such as prunasin in
Prunus (Errea et al; 2000) and can be stimulated by ABA and GA. All
these macro-molecules (phloem proteins, RNA, hormones) present in the
phloem sap appear to be important during vascular differentiation in
compatibility process. The involvement of certain enzymes in the cellular
behavior during the first steps of graft formation has also been studied
in different species, although the specific role and effects on
incompatibility is still not clear. Studies on proteins, peroxidases in the
phloem and acid phosphatases in the Prunus avium/ Prunus cerasus
grafting by (Schmid and Feucht, 1985) confirmed that it is possible to
define a good union not only by morphological characters but also by
biochemical methods. Based on biochemical assays, Gulen et al. (2002)
concluded that the absence or presence of peroxidase isozyme profile in
the pear qunice combination is an experimental approach to predict
incompatibility reaction. Recent investigations on graft development
indicate the involvement of increased peroxidase and catalase activities.
2. Differences in stock and scion growth: The different growth characteristics
of the stock and scion vigour is one of the possible mechanisms of
incompatibility i.e if marked differences occur in vigour or in the time or
completing vegetative growth for the season. Cambial activity during spring
originating with swelling of buds of scion spread through the graft union
activates the cambium of the root stock in both compatible and incompatible
combinations. However, the growth difference between stock and scion are
not the principle cause of incompatibility.
3. Environmental aspects: The success of grafting is also determined by various
environmental factors such as drought, water logging, heavy soils, poor soils,
low temperatures, frost or heavy rainfall.
a. Frost injury: In peach grafts the root stock is often more susceptible
than the above ground shoots to the frost injury and results in complete
failure of graft union in different parts of the world (Layne, 1984).
 Graft incompatibility – Causes and Remedial Measures 243

b. Salt stress: In citrus the incompatibility among the stock and scion is
due to their sensitivity towards higher concentration of salt in the soil.
c. Low temperature: Phloem disintegration, breakage and distortion are
prevelent in peach trees when exposed to low and extremely low
fluctuating temperatures. It also results in serious trunk and branch injuries.
d. Excessive rainfall: Incompatibility in cherry has been attributed to
excessive rainfall on poor drained loam soils (Futch et al., 1988).
4. Virus and mycoplasma problems:
a. Citrus: Incompatibility in citrus occurs due to infection by viral diseases
like tristiza, porosis, xyloporosis etc. Quick decline in citrus is caused by
tristeza virus due to the production of substances by the scion that are
toxic to root stock resulting in incompatible reaction (Toxopcus, 1936)
but sweet oranges are tolerant while lethal to sour orange root stocks.
b. Blackline in English walnut: It is a type of delayed incompatibility and
is due to cherry leaf roll virus infecting symptom less pollens of English
walnut which roll down to graft union and kills Juglans regia grafted on
J. hindsii seedlings.
c. Apple necrosis and decline (ANUD): Tomato ring spot virus inoculated
by nematodes affects the union.
d. Prune brownline: Caused by tomato ring spot virus.
e. Pear decline: Pear decline disease on Bartlett trees grafted on P. pyrifolia
root stock occurs at the graft union due to viruses.
5. Nutritional deficiency:
a. Deficiencies of certain nutrients like N, P, K, Mg, Mo etc. may result in
incompatible union's e.g when Jonathan apple is grafted on EM-IX root
stock the scion develops the Mo deficiency. However, Jonathan on other
root stocks does not show these deficiency symptoms.
b. When peach is grafted on Myrobalan B plum root stock deficiency of N,
P, K and Mg occurs resulting in incompatibility symptoms.
6. Biochemical reasons of graft incompatibility:
Biochemical substances in root stock: Prunasin a cynogenic glucoside is
responsible for incompatibility in certain pear cvs. on quince root stock. It
is found in the quince, but not in pear tissues. It is translocated from the
quince into the phloem of the pear. Prunasin breaks down in the region of
the graft union and forms hydrocyanic acid as one of the decomposition
products which leads to lack of cambial activity at the graft union and causes
anatomical disturbances in the phloem and xylem at the resulting union.
244 Fruit Tree Physiology

There is destruction of the phloem tissues at and above the graft union
leading to reduced conduction of water and minerals in both xylem and
phloem (Gur et al; 1968). In roots there is further decomposition of prunasin
leading to production of hydrocyanic acid and killing of quince phloem.
Biochemical substances present in scion: High cyanogenic glycoside content
in the bark of peach cultivars and some of it is tranolocated into almond root
stock leads to incompatibility in case of peach scion on almond root stock.
With increased glycosidic content, the 3-glycosidase activity of almond root
stock bark increased. Due to this non glycosidic hydrocynic acid accumulates
in root stock bark, particularly in incompatible scion root stock combinations
leading to death of tissues at the union of incompatible combinations
(Gur and Blum, 1973).
Phenols and incompatibility: Phenolic compounds play an important role in
different steps of healing of a graft union:
Production of callus tissues: Healing and lignification at graft union is
determined by the content and nature of callus tissues (Deloire and Mebant
1986, Machiex et al; 1986). There is diffusion of phenols from both sides of
the union to the other, diffusing into the cell walls and resulting in a firm
cohesion between autografts due to mutual metabolic interaction of the
interfacing cells. A steep gradient is established at the line of union because
the compounds produced by the phenolic heterospecific grafts are
quantitatively different from each other. Relatively small quantities of phenols
could be enough to produce locally limited disfunctions between two cells.
As a consequence, the micro tubule and internal side of the plasmalemma
may be affected, changing the structure and orientation of cell wall micro
tubule. These alterations could result in problems in the permeability of the
cell wall and this membrane deterioration is associated with anthocyanidins
(Trippi et al; 1988). In callus cultures of fruit trees the membrane damage
has been observed due to presence of flavonon which also interferes with the
permeability of tissues (Errea, 1998).
Differentiation into vascular tissues: Various phenolic compounds have been
found to be associated with the processes of division, development and
differentiation into new tissues (Mossela and Machiex, 1979). High level of
prunasin limits the proliferation and differentiation of the cells in callus cultures
of P. avium (Feucht et al; 1988). Other compounds like chlorogenic acid, which
might inhibit the development of the callus and stimulation of cell division
have been observed in P. avium (Jordan et al; 1980). Compounds like
flavanoles, cataquins and proanthocyanidins, that affect cell division intensity
have been observed in callus cultures of P. avium (Fecht and Nachit, 1977).
 Graft incompatibility – Causes and Remedial Measures 245

These events are directly related to the micro localization of these compounds
at the cellular level which results in oxidation of these compounds and
peroxidases and phenoloxidases (Elsnter et al; 1994).
Accumulation of phenols in cambium tissue of some graft combinations
leads to disorganization at the sub cellular level leading to cellular damage
and alter the complete phloem cambium system around graft union e.g. apricot
grafts (Errea et al; 1994 b). Differentiation into new xylem and phloem
incompatible grafts can grow for several years without any external indication
of incompatibility, denoting the presence of functional vascular connections
in incompatible grafts (Hartman et al; 1990). Phenolic compounds are involved
in these steps due to their important role as crosslink agents between
polysaccharides (Markwalder and Neukom 1976., Fry 1988) and their
implication in liginin synthesis pathway (Tomaszewski, 1962). The lignification
of cell wall could be the main event leading to the formation of a solid union
and therefore resulting in compatible grafts (Feucht and Schmid, 1979).
Sap exudation during grafting: In walnut exuding sap from root stock grafts
often results in poor take of the scions/ grafts. Bleeding results in lack of union
of the scion to the stock due to the inhibition of callus formation. A toxin Juglane
(5-hypromyl-1, 4-nepthoquinone) exuding in the walnut sap has been found to
be responsible for this inhibition of callus formation (Pratariera et al; 1983).
Low auxin concentration: Auxins (IAA) play an important role in the
development of compatible union as these induce the differentiation of vascular
tissues (Quessada and Macheix 1984., Aloni, 1987). Accumulation of
polyphenols (which are known to affect auxin transport) above the union
leads to incompatibility of P. avium / P. cerasus grafts (Stenlid, 1976). The
differentiation of xylem and phloem and lignification are also affected by
low auxin content and results in incompatibility.
Biochemicals related to soil type: Apricot cvs. on peach and myrobalan are
short lived, and when grafted on hybrids of P. bessay x P. salciana or
P. cerasifera x P. spinosa often develop incompatibility symptoms such as
brown plates at the union or even breakage of stem on heavy silty loam soils
(Cummins, 1985). The concentration of flavanols increases under these stress
conditions. These phenols escape from the vacuole into the cytoplasmic
matrix where they are oxidized by peroxodases and phenoloxidases (Feucht
and Treutter, 1989) forming a quinone which polymerizes forming toxic
compounds for a number of chemical reactions (Possel et al; 1980). These
quinones form covalent links with neophlic groups in protein and other
macro molecules and their irreversible unions affect posterior development
of lignin pathway, imparing a good connection between stock and scion
246 Fruit Tree Physiology

(Possel et al; 1980., Feucht and Treutter, 1989). There is protein precipitation
resulting in cellular necrosis in xylem of incompatible combinations due to
these irreversible unions (Feucht and Treutter, 1989).
Low temperature: Exposure to low and exteremly fluctuating temperatures
leads to trunk and branch injuries in peach trees. The phloem gets disintegrated,
breakage and distortion of phloem occurs and the cambial cells indicate
absorption of safranin leading to graft incompatibility.
Thus, the various reasons leading to graft incompatibility can be summed up
as follows:
1. Use of genetically different plant material for scion and root stock. The
family, genus and species of plants must be considered while grafting.
2. Plant viruses especially latent type (Juglanus regia clones grafted on J.
hindisi), can reduce overall percentage success quite dramatically. So
production of virus free plant material should be performed e.g EMLA series
in apple.
3. Use of vigorous root stock with very weak growing scion cultivar.
4. Formation of cork tissue layers between scion and stock. It acts as a barrier
to the transport of water and nutrients as well as causing mechanical weakness.
5. Failure of fibres of stock and scion to interlock effectively.
6. Death of cambium and phloem tissues at the graft union.
7. Abnormal distribution of starch at graft union and differences in biochemical
compounds or reactions between stock and scion.

DETECTION OF INCOMPATIBLE COMBINATIONS

Various techniques used to detect the incompatible graft combinations are as:
1. Electrophoresis: It is used for testing cambial peroxidase banding pattern of
the scion and root stock e.g in chestnut. Peroxidase produces specific lignin
production. In incompatible grafts there is increased peroxidase activity as
compared to the autografts, and adjacent root stock and scion cells must
produce similar lignin and should have identical peroxidase enzymes patterens
to ensure the development of functional vascular system across the graft
union. If the peroxidase bands match the combination it indicates compatibility,
otherwise incompatibility may be predicted (Noblitt and Henry, 2008). In
case of grapevines this method was used by Gokbayrak et al. (2007).
2. Invitro callus fusion method: Callus tissues of stock and scion having
different degrees of incompatibility are used for easy detection of
incompatibility. Incompatibility is manifested by the breakdown of trees at
 Graft incompatibility – Causes and Remedial Measures 247

the union area after many years of grafting. The differences of two callus
partners, development of cells at the contact surface (cell arrangement,
intensity of cell wall staining) and the presence of lipid and phenolic
compounds of callus tissues help in early detection of compatible/ incompatible
combinations e.g in apricot and different prunus root stocks (Errea et al;
2001) .
3. Microscopic or histological detection: very little callus differentiates into
cambium and vascular tissue in incompatible apricot / plum grafts, however
a large portion of callus never differentiates. Histological detection within
weeks after grafting helps in early detection of graft incompatiblity in fruit
trees (Dorsmann, 1966).
4. Magnetic resonance imaging (MRI): Detection of vascular discontinuity
(incompatibility) can be done by MRI in bud unions of apple. A high MRI
signal intensity indicates bound water in live tissues and establishment of
vascular continuity between the root stock and scions. Graft incompatiblity
caused by poor vascular connections was detected through MRI technique
by Warmund et al. (1993).
5. Macroscopic detection: Macroscopic or external examination for discontunity
of inner bark tissue at the graft union is simplest and non destructive method
of incompatibility detection.
6. Breaking strength test: If the union breaks while applying slight pressure
by hand over it such a graft will be considered as incompatible.

METHODS TO OVERCOME INCOMPATIBILITY

1. Double working: To overcome the localized incompatibility of the scion
and root stock a mutually compatible interstock can be used. The interstock
prevents physical contact of the root stock and scion, and affects the
physiology of normally incompatible scion and root stock e.g Bartlett pear
grafted on dwarfing Quince root stock shows incompatibility but when 'Old
Home' is used as an interstock, this three graft combination is completely
compatible and satisfactory tree growth take place (Mosse, 1958).
2. Time of grafting: Dormant season is best for grafting when no exudation
occurs at the juncture of stock and scion. A ring of callus tissue is formed
from the cambium of the scion and on the surface of stock wood, forming
a successful graft union. When xylary sap excudes no callus is formed and
vascular connections between scion and stock do not form.
3. Presence of nurse limbs: Bleeding at the graft union is reduced by nurse
limbs and results in greater success. They also supply photosynthates to rest
of the tree until the grafts are large enough to take over this function.
248 Fruit Tree Physiology

4. Inarching / bridge grafting: Inarching/ bridge grafting with a seedling of

compatible root stock if the incompatibility is discovered before the tree died
or broke off at the union could be done with a mutually compatible root
stock.
5. Remove and top work the root stock: Incompatible scion should be removed
and the stock should be top worked by a compatible one through grafting.
6. Use of correct root stock: Root stocks which show compatibility with scion
shall only be used and incompatible ones shall be avoided.
7. Virus free planting material of both root stock and scion: Both stock and
scion used for grafting shall be disease and viruses free so as to have a
compatible graft union.

CONCLUSION
The inability or failure to produce successfull graft is called graft incompatibility.
There is no clear cut distinction between a compatible and an incompatible graft
union. The stock and scion of related species unite readily and form a compatible
graft union and grow as a composite plant while unrelated species fail to do so.
However, sometimes unrelated stock and scion initially show compatibility but
develop incompatibility later and die eventually. Several visual symptoms like
premature death, poor tree growth, low graft success, dieback, nutritional deficiency
etc. are observed in incompatible combinations. Incompatibility may be either
translocated or localized. The exact causes of graft incompatibility are not yet
fully understood, however, many researchers suggest that the possible causes
include structural, physiological, biochemical, pathogens, insects or combination
of all these factors. Several techniques used for early detection of incompatibility
include electrophoresis, invitro callus fusion, histological detection, breaking
strength test etc. Inompatibility can be overcome to some extent through double
working, inarching, use of correct root stock, virus free planting material etc.

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 10

PHYSIOLOGY OF DWARFISM
IN FRUIT PLANTS

D warfing is an alternation in the normal growth pattern. Dwarfing fruit trees

play an important role in fruit growth, development and quality. Large and
vigorous fruit trees are a problem to the fruit growers, because ladders have to
be used during fruit thinning, bagging, harvesting pruning and other cultural
operations. In dwarf, small and compact trees operations like pruning, thinning,
spraying and harvesting etc. are easy and could lead to production of high grade
fruit at lower production cost. A dwarf plant is that which is smaller than normal
size at full maturity and possesses other characteristics like precocity, canopy
architecture, and time of flowering and altered fruit size. Anatomically and
histologically the dwarf plants are like that of standard ones but horticultural
practices can alter the physiology of a tree, resulting in dwarfing. Genetic changes
(natural or imposed through selection breeding) or horticultural manipulations e.g
root stock, pruning, training, deficit irrigation and plant bioregulators etc. can be
used to produce dwarf trees. In modern orchards, dwarfing is considered a desirable
characteristic, where genetic dwarfs may be selected and propagated, or more
often scions are grafted on to dwarfing root stocks. Almost all modern apples in
commercial use are propagated as dwarf or semi dwarf trees for the ease of
picking, spraying and high productivity per unit of land. The principle underlying
dwarfing is to make the best use of vertical and horizontal space per unit time and
to harness maximum possible returns per unit of inputs and natural resources.

PHYSIOLOGY OF DWARFISM
Various horticultural practices are used to achieve dwarfing in fruit crops such as
selection of spur type scion cultivars, use of interstocks, pruning, root pruning,
252 Fruit Tree Physiology

use of growth retardents, control of nutrient elements, girdling, scoring, bark

inversion, use of dwarfing/ ultra dwarfing/ semidwarfing root stocks.
The mechanism underlying dwarfing involves anatomical, physiological and
biochemical changes. It has been observed in apple that bark and phenolics play
important role in dwarfing. Some researchers postulated that dwarfing occurs
when auxin produced by the shoot tips is translocated basipetaly downwords to
the roots through phloem. This influences the root metabolism and affects the
amount and kind of cytokinin synthesized in roots and translocated to shoots via
xylem. Since auxin is synthesized in shoots and young leaves and flows basipetally
through phloem and cambial cells to the roots. Some auxin is degraded in the
bark by IAA oxidase, peroxidase and phenols and other compounds whose levels
are genetically controlled. The level of auxin that reaches roots influences root
growth and metabolism including synthesis of other hormones like cytokinins,
gibberellins and ABA which in turn maintain balance in shoot and root growth.
It has been found that thicker bark and higher starch levels in dwarfing root
stocks indicates a low level of auxin in these tissues.
It has been proposed that dwarfing root stocks or interstocks control tree size
by controlling the auxin passing through the bark of the root stock or interstock.
Tissue culture experiments have indicated that the calli of dwarfing root stocks
produced larger amounts of growth inhibiting compounds than those of vigorous
ones. The strong evidence for this theory is that insertion of 20 cm length of bark
of M26 in the scion of Gravenstein / MM111 trees resulted in more dwarfing than
did a 10 cm length of bark (Jones, 1976).
Effects of interstocks on dwarfing have been investigated and it was found
that higher per centage of bark on new growth (scions and interstocks) in plants
having M9 (dwarf root stock) as interpiece than those with Red Delicious or
MM106 (vigorous root stocks than M9). It has also been observed that there is
change in the orientation of phloem-ray cells and sieve tubes of the scion from
their normal conducting pattern. Anatomical studies on dwarfing mechanism
indicate large number of vessels and twice as many xylem fibres are produced in
vigorous root stocks than the dwarfing ones (Beakbane et al; 1974). Besides, the
most dwarfing root stocks posses the maximum percentage of living tissues than
the vigorous ones.
Nutrient uptake studies showed that more Ca is present in conducting phloem
than in non-conducting phloem, cambium or xylem. The non-conducting phloem
are more in dwarfing root stocks and Ca accumulates in necrotic tissues. A greater
proportion of non-functioning phloem has been found in root stocks with thick
bark (M9) than those with thin bark i.e MM106 or M7 (Chu and Simons, 1984).
 Physiology of Dwarfism in Fruit Plants 253

Application of PGRs such as Prohexadione-calcium can prevent excessive

shoot growth and maintain size control in apple crowns by inhibiting gibberellin
biosynthesis. Training also will reduce tree size and decrease self shading, but
pruning can remove apical dominance by reducing auxin concentrations and
rapid regrowth (e.g water sprouts) from axillary or latent buds can occur. In
apple, dwarfing root stocks have been demonstrated to reduce auxin in cambial
sap and in movement through the scion/ root stock graft and to reduce cytokinin
concentration in xylem exudates although other physiological processes, such as
reduced hydraulic conductivity, may contribute to the dwarfing effect. Peach tree
crown size and form can be modified when root systems are restricted by deficit
irrigation, physical barriers, or by roots of competing plants. Crown growth may
decrease allometrically with reduced root growth but reduced growth may also be
associated with stress induced hormones such as abscisic acid and with inhibited
cytokinin production. There is tremendous diversity in apple and peach tree crowns
and hormones were suggested to play a role in the diverse tree architectures that
could be selected in breeding programs. Recent research has demonstrated
quantitative differences in auxin and cytokinin concentrations in different growth
habits of peach trees. Trees with less sylleptic branching and more upright growth
had high auxin concentrations as the growing season progressed and low cytokinin
to auxin ratios throughout the season. The effects of cultural practices on
endogenous hormone concentrations of peach and apple trees with different growth
habits are virtually unknown. Significant progress is being achieved to discover
genes that affect morphological traits and to include them in breeding programs
for fruit crops. Cultural practices which affect tree form will likely have to be
adapted to new genotypes and endogenous hormones may be an important link
for crown management by both genetic and cultural techniques (Tworkoski, 2003).
From the above discussion, it is concluded that physiological mechanisms of
dwarfness are not fully understood yet, however, a number of correlations have
been claimed for predicting dwarfing capacity of potential root stocks e.g
1. Through correlation between dwarfing propensity and various structural
features as:
a. Percentage of live tissues of the root cross-section.
b. Bark/wood ratio.
c. Percentage of medullary ray tissues in root cross section.
d. High stomatal density of leaves.
2. Bark is key factor for dwarfness as it reduces translocation of auxins, sugars
and other compounds.
3. Genes exert their effect on dwarfness.
254 Fruit Tree Physiology

METHODS TO ACHIEVE DWARFISM

Dwarfness can be achieved in fruit plants through various ways such as:
A. Use of dwarfing root stock/ interstock and genetically dwarf scion cultivars.
B. Use of bioregulators.
C. Use of incompatible root stock.
D. Induction of viral infection.
E. Pruning and training.
F. Nutrients.
G. Phenols.
H. Invitro techniques.
I. Genetic engineering.
J. Others.
A. Use of dwarfing root stocks, inter stock and genetically dwarf scion
cultivars : The use of vigour controlling root stocks is one method used to
promote early fruit bearing, reduce vigour and increase yield. Although
dwarfing root stocks have been widely used for many decades, the mechanisms
involved and specific effects on fruit tree (apple) development remain poorly
understood (Webster, 2004). The difficulty in understanding root stock effects
may be because mature trees result from successive cycles of shoot growth,
flowering and fruiting that lead to cumulative and compound effects on tree
structure and functions (Webster, 1995). Apple dwarfing root stocks induce
two most important effects i.e pre-cocity and reduction in tree size (Lauri
et al., 2006) e.g some root stocks induce flowering in apple in the second
year of tree growth (Seleznyova et al., 2004). It is hypothesized that pre-
cocity in flowering induced by dwarfing root stocks is likely to have significant
effects on the ensuing development of the apple tree form, by shifting annual
shoot development from mainly monopodial (vegetative) to sympodial (floral)
shoots (Seleznyova et al; 2007). These shoot types differ in structure and in
the axillary bud outgrowth during the following year (Lauri and Terouanne,
1998). In addition, sympodial annual shoots subtend flowers and fruit that
compete with vegetative growth for resources (Ferree and Palmer, 1982).
Various studies conducted suggested a correlation between early flowering
and smaller trees size (Lauri et al., 2006.; Costes and Garcia Villanueva,
2007). The use of inter stems (grafting a third genotype in between a root
stock and a scion) is another method used to induce scion dwarfing, and also
to induce tolerance to winter cold and resistance to disease. Malling 9 (M9)
inter stems have been found to be effective for reducing the vigour of apple
tree but the influence of M9 as an inter-stem is generally less than that of M9
 Physiology of Dwarfism in Fruit Plants 255

used as a rootstock. The effect of some vigour-controlling root stocks used

as inter-stems was reported to be related to the length of the inter-stem piece
in apple (Di Vaio et al; 2009) and peach. Dwarfing apple root stocks reduced
the formation of nodes during shoot growth and the cumulative effect of
such reduction over time can have a dramatic effect on branch development.
Furthermore, dwarfing root stocks and inter stems reduced the number of
extension shoots and promoted the formation of floral shoots. The
physiological mechanisms of vigour reduction by root stocks or inter stems
are not clearly understood.
During the last century, many mechanisms for dwarfing have been proposed.
It has been proposed that dwarfing might be caused by water supply restrictions
to the scion induced by anatomical characteristics of the root stock. On the
other hand, others suggested that dwarfing was caused by a reduction of solutes
transported to the scion through the root stocks. This theory contradicted a
previous study that reported an increase of solutes in scions grafted on dwarfing
root stocks. A long series of studies reported possible mechanisms related to
the production and translocation of hormones in the plant. However, exactly
how hormones act in the dwarfing process is not well understood. It has also
been reported that in some cases dwarfing might be caused by partial
incompatibility between the scion and the root stock, which may alter the
transport of minerals and hormones. It is well known that root restriction or
pruning can influence scion vigour and the root: shoot ratio. While the root:
shoot ratio is sometimes less in trees on dwarfing root stocks, this is not always
the case and hypothesized that dwarfing apple root stocks may impose scion
dwarfing by changing the hydraulic conductance of the whole tree. However,
Cohen et al. (2007) later concluded that apple root stock vigour is not related
to hydraulic architecture or hydraulic conductance of the root stock. They
suggested that the graft union may play a central role in limiting water transport
and thus induce scion dwarfing. Experiments with selected peach and olive
root stocks have indicated that there was no influence of the graft union on
stem hydraulic conductance but that conductance of the root stock was
apparently directly related to peach root stock vigour. On the other hand,
Clearwater et al. (2004) reported that differences in root stock induced vigour
in kiwifruit vines were related to differences in the timing of phenological
development of the root stocks. Nardini et al. (2006) also reported that dwarfing
in specific olive root stocks was unrelated to hydraulic conductance, while
Trifilo et al. (2007) suggested that the dwarfing characteristics of a dwarfing
olive root stock obtained through mutagenesis may also be related to delays in
development. From these varied accounts it is clear that there is probably more
than one mechanism involved in causing root stock mediated dwarfing effects
in different fruit crop species.
256 Fruit Tree Physiology

Recent studies with peach have demonstrated that daily patterns of stem
water potential are directly related to patterns of shoot growth. The pattern
of stem water potential occurring during the afternoon hours strongly affects
shoot growth rates in the field. Subsequently, a strong correlation was found
between stem water potential and shoot growth over a day among trees on
root stocks that imparted differing amounts of vigour. Vegetative growth was
also found to be correlated with cumulative water potential differences during
the first half of a growing season. Subsequently, it was demonstrated that
differences in stem water potential are causally related to differences in
relative shoot growth rates among peach trees on different root stocks. Stem
water potential is strongly influenced by stem hydraulic conductance, and
differences in measured stem hydraulic conductance among root stocks
correspond to differences in xylem vessel characteristics and theoretically
calculated hydraulic conductance of the same root stocks. Furthermore, it
appears that xylem vessel diameter and number are the main root stock
characteristics influencing scion vigour in peach trees growing on graft
compatible root stocks.
Xylogenesis has been hypothesized to be regulated by hormones produced
both in leaves and in roots. Indoleacetic acid produced in leaves and transported
basipetally appears to control xylem regeneration around wounds. In the
basipetal direction, high concentrations of auxin in close proximity to leaves
have been proposed to cause the formation of many vessels of small diameter
while low auxin concentrations farther from leaves appear to promote the
growth of fewer large vessel. These results were confirmed by experiments
on transgenic petunia plants. Cytokinin appears to promote vessel regeneration
in the acropetal direction in the presence of indoleacetic acid by increasing
the sensitivity to auxin and stimulating xylem formation. These results suggest
that in compound trees the scion could influence xylogenesis in the root stock
or vice versa. The comparison of xylem anatomy among peach root stocks
reported by Tombesi et al. (2010) focused on anatomical differences among
root stocks, but it did not determine whether the xylem characteristics of the
root stocks influenced the xylem characteristics of a scion grafted onto those
root stocks, or whether the xylem characteristics of a vigorous root stock
could influence the xylem characteristics of a size-controlling root stock
genotype grafted onto it. The primary aim of this work was to study the
influence of the root stock on the xylem anatomical characteristics of the
scion to determine whether the vigour and xylem vessel diameters of the root
stock induce changes in the xylem structural characteristics of the scion. Further
it was determined that whether a vigorous root stock can influence the xylem
vessel characteristics of a dwarfing genotype used as an inter stem.
 Physiology of Dwarfism in Fruit Plants 257

Most of the work has been done on apple dwarfing root stocks. There is a
combination of anatomical, physiological and biochemical factors that appear
to be the cause of the dwarfing effect.
1. Anatomy of dwarfing root stock : Dwarfing root stocks have smaller
xylem vessels and less xylem fibres than vigorous root stocks, besides
having higher percentage of bark and wood ray tissues per unit cross
sectional area of root system. Root stock genotype only marginally affects
scion xylem vessel characteristics. Thus the xylem vessel characteristics
of the dwarfing root stock genotypes appear to influence tree growth
directly rather than through an effect on the xylem characteristics of the
scion. A dwarfing root stock genotype used as an inter stem appeared to
work as a physical restriction to water movement, reducing potential
xylem flow and conductance of the whole tree. Diameter of xylem vessels
in dwarfing root stocks (M9) shows that there is a greater percentage of
living xylem tissue compared to dead, lignified vessels in more vigorous
root stocks. At the graft union of a scion on M9 the xylem vessels are
also smaller in diameter than on other root stocks. The theory is that
there is less water transport in trees on M9.
2. Nutrition : Partitioning of carbohydrates between above and below graft
union parts of the tree is affected by root stock. More dry matter is
partitioned to root system by vigorous root stock than dwarfing root
stocks while more carbohydrates are partitioned to fruiting structures
such as spurs and fruits and less to tree frame by dwarfing root stocks.
3. Hydraulic conductivity : Reduced root hydraulic conductance helps to
induce dwarfism e.g olive (Nardini et al; 2006).
4. Translocation of water and minerals : Dwarfing root stocks are less
effective than vigorous ones in uptake of nutrients and water and their
supply to shoots. Partial blockage at graft union affects water and nutrient
translocation e.g reduction of translocation of P and Ca (Bukovac et al;
1958).
5. Phytohormones : Auxin is the main hormone affecting scion/ stock
interactions. Less auxin flow takes place through bark of dwarf root
stock as compared to vigorous ones. M9 a dwarfing root stock showed
greater ABA: IAA ratios (2.0) compared to vigorous root stock M111
(1.4). Lower concentrations of both hormones, IAA and ABA are observed
in dwarfing root stocks like M9 and M27 while more vigorous root
stocks like MM106 showed higher cytokinin concentration in root pressure
exudate and shoot xylem sap (from above graft union in grafted trees).
M9 had mostly zeatin and MM106 had mostly zeatin riboside. The greater
258 Fruit Tree Physiology

amount of cytokinins in vigrous root stocks helps in their maximum

growth rates compared to dwarfing ones.
Lack of ability to produce soluble sugars results in reduced growth
e.g starch hydrolysis. NAA suppresses the conversion while IAA promotes
it that is why NAA causes dwarfing. In general higher ABA levels have
been observed in dwarfing apple stocks and cultivars. It controls tree
height by influencing the translocation of other growth substances.
In apple and pear culture the dwarfing root stocks have been widely
used for a very long time. However, the mechanisms responsible for
dwarfing effect on the scion are still not fully understood. In general, the
dwarfing root stocks reduce the amount of scion dry weight through
reduction in both the rate at which vegetative shoots grow (extend) and
the time period over which they grow. A dwarfing root stock directs a
greater proportion of dry weight into fruit production rather than into
vegetative growth unlike vigorous root stocks which diverts the dry
weight into vegetative growth mostly. The reason is that dwarf scions
have less leaf area than those on vigorous root stocks. Due to reduction
in leaf area the scion shoots on dwarfing root stocks produce less
photoassimilates relative to scions on vigorous root stocks. The ability to
intercept solar radiation and convert this into fruit production via
photosynthesis is also influenced by the differences in leaf orientation on
the tree. The factors associated with dwarfing root stocks result in
reduction in tree size. Kamboj et al. (1999) observed that levels of
cytokinin in the shoot xylem sap increased with increase in vigour of the
root stock as it is produced in the root system and translocated to shoots.
6. Dwarfing root stock : These helps to increase the proportion of bark that
results in dwarfing e.g in avocado the dwarfed trees had 22.7 percent of
bark area while tall ones had 12.9 per cent only. This is because more bark
leads to more destruction of auxins. Lockhard and Schneider (1981)
proposed that the greater thickness in the bark of the dwarf trees is associated
with a higher degradation of auxins by lAA-oxidase, peroxidase, and
phenolic compounds that are present in the bark. As a consequence of this
reduction of auxin supply, there is a reduction in the production of cytokinins
by the roots that alters the normal growth pattern of the tree.
7. Reduced root systems of dwarfing root stock : In general, the more
dwarfing a root stock the smaller is its root system e.g M9 has limited
root system compared to seedling root stock (Fernandez et al; 1995).
8. Depletion of solutes in xylem sap of scions by dwarfing root stocks :
It has been observed that dwarfing root stocks and inter stocks lead to
depletion of xylem sap with respect to N, P, K, Ca, and Mg (Jones, 1976).
 Physiology of Dwarfism in Fruit Plants 259

9. Dwarfing root stocks induce restricted canopy development of scion

variety : It has been documented by various workers that trees on dwarfing
root stocks show restricted canopy development and has been attributed
to several factors:
I. Induction of wider crotch angles to lateral branches (become more
horizontal) and spreading habit by dwarfing root stocks. Horizontal
branches show both decreased vegetative growth and enhanced
flowering and fruiting (Crabbe, 1984).
II. Reduced growth rate of both vertical and horizontal branches, the
effects being greater on vertical ones (Webester, 1995).
III. Reduced shoot growth due to heavy cropping induced by dwarfing
root stocks in the following year.
IV. Earlier termination of leaf production, earlier leaf senescence and
abscission of spur leaf.
10. Reduced net assimilation rates : Trees on dwarfing root stocks have
lower net assimilation rates per unit leaf area than those on vigorous root
stocks. It has been found that the balance between photosynthesis and
respiration were found to be higher for M16 than for M9 suggesting early
leaf senescence and low proportion of young leaves with high
photosynthetic rates (Wunche et al. 1996).
11. Precocious and heavy fruiting at the expense of vegetative growth : At
maturity trees on dwarfing root stocks show a much higher ratio of fruit
yield to vegetative growth increment than those on vigorous root stocks.
This heavy fruiting checks the growth in the cropping year especially the
root growth and also the growth in the following year.
12. Reduced carbohydrates/ phloem transport from leaves to roots by
dwarfing root stocks or inter stocks : Dwarfing root stocks have been
found to block the transport of photosynthates (carbohydrates etc.) from
leaves to roots.
Effects of root stock on the scion cultivar :
(i) Size and growth habit : In citrus, pear and other species the most
significant effect of root stocks is tree size control e.g in apple root
stocks produce a complete range of tree size from dwarfing to very
vigorous (Gardner and Horarnic, 1963). In citrus root stocks (F and
A418 and No. 23) it was observed by Liiso et al. (2004) that dwarfing
mechanism is regulated by enhanced reproductive development and fruit
growth resulting in reduced vegetative development. Thus a change in
260 Fruit Tree Physiology

patteren of assimilate distribution appears to be one of the main

components of dwarfing mechanisms.
The mechanisms are complex and poorly understood, but a common
hypothesis is that the root stock that reduces scion vigor have low
hydraulic conductance (Syvertsen and Graham, 1985) which reduces
water transport to the shoots, ultimately decreasing stomatal conductance,
photosynthesis and shoot growth for a given investment in root biomass.
The effect of root stocks on water and mineral transport to the shoots,
growth regulator signals and the direct influence of the graft union on
phloem and xylem transport are the mechanisms through which root
stocks induce dwarfing (Atkinson et al; 2003).
Use of dwarfing root stocks also reduces scion shoot growth and the
longer the interstock the greater is the effect (Parry and Rogers, 1972).
When height of budding on the stem or shank of a dwarfing root stock
is increased it also leads to scion dwarfing (Parry, 1986). Differences in
stem xylem or phloem anatomy and function, the production of inhibitors
or the inactivation of promoters within this root stock/ interstock stem
piece are some of the mechanisms that lead to dwarfing e.g in sweet
cherries (Prunus avium) and plums (Prunus demestica L.) the dwarfing
root stock Pixy reduced the vigour of European plum (prune) trees by up
to 50 per cent (Webster, 1980) but had only a very small effect on scion
vigour when used as an interstock indicating that the dwarfing effect is
largely attributable to root rather than shank (root stock stem) effects in
cherry and plum root stocks.
Apple : The root stock effects were evaluated in apple by Samad et al.
(1998) and found that the use of interstock (M9) in four apple cultivars
viz. Granny Smith, Cox’s Orange, Gala and Oregon Red Delicious on
root stock MM106, M793 or Northern Spy reduced shoot growth by about
20 per cent. Thus interstock bridge grafting can be used as a viable
method of reducing growth in apple. The root stocks affect vigour of
scion cultivars grafted on them according to their potential. Apple clonal
root stocks have been classified into four groups i.e dwarf, semi-dwarf,
vigorous and very vigorous. The vigour potential of root stocks is related
to stomatal density in leaves (Beakbane and Majumdar, 1975., Pathak
et al., 1976) which is low in dwarfing root stocks and high in vigorous
root stocks (Rana, 1979., Jindal & Rana, 1984).
Root stocks like M9 and M27 are dwarf and produce smallest trees,
whereas trees on M25 are vigorous and attain significantly more height,
trunk girth and annual shoot growth than those on MMl06 root stock.
 Physiology of Dwarfism in Fruit Plants 261

Among the M7, M9 and MM111root stocks, M9 root stock induced the
minimum scion growth in terms of shoot length, trunk girth and shoot
dry weight, M7 was found to be intermediate between M9 and MM111.
Similarly, EMLA106 recorded maximum growth in terms of tree height,
spread, volume, and trunk girth while EMLA 26 had minimum growth
(Al Hinai and Ropar, 2004).

● ● ●

Fig. : Apple root stocks.

P.22 Mark M.9 EMLA Ott.3 M.7

M.9 337 M.9 PAJ. 2 M. 26
Bud.9 M.9 RN. 29
G. 65 G. 16 G. 11 G. 30*
Fig. : Cornell Geneva apple root stocks (http://www.ces.uga.edu/pubcd/b949-w.html).

Pear : The standard dwarf root stock for pear is Quince. Quince A is
standard root stock while Quince B is semi vigorous and Quince C is
very dwarf. Rivalta et al. (1986) observed that trees of William’s Bon
Chretien were the smallest on EMC and ADAMS, of medium size on
BA29 and EMA and the largest on Ct.S. 214 and Ct. S. 212. The own
rooted William’s showed greatest trunk diameter, while it was least on
BA29 after seven years of growth (Urbina et al; 2003). Maas (2008)
while testing different root stocks found that Quince root stock C 132
shows promise because of its control of tree growth in combination with
good fruit size and reduction of fruit russeting in Conference.
262 Fruit Tree Physiology

Fig. : Pear root stocks

100% 80% 70% 60% 50% 40% 30%

Standrd / Seedling <<<<<<<<<<<<Semi-Dwarf range>>>>>>>>>>>>>><<Full dwarf>

(http://www.ces.uga.edu/pubcd/b949-w.html).

Peach : The dwarfing root stocks used for peach are Siberian C, St
Julien X, P. besseyi and Rubira. The trees growing on Red Leaf lines
produce larger trunks than on Lovell. Three dwarfing root stocks, Inmil
(GM9), Damil (GM69/ l) and Camil (GM79) which were much more
dwarfing than F12/1 were produced through breeding and selection
(Trefois, 1985). In peach, trees grafted on dwarfing root stocks, Murase
et al. (1990) noticed shorter internodes as compared to trees on the more
vigorous root stocks. Weibel (2003) observed that the trunk cross sectional
area and growth of two peach cvs. Flavorcrest and Loadel on two small
root stocks (K-146-43 and K-146-44) was only 25-37 per cent of the
growth of trees on Nemaguard while tree on Hiawatha, P-30-135 and K-
119-50 root stocks provided an intermediate level of size control. European
root stocks that are mildly dwarfing include Ishtara (70%), Rubira (90%),
 Physiology of Dwarfism in Fruit Plants 263

Tetra (90%), Adara (80%) etc and semidwarfing ones like Pumiselect
(60%), Sirio (60%), Adesolo (60-80%) etc. (Layne and Bassi, 2008).
Plum : Among a wide variety of root stocks used in plum Prunus inistita
and their clones are dwarfing in nature like St. Julien A, St. Julien K and
St. Julien GF 655-2. The plum trees on Marianna plum were found to be
most vigorous as compared with trees on Okinawa and Nemaguard peach
seedling root stocks as reported by Starsen and Bester (1981). Dixy a
dwarfing root stock for both plums and gages (Prunus domestica) produces
trees half to two third the size of trees on St. Julien. Colt (P. avium x P.
pseudocerasus) - a hybrid root stock is of intermediate vigour producing
trees half to two third size as compared to trees on Fl21l. Beach Plum
(Prunus maritime) was classified as a very dwarfing root stock by Okie
(1987).
Cherry : The main root stock for sweet cherry (Prunus avium L.) and
sour cherry (Prunus cerasus L.) are seedling or clonal selection of
P. avium and P. mehaleb throughout the world. Under irrigated conditions
colt (P. avium X P. pseudocerasus) a dwarfing root stock may be used
(Singh et al; 1971) while new hybrid root stock of M x M (P. avium X
P. mehaleb) series which are semi dwarf to dwarfing are very useful
under rainfed conditions of north western Himalayas (Westwood et al;
1976). Stockton Morello- a clone of P. cerasus is best dwarfing root
stock and F121l is a vigorous root stock for sweet cherry (Parnia et al;
1982). Oppenheim (P. fructicosa X P. cerasus) and eM clones (P. incisa
X P. serrulata) are some other examples of dwarfing and precocious root
stocks. The Giesela root stock series produces true dwarf trees 40 to 70
per cent smaller than standard trees. Giesla 5 is hardy and high yielding
root stock producing over forty pounds of high quality cherries as early
as the first year. Gisela 5, producing a tree that is 45 per cent standard
size, Gisela 6, 70 per cent, and Gisela 7, 50 per cent (Riker, 2009). PIKU
root stock series like 1, 3 and 4 which are hybrids between different
Prunus species also impart dwarfness to cherry with varying degrees.
Mango : To achieve dwarfness in mango following criterion are essential
(i) high bark percentage (ii) small area of xylem vessels (iii) lower
stomatal density. Olour is a dwarfing and productive root stock for
Himsagar, Langra and Alphonso. The cultivars Kalapady, Olour and
Ambalavi, Vellaikollumban and Belkhas and Parikhas too have potential
for imparting dwarfness to scion. Use of interstock has been found to
produce promising results in regulating the tree height (Avila et al; 2010).
The nucellar seedlings of mango cv.Vellaikolamban were found to be
dwarfing when used as a root stock for cv. Alphonso scions, while those
264 Fruit Tree Physiology

of cvs. Milvandan, Bappakai and Olour behaved as vigorous root stocks

in decreasing order and when Alphonso was grown on this dwarfing root
stock there was more flowering and less vegetative flushing (Kurian
et al; 1996). The performance of Alphonso mango grafted on eight root
stocks showed that nucellar seedlings of Muvandan, Bappakai and Olour
were vigorous root stocks in decreasing order of vigour, while those of
Vellaikulamban imparted dwarfing, in comparison with the scion grafted
on its own open pollinated zygotic seedlings. The vigorous root stocks,
particularly Muvandan and Olour produced higher fruit yield per tree.
Olour also resulted in early higher yields of the scion. However, the
dwarfing root stock Vellaikulamban produced higher fruit yields per unit
canopy volume and per unit land area. From these studies, it was observed
that nucellar seedlings of Vellaikulamban and Olour are potential dwarfing
root stocks in mango. The performance of Dashehari was evaluated on
24 root stocks and it was observed that Ambalvi, Pahutan, Olour, Nakare,
Rumani, Willard, Moovandan etc. caused dwarfing effect (Litz, 2009).
Citrus : Tree size can be controlled through the use of tetraploid root
stock selections. A hybrid between grapefruit and trifoliate orange
(Citrumelo, 4475) was found useful as a dwarfing root stock for Valencia
orange in Florida (Gardner and Horanic, 1968) while Trifoliate orange
was found to be most dwarfing root stock for grapefruit, Washington
Navel orange and Valencia oranges. Root stocks like Rubidoux trifoliate
orange, Rusk citrange, Sweet orange x Rubidoux and Rangpur x Troyer
Citrange were found to be the most promising root stock for high density
planting of nucellar Marsh grapefruit and Valencia. When P. trifoliate cv.
Flying Dragon were used as root stock for lemon cv. Bnioneira 8A it
produced least vegetative development and the tree reached 3.5 m in
height after 12 years and produced more number of fruits (Foguet et al;
2000). Sonkar et al. (2004) observed dwarfing effect on growth in acid
lime budded on strain of Cleopatra mandarin (Coorg) root stock. Dwarfing
mechanism induced by the F&A 418 and No.23 root stocks is mediated
by enhanced reproductive development and fruit growth, resulting in
reduced vegetative development in the summer. One of the main
components of the dwarfing mechanism is change in the pattern of
assimilate distribution as suggested by Lliso et al. (2004).
Guava : Root stocks exert a great influence on vigour of the guava plants.
For commercial planting the Chinese guava (P. fredrichsthalianum) has
been used as dwarfing root stock in many orchards (Edward and Shankar,
1964). Teaotia and Phogat (1971) reported that P. pumilum stock produced
 Physiology of Dwarfism in Fruit Plants 265

dwarfing effect and small canopies when it was used as root stock. Dwarfing
of Allahabad Safeda in terms of plant height, spread and volume was
obtained by the use of potentially dwarf Aneuploid no. 82 root stock of
guava and yield/ unit volume was maximum. Several species of Psidium
such as P. cujavillis, P. molle, P. cattelianum and P. guineens can be suitably
used as root stock in India but the Chinese guava (Psidium
friedrichsthalianum) has been found to induce dwarfing effect and is
resistant to wilt disease and can be used for commercial planting.
Ber : Seedlings are mostly used as root stocks in ber, however, it can
also be propagated on some different root stocks like Boradi (Zizyphus
rotundifolia) and Jhar ber (Z.nummularia). Jhar ber genetically shows
dwarfism and variable degrees of incompatibility. Dwarf and spreading
trees were produced on Z. mauritiana (Coimbatore) whereas those on
Z. mauritiana (Dehradun) were found to be vigorous by Singh et al.
(1978). Bal et al. (1997) found that the plants on Z. mauritiana
(Coimbatore), Z jujuba and Z. numularia from Punjab, Rajasthan and
Haldwani were dwarf than mauritiana root stocks. Prasad et al. (2004)
reported that cv. Gola recorded maximum tree height spread and volume
with Z. rotundifolia followed by Z. mauritiana which is significantly
higher than Z. nummularia. Verma et al. (2001) studying the effect of
stomatal density on different stionic combinations of ber (Zizyphus
mauritiana) found significant differences in stomatal density, internodal
length and plant growth in plant samples comprising root stock-scion
combination of root stocks like Z. nummularia, Z. mauritiana ecotype-
291 and Z. mauritiana ecotype-Assam-Gauhati and commercial cultivars
like Banarasi, Karaka, Ponda and Gola. Gola budded on Z. nummularia
recorded the minimum stomatal density, internodal length and plant height
and was found the most dwarfing combination. Whereas Ponda budded
on root stock Z. mauritiana ecotype-Assam-Gauhati had maximum
stomatal density, internodal length and plant height and was found to be
most vigorous stionic combination.
Dwarfing root stocks in different fruit crops :
Apple
Super dwarfing : M27, Bud 57-491
Dwarfing : M9, M26, P2, MAC 9
Semi dwarfing :M4, M7, MM106
Pear : OH× F 51, Oregon 211, Oregon 249, OH× F 34, OH× F 87, BP-
1, Quince C
266 Fruit Tree Physiology

Stone fruits :
Almond : Adafuel bitter almond
Apricot : P. besseyi, Hyb. P 2038
Cherry : Colt, M× M14, Charger, CAB 6P, CAB 11 EW
Peach : Siberian C
Plum : Pixy, St. Julien K, GF43, P. maritima, P. besseyi,
P. cistena, P. tomentosa.
(ii) Interstock effects on scion and root stock : The use of a dwarfing
interstock such as M9, MM111 etc. with different lengths has been widely
used to produce dwarfing in apple trees. It allows the use of well anchored,
vigorous root stock rather than a brittle, poorly anchored dwarfing clone.
With the removal of the bark the flow of carbohydrates down to the root
is restricted and, therefore, carbohydrates accumulate above the bridge
producing differential effects on root development. This in turn, leads to
differential effects on shoot growth. The greater amount of carbohydrates
above the cut results in the diversion of carbohydrates to the upper portion
of the tree (Greene, 1937) which directly affects the reproductive status
of the tree. High carbohydrate level in the upper tree increases flower bud
initiation as high carbohydrate levels in the upper tree were not translated
to increased vegetative growth. Reduced vegetative growth, fruit size and
improved reproductive growth after interstock bridge grafting was also
observed by Samad et al.(1999). Dann et al.(1984) observed in peach that
severing the phloem reduced the flow of growth regulators to the roots,
which in turn reduced the production of root produced gibberellins and
cytokinins. Cutting and Lyne (1993), also supported the hypothesis that
less cytokinins and gibberellins above the girdle are likely to reduce
meristematic activity and cell elongation, leading to a reduction in
vegetative growth. Dwarfing root stock genotype used as an inter stem
function as a kind of physical restriction to water movement, reduce
potential xylem flow and hydraulic conductance of the whole tree. The
interstem effect also depends on its length and doubling the length halves.
The hydraulic conductance e.g a 35 cm inter stem piece produced a greater
dwarfing effect than a 5 cm inter stem in apple (Parry and Rogers, 1972),
and similar effects were reported by Di Vaio et al. (2009).
B. Use of bioregulators :
1. Paclobutrazol : It inhibits the GA synthesis and thereby retards growth,
shortens elongation and reduces the intermodal length. Dwarfing
mechanism due to paclobutrazol is based upon:
 Physiology of Dwarfism in Fruit Plants 267

(i) Shorter internodes due to less GA in the tissues resulting from the
action of paclobutrazol.
(ii) Reduces ABA levels in the shoot tips (Kurian et al. 1993).
(iii) Reduces the levels of cytokinins in treated trees (Kurian et al.
1993).
(iv) Enhances the total phenolic content of terminal buds and alteres the
phloem to xylem ratio of the stem (Kurian and Iyer, 1993c).
(v) Paclobutrazol treated sugar beet leaves were thicker, smaller and
darker due to increase in the chlorophyll concentration per unit area
thus accounting for higher rate of photosynthesis.
(vi) Reduction in the lateral growth resulting in decreased whole plant
CO2, uptake due to reduced total leaf area.
(vii) Reduction in trunk cross section area.
(viii) Alters endogenous growth promoters and inhibitors, besides altering
xylem and phloem tissues thereby restricting vegetative growth and
enhancing flowering by altering assimilates partioning and patterns
of nutrient supply for new growth. Examples :
(a) Paclobutrazol (15 mg/ litre) significantly reduced shoot length
in plum, sour and sweet cherries, apricot and pear trees and
TIBA (3.5 mg/ tree) in plum and sour cherry respectively, for
three consecutive years (Grochowska and Hodun, 1997).
(b) Paclobutrazol @ 10 g a.i. for 5 years old and 1.25 and 2.5g per
litre for 1 and 2 year old grafts was safe and effective in 3
mango cultivars (Kulkarni, 1988).
(c) Paclolutrazol @ 2.5 or 5.0 g/l tree resulted in 50 per cent
reduction in tree volume expansion with no phytotoxicity in
mango Alphonso (Kurian and Iyer, 1993)
(d) PBZ inhibited length (21 %) and dry weight (19%) of the stem
whereas GA3 increased both (79 and 29% respactively) in the
absence or presence of PCZ in citrus root stock seedlings
(Mehouachi et al; 1996).
(e) Cycocel and etherel reduced the trunk diameter and tree volume
of Golden Delicious apple.
(f) Soil applied PBZ (8 mg a.i. tree) inhibited the growth of branches
and intensified green color of the foliage in ber plants.
(g) 2 sprays at 1000 ppm, one at petal fall and one 2-3 weeks later
suppresses vegetative growth and increaes fruiting on pears.
268 Fruit Tree Physiology

2. Alar :
(i) The frequency of mitotic figures in the shoot apex decreased
significantly in apples treated with Alar. The rate of penetration of
Alar and its effectiveness in altering the rate of gibberellin synthesis
may account for rapid mitotic activity.
(ii) It affects the auxin and diamine oxidases forming the basis for growth
inhibition.
(iii) It lowers the membrane integrity allowing vacuolar contents to diffuse
rapidly into the external medium.
Ethephon at higher concentration (500-3000 ppm) effectively reduces
plant height and increases flower production in guava and the density of
27000 plants/ ha through the use of various growth retardents has been
achieved. Paclobutrazol @ 2000 ppm produces dwarfed mangoes. Okuda
et al. (1996) observed that paclobutrazol @ 500 and 1000mg application
significantly reduces the vegetative growth in Satsuma mandarin.
Paclobutrazole applied through foliar spray as well as soil drenching to
the pollarded trees of guava cv. Round and L-49 produced a positive
response for tree size control and fruit yield by reducing competition
from vegetative growth. Reduced vegetative growth, induction of
flowering and increased yield in mango has been obtained by
paclobutrazol application. Soil application @10 g a.i./ tree gave the desired
effects for at least 2 years in 5-year-old bearing trees whereas in 1- and
2-year-old trees, 1.25 and 2.5 g a.i./ trees, respectively were safe and
effective (Kulkarni, 1988).
The dwarfing effect is complex and most likely regulated by a number
of signaling pathways acting in tandem and can not be readily explained
in terms of changes in individual signaling molecules. Plant growth
retardants are applied in horticultural crops to reduce unwanted
longitudnal shoot growth without lowering plant productivity. 0.5 - 1 per
cent NAA in 40 per cent white latex paint solution or NAA as 1 per cent
asphalt based aerosol significantly reduced the number of sprouts in
Delicious apple trees (Miller and Ware, 1980). Marini and Barden (1982)
reported that 1 per cent NAA to summer pruning cuts reduced regrowth
in young apple trees. Brenner (1973) obtained successful control of root
suckers with a mixture of 2 per cent benefin + 0.25 per cent NAA.
The application rate of NAA was increased to 0.5 per cent in case of
pear.
 Physiology of Dwarfism in Fruit Plants 269

3. Gibberellin biosynthesis inhibitors :

1. Onium compounds : They block the cyclases copalyl - diphosphate
synthase and ent-kaurene synthase involved in the early steps of GA
metabolism. The inhibition of the cyclization of geranyl geranyl
pyrophosphate to copallyl pyrophosphate is the primary mode of
action of onium compounds leading to inhibition of GA formation
e.g cycocel, mapiquat chloride, AMO-1618, phosphon D and
piperioxim bromide.
2. Triazoles : They inhibit the microsomal oxidation of kaurene, kaurenol
and kaurenal which is catalyzed by kaurene oxidase, thus reducing
the plant growth e.g Paclobutrazol, uniconazole, tripenthenol, BAS
111 and XE1019.
3. Pyrimidines : They inhibit cytochrome PASO which controls the
oxidation of kaurene to kaurenoic acid. They inhibit gibberellin
biosynthesis and also interfere with sterol and abscisic acid
biosynthesis.
4. Others :
1. Tetcyclacis - They reduce gibberellin biosynthesis by blocking
microsomal oxidation of kaurene to kaurenoic acid.
Inabenfide - It also inhibits GA biosynthesis by blocking the oxidative
conversion of kaurene to kaurenoic acid.
2. Growth retarding compounds not involved in inhibiting gibberellin
biosynthesis include: abscisic acid, cimetacarb, daminozide, dikegulac,
maleic hydrazide, mefluidide.
3. Morphactins: chlorfluron, chlorflurenol, dichorflurenol and flurenol.
They have the ability to affect plant morphogensis. They act as
competitive antagonists of GA.
C. Use of incompatible root stocks : Dwarfness can also be imparted by the
use of incompatible scion and stock. However, it is not commercially exploited
for this end e.g Jhar ber produces genetic dwarfism and shows variable
degrees of incompatibility i.e. inverted bottleneck formation at bud union
and induces dwarfness and can be exploited for high density planting (Verma
et al; 2000). When pear is grafted on quince similar effects are produced
(Francescatto et al; 2007).
D. Induction of viral infection : Dwarfing can be induced by inducing viral
infection, although, it is not commercially adopted e.g citurs, apple. The isolated
cultures containing cachexia pathogenic variant of the HSVd viroid (CVd -
lIb) forming a complex with CV d-W or with exocortis (CEV d) induced
270 Fruit Tree Physiology

shortening of shoots and reduction of the canopy diameter, height and volume
with respect to healthy trees of Persian lime and inoculated trees showed higher
production efficiency. East MaIling Long Ashton (EMLA) series in apple are
virus free and vigorous than their infected counterparts. Dwarfing produced by
viroids can be considered a disorder, however, these dwarf trees are healthy in
appearance and produce yields comparative to or better than healthy trees.
Viral infection, induces dwarfism has been reported by many workers (Manes;
2004., Nishida and Sugiyama; 1993., Crespi et al; 1992).
Viroid inoculated trees may give high yields while restricting tree size in
high density plantings. They reduce scion trunk cross sectional area, tree
height, canopy volume and yield and produce no effect on cropping efficiency
or yield efficiency. Such trees have smaller summer and autumn growth
flushes, fewer spring shoots, less annual vegetative growth and are less
damaged by frost. The timing and duration of flushes is unaffected. The
timing and intensity of flowering, ratio of leafy to leafless inflorescences etc.
are unaffected. Effects on canopy development and tree performance are not
pronounced until 4–5 years (P. trifoliate) or 7-8 years (citrange) after
inoculation, while effects on yield are delayed for further 1–2 years. The
extent of the dwarfing response is influenced by the time of viral inoculation
and earlier it is done the greater will be the dwarfing response. However, in
late inoculated trees (5-6 years after planting) the reduction in tree size is not
apparent. Infected bud or bark from the inoculated bud stick can be used for
induction of infection, however, bark shield is preferred, because no subsequent
removal of shoots is required. Inoculation is done within 6–18 months of
field planting in citrus and dwarfing effect is observed after a period of 18
months. However, late (4 or 5 years after planting) inoculation produces little
effect on tree size control. Further double bud is prefered over single bud to
have better infection.
E. Pruning and training : Dwarfing can be achieved through pruning and it
has been observed that slow growing trees respond more. To reduce excessive
shoot growth and promote flowering and cropping manipulation of roots has
been practised since long time e.g figs have been grown in pots for about
2000 years because of the restriction of root growth by the container which
makes them to fruit more profusely. To propagate recalcitrant rooting clonal
scions root stocks have been used for many centuries. Selective breeding and
development programs have produced many apple root stocks that are able
to with stand various biotic and abiotic stresses. The primary targets for root
stock breeding are: inducing dwarfness, enhancing the cropping efficiency
(yield per tree size) and precocity. Work at HRI-East Malling showed that
root stocks influence the scion and pruning and training are used to maintain
 Physiology of Dwarfism in Fruit Plants 271

at a given size and shape without sacrificing yield. A compact and bushy tree
growth is achieved by removal of apical portion which stimulates lateral bud
growth. Tree size control through pruning is limited to grape, apple and some
other temperate crops. Lauri and Claverie (2005) proposed training proposals
in cherry based on two principles i.e maintaining the natural hierarchy between
the trunk and side branches and controlling the growth of the trunk and the
branches by bending rather than by heading cuts. Solaxe training system
developed initially for apple is also based on these principles. In traditional
systems, such as Goblet, the balance of fruiting to vegetative growth is
obtained by annual repeated heading or thinning cuts, usually during winter
because of the greater time available during this period resulting in marked
vegetative reiteration mechanisms, which may generate an imbalance between
growth and fruiting.
Of the various training systems being followed in apple e.g spindle bush
and SVA (Single Vertical Axis) raised on M9, M7 and M4 root stocks has
been found to be promising for HDP w.r.t size control and higher yields per
unit land area (Gyuro, 1978).
F. Nutrients : Moderately vigorous trees utilize less nitrogen (Whitney apple)
while very vigorous ones use more nitrogen (Snow and York Imperial).
G. Phenols : A number of phenolic compounds reduce plant growth e.g coumarin.
Phenols inhibit growth through inhibition of mitosis, cell division, cell
elongation and increased oxidative decarboxylation of IAA. Another phenol
phloridzin stimulates IAA oxidase and is present in large amounts in the bark
of dwarfing root stocks. They also retard IAA synthesis. Some phenols also
inhibit translocation of sugars and auxin or act by regulating the polar auxin
transport.
H. Invitro techniques : Seeds of apple cultivar USDA4-20 were subjected to
different levels of (0, 20, 40 and 60 muM) BA (N6 Benzyl adenine) and TDZ
(Thidiazuron) to induce dwarfism via in vitro technique. It was observed that
TDZ was more effective chemical in inducing dwarfism than BA. The plants
produced by TDZ and BA were slower in growth and dwarf in size than non-
treated plants (Khattak et al; 2004).
I. Genetic engineering : A cDNA encoding sorbitol-6-phosphate dehydrogenase
(S6PDH), which is a key enzyme in sorbitol biosynthesis introduced into the
Japanese persimmon (Diospyros kaki) induced dwarfism. The physiological
mechanism behind this dwarfing effect has been attributed to accumulation
of sorbitol which might have caused an osmotic imbalance between the
cytosol and vacuole (Deguchi et al; 2004). Various genes have been identified
in fruit crops that can induce dwarfism e.g ipt, OSHI, rolABC etc. that
272 Fruit Tree Physiology

induce compactness and bushy appearance in fruit crops like apple, kiwi,
peach etc. In apple reduction in stem length, internode length and node
number was achieved through overexpression of gaai gene isolated from
Arabidopsis (Zhu et al; 2008).
J. Others :
● Proteins, tengu-su inducer (TENGU) secreted by bacterium phytoplasma
it inhibits auxin related pathways thereby affecting plant development
(Hoshi et al; 2009).
● Down regulation of GA synthesizing genes e.g reduction of CND41
proteins reduces the GA levels.
● Dwarfing through mutations induced by irradiation e.g a dose of 2-kR
γ-irradiation applied twice induced dwarfism in Irwin cv. of mango seedlings
and the survival rate was 65 percent (Yitzu and Loonshung, 2000).

CONCLUSIONS
The mechanism underlying dwarfing is obscure, however, it involves anatomical,
physiological and biochemical changes. Dwarfing in fruit crops can be achieved
through various approaches like selection of spur type scion cultivars, use of
interstocks, pruning, root pruning, use of growth retardants, control of nutrient
elements, girdling, scoring, bark inversion, use of dwarfing/ ultra dawrfing/ semi
dwarfing root stocks. Growth retardants like paclobutrazol are most effective and
widely used growth retardant which acts through inhibition of GA biosynthetic
pathway. It acts as the potential growth retardant and effectively controls canopy
volume increment and canopy structure. Although these chemicals provide effective
control of growth as well as increasing fruit yields but they have extreme
persistence in the soil and hence not approved for use in many countries. Thus,
long-term future use of growth retardants in fruit production and consumer opinion
needs to be evaluated. Recently the recombinant DNA technology has widen the
gene pool which can be manipulated to induce dwarfism and harvest maximum
benefits in horticultural crops e.g the GA20 oxidase gene has been inserted into
the Greensleeves variety of apple to produce dwarf trees that are dwarf but
identical to the parent in every respect.

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 11

FRUIT RIPENING – BIOCHEMISTRY,

PHYSIOLOGY AND REGULATION

R ipening is a physiological process that transforms a physiologically mature

but inedible plant organ into a visually attractive olfactory taste sensation. It
marks the completion of development of a fruit and commencement of senescence.
Fruit ripening is a highly co-ordinated, genetically programmed, and an irreversible
phenomenon involving a series of physiological, biochemical, and organoleptic
changes that lead to the development of a soft and edible ripe fruit with desirable
quality attributes. A wide spectrum of biochemical changes such as increased
respiration, chlorophyll degradation, biosynthesis of carotenoids, anthocyanins,
essential oils, and flavor and aroma components, increased activity of cell wall
degrading enzymes, and a transient increase in ethylene production are some of
the major changes involved during fruit ripening (Brady, 1987).
The metabolic changes during fruit ripening include increase in biosynthesis
and evolution of the ripening hormone, ethylene (Yang and Hoffman, 1984),
increase in respiration mediated by mitochondrial enzymes, especially oxidases
and de novo synthesis of enzymes catalyzing ripening specific changes (Tucker
and Grierson, 1987). Alteration of cell structure involves changes in cell wall
thickness, permeability of plasma membrane, hydration of cell wall, decrease in
the structural integrity, and increase in intracellular spaces (Redgwell et al; 1997).
The major textural changes resulting in the softening of fruit are due to enzyme
mediated alterations in the structure and composition of cell wall, partial or complete
solubilization of cell wall polysaccharides such as pectins and cellulose and
hydrolysis of starch and other storage polysaccharides. The changes in gene
expression during ripening involves the appearance of new ripening specific mRNAs,
tRNA, rRNA, poly A+RNA, and proteins, and the disappearance of some mRNAs
(Wong, 1995). These changes during fruit ripening are activated by plant hormones.
 Fruit Ripening – Biochemistry, Physiology and Regulation 279

Based on their respiratory pattern and ethylene biosynthesis during ripening,

fruits can be classified as climacteric and non-climacteric type. Climacteric fruits,
harvested at full maturity, can be ripened off the parent plant. The respiration rate
and ethylene formation though minimal at maturity, rise dramatically to a
climacteric peak, at the onset of ripening, after which it declines (Gamage and
Rehman, 1999).
Non-climacteric fruits are not capable of continuing their ripening process,
once they are detached from the parent plant. Also, these fruits produce a very
small quantity of endogenous ethylene, and do not respond to external ethylene
treatment. Non-climacteric fruits show comparatively low profile and a gradual
decline in their respiration pattern and ethylene production, throughout the ripening
process (Gamage and Rehman, 1999).

Climacteric Non-climacteric

1. They show an increase in respiration 1. They do not show respiratory climacteric.

during ripening i.e. respiratory
climacteric.
2. Produce high amount of ethylene 2. Produce less amount of ethylene during
during ripening. ripening.
3. Internal ethylene concentration varies 3. Internal ethylene concentration changes
widely. little during development and ripening.
4. Ripening may occur even after 4. Compelete ripening occurs on the tree.
detachment.
5. External application of ethylene 5. External application of ethylene merely
(0.1 to 1.0 ul/ L for one day) hastens causes a transient increase in respiration
ripening but the magnitude of but magnitude of respiration is depen-
climacteric is relatively independent dent on the concentration of ethylene.
of the concentration.
6. Ethylene application causes a single 6. Rise in respiration may occur more than
increase in respiration. once.

Classification of Fruits

Climacteric fruits Non-climacteric fruits

Apple Cherry
Apricot Cucumber
Banana Grape
Guava Grapefruit
Kiwifruit Lemon
[Table Contd...
280 Fruit Tree Physiology

Contd. Table]

Climacteric fruits Non-climacteric fruits

Mango Lime
Papaya Litchi
Passion fruit Mandarin
Peach Melon
Pear Orange
Persimmon Pineapple
Plum Pomegranate
Sapodilla Raspberry
Tomato Strawberry

Senescence

Fig. : Growth, respiration and ethylene production in climacteric and non-

climacteric fruits (Wills, 2007).

Climacteric ripening: In many fruits, ripening is associated with sudden and

dramatic increase in ethylene biosynthesis as well as a rapid increase in respiration.
This sudden increase or upsurge is called as climacteric rise in respiration and the
fruits are refered as climacteric fruits (examples listed above). The rise in the rate
of respiration depends on species and cultivars. Respiration is a catabolic process
and results in breakdown of complex materials such as starch into simpler molecules
along with production of carbondioxide and energy. Because of dynamic nature
of ripening in several species, it is likely that endogenous control system also
 Fruit Ripening – Biochemistry, Physiology and Regulation 281

function to initiate ripening at the proper stage of ontogeny. Climacteric is therefore

a period during which certain biochemical changes are initiated but autocatalytic
production of ethylene, marking the changes from growth to senescence and
involving an increase in respiration leading to ripening of fruits (Rhodes et al;
1981). Some fruits such as avacados do not ripen while attached to trees and
gradually increase their sensitivity to ethylene with time after harvest (Biale,
1975). Similarly, pear develops the best organoleptic characteristics when harvested
mature green and ripened off the tree. The respiratory curve of climacteric fruits
at a suitable temperature shows a decreasing trend to the lowest value termed as
pre climacteric minimum followed by more or less sharp rise in respiration called
climacteric peak (depending upon species and or cultivars) and subsequent post
climacteric decline in the rate of respiration during fruit senescence. Ethylene
levels at the harvest influence the magnitude of climacteric curve and the final
product quality (Lalel et al; 2006). Fruits harvested too early do not undergo the
desired ripening changes and a late harvest will lead to off flavor and reduced
shelf life. In apple the rise in respiration occurs in fruits both detached and
attached form whereas avocado fruits shows climacteric rise in respiration only
after detachment from the tree. In climacteric fruits, ethylene generally regulates
fruit ripening by coordinating the expression of genes responsible for 1) enhancing
rise in the rate of respiration, 2) autocatalytic ethylene production,
3) chlorophyll degradation, 4) pigment synthesis, 5) conversion of starch to sugars
6) production of aroma and 7) increased activity of cell wall degradation.
Non- climacteric ripening: There is no distinctive and well co-ordinated
pattern of ripening in these fruits as in climacteric ones. These fruits show neither
a peak in respiration nor an associated production of ethylene during the ripening
process. The ripening is normally completed on the parent plant itself and fruits
normally contain no or very little starch e.g cherry, citrus, grapes, pineapple,
litchi, olive etc.

MECHANISM OF RIPENING
The mechanism by which ripening is regulated and co-ordinated is not clear.
However, two hypothesis have been proposed to explain the same. According to
one hypothesis, ripening is considered as an ultimate result of progressive increase
in cell permeability leading to increased contact between enzymes and substrate
already present in the tissues. The second hypothesis suggests that ripening involves
denovo protein and RNA synthesis. It also suggests that ripening may be the
result of both these processes.
282 Fruit Tree Physiology

There is no clear metabolic difference of the climacteric and non-climacteric

fruit ripening. The ripening changes occur from the last stages of growth and
development before senescence and results in characteristic food quality (Watada
et al; 1984). There is an indisputable evidence that ethylene is involved in induction
of ripening. In several fruits, measurable concentration of endogenous ethylene
is reported during development and maturation.
The treatments that decrease the level of ethylene also retard ripening such
as:
1. Hypobaric condition, where the partial pressure of O2 is maintained at the
normal atmospheric level, but the internal ethylene concentration is reduced.
In mango cv. Okrang at 60-100 mm Hg and 13°C was found effective for
long term storage (four weeks) with no adverse effect upon removal to
normal pressure.
2. Storage at low O2 concentration, where ethylene synthesis is inhibited.
3. Storage at high CO2 concentration, which inhibits ethylene.
4. Treatment with auxins inhibits ethylene synthesis and retard ripening.
5. AVG (Aminoethoxyvinyl Glycine) and AOC (Aminooxy acetic acid).
The retardation of ripening could be reversed by the addition of small amount
of ethylene. Exogenously applied ethylene above a threshold level induces ripening
and simultaneously endogenous ethylene production. The climacteric and non-
climacteric fruits are distinguishable by their response to treatment with ethylene.
Ripening can be considered as a specialized stage of plant senescence and
takes place after certain stage of maturity has been reached. Major compositional
changes occur in fruits during ripening and these affect their attractiveness, storage
life and nutritional value. These changes are a consequence of alterations in the
physiological and biochemical processes occurring in the cells of the fruit. Most
fruits undergo similar changes that may include seed maturation, degradation of
cholorophyll and colour change, abscission, change in respiratory rate, change in
rate of ethylene production, change in tissue rigidity, change in composition of
pectic substances, change in carbohydrate composition, change in organic acids,
change in proteins, production of flavor volatiles, development of wax on skin
etc. Generally, the fruits displaying the highest respiration rates tend to ripen
faster and are more perishable. Non climacteric fruits such as citrus display a
gradual decline in respiration during ripening. There is loss of reserve food during
respiration and it indicates that during climacteric rise the senescence is hastened
as the stored energy reserves are used up, loss in the energy value of the food for
the consumer and flavor quality especially sweetness is reduced.
 Fruit Ripening – Biochemistry, Physiology and Regulation 283

Ethylene biosynthesis and ripening: Ethylene, a fruit ripening phytohormone,

in minute amounts can trigger many events of cell metabolism including initiation
of ripening and senescence, particularly in a climacteric fruit.
The pathway for ethylene biosynthesis has been elucidated in apple, and
other fruits such as avocado, banana, and tomato (Ke D, 1993; Yang and Hoffman,
1984). The first step is the conversion of S-adenosylmethionine (SAM) to 1-
aminocyclopropane carboxylic acid (ACC) by the enzyme ACC synthase (Fig.).
At the onset of fruit ripening, expression of multiple ACC synthase genes are
activated, resulting in increased production of ACC. In most cases, it is the ACC
synthase activity, which determines the rate of ethylene biosynthesis. ACC is then
oxidized to ethylene by ACC oxidase. Inhibition of ethylene biosynthesis by
antisense RNA for ACC synthase and ACC oxidase was demonstrated first in the
tomato fruit. Deamination of ACC to α-ketobutyrate by over expressing ACC
deaminase enzyme also inhibited ethylene formation and fruit ripening. The
resultant transgenic fruit did not over ripe as normal controls, though some color
change occurred and a mere ethylene boost triggered back all the ripening related
biochemical changes in a similar way as in normal fruit. Recently the cDNA
encoding for ACC oxidase has been isolated and characterized from mango (Zainal
et al; 1999). Down regulation of ACC synthase and ACC oxidase genes in mango
are now being used for extending the shelf life of this fruit.

Fig. : Pathway for ethylene biosynthesis and metabolism.

Changes Occurring During Ripening

The various changes occuring during ripening of fruits are: Seed maturation,
degradation of cholorophyll and colour change, abscission, change in respiratory
rate, change in rate of ethylene production, tissue rigidity, composition of pectic
substances, carbohydrate composition, organic acids, proteins, production of flavor
volatiles, development of wax on skin etc.
284 Fruit Tree Physiology

Some of the important changes which occur during ripening are discussed
here:
1. Degradation of chlorophyll and pigment synthesis : The change from a
green mature fruit to a ripe apple, mango, and banana involves a transition
of chloroplasts into chromoplasts. Thylakoid membranes and chlorophyll
pigments are broken down, and there is a progressive accumulation of new
carotenoid pigments in the plastids or anthocyanin in vacuoles. Ethylene
treatment in immature climacteric fruits accelerates the onset of climacteric
and associated ripening changes without appreciably changing the pattern or
magnitude of respiration. However, in non-climacteric fruits, it leads to
increase in respiration and ethylene production and the magnitude of response
depends on concentration of ethylene applied. In climacteric fruits, if ethylene
is applied over a sufficient period, no return to preclimacteric will result on
the removal of gas, but in non-climacteric fruits, removal of ethylene at any
time will result in a return of the rate of gaseous exchange to the level of
untreated fruits.
Various ways through which chlorophyll get degaraded and leads to
synthesis of other pigments are as:
1. Degradation of chlorophyll due to chlorophyllase enzyme.
2. Splitting of chlorophyll into phytol chain and porphyrin.
3. Loss of Mg++ ion and conversion of porphyrin into phaeophytin.
4. Change in tetrapyrolic chain and it becomes bilviridin.
5. Oxidation or saturation of double bonds.

Fig. : Pathways of chlorophyll degradation

 Fruit Ripening – Biochemistry, Physiology and Regulation 285

In fruits of early ripening cultivar, Wase Satsuma mandarin, the peel is

green when the flesh matures and fruits attain harvest maturity. Ethylene is
used for the enhancement of peel degreening and chlorophyll content decreased
greatly with a concomitant increase in chlorophyllase and chlorophyll
peroxidase activities (Yamauchi et al; 1997). Thus, chlorophyllase and
chlorophyll peroxidase could be involved in chlorophyll degradation of
ethylene treated Satsuma mandarin. Wase Satsuma mandarin matures on
trees without ethylene treatment.
2. Hydrolysis of starch to sugars : During ripening, fruits show an increase
in the concentration of sugars, either by hydrolysis of starch within the fruit
(e.g banana, mango, kiwi fruits etc.) or by continued import of sugars from
other part of the plant (e g. that ripen on the vine, melon) depending on
whether the fruit ripens on or off the plant. The predominant sugar in fruits
are sucrose, glucose and fructose. An increase in sugars content makes the
fruits sweeter and palatable. In climacteric fruits, the starch content generally
increases during development but sucrose decreases during storage due to its
usage for metabolic processes. In case of non-climacteric fruits quality is
poor if harvested at the onset of ripening than those if allowed to ripen on
the tree.
Various starch mobilizing enzymes and their response.

Enzyme Mode of action

α-Amylase Hydrolysis α (1-4) linkage of amylase at random to produce

mixture of glucose and maltose.
β-amylase Attacks penultimate linkages and releases maltose
Starch phosphorylase Hydrolyses α (1-4) linkages to give glucose-1, phosphate which
can be coverted into glucose 6-phosphate by the action of glucose
phosphate mutase.
α (1-6) glucosidase Attacks α1-6 glucose linkage of amylopectin.

3. Textural softening : Textural change is the major event in fruit softening,

and is the integral part of ripening, which is the result of enzymatic degradation
of structural as well as storage polysaccharides. Depending upon their inherent
composition and nature, different fruits soften at different rates and to varying
degrees (Tucker and Grierson, 1987). Softening occurs due to breakdown of
starch and other non-pectic substances resulting in reduced cellular rigidity.
Due to changes in pectic substances and hydrolysis starch fruits loose firmness
and become soft. Fruits such as mango, papaya, avocado, sapota, and banana
undergo drastic and extensive textural softening from stone hard stage to a
soft pulpy stage, whereas apple and citrus fruits do not exhibit such a drastic
286 Fruit Tree Physiology

softening, though they undergo textural modifications during ripening. Fruit

ripening with special reference to textural softening has been diagrammatically
shown in Fig.

Fig. : Fruit ripening with particular emphasis on textural softening. Control points
at ethylene (1) and post-ethylene (2) levels.

The process of textural softening is of commercial importance as it directly

dictates fruit shelf life and quality (Tucker, 1993). This should be considered
to avoid mechanical damage during harvesting and transportation. The textural
properties of fruits in general play a very significant role in the consumer
acceptability.
4. Production of aroma volatiles : During ripening, there is increase in volatile
compounds that leads to characteristic flavor. The major chemical compounds
found are esters of aliphatic alcohols and short chain fatty acids. In fruits,
major volatile compounds are isoamyl acetate, aldehydes and terpenoid
compounds. The ripe fruit develops unique flavours that result, at least in
part, from increased synthesis of organic acids, principally citric acid but
also malic acid and volatiles increase during ripening and combine to produce
the unique flavour and aroma of the ripe fruit. Phenolics such as tannins
provide astringency to unripe fruit and have an important influence on the
flavour and colour of mature fruit. A complex mixture of alcohols, aldehydes,
 Fruit Ripening – Biochemistry, Physiology and Regulation 287

C1-C6 esterified acids, estragol and terpenes contribute to the flavour of

apples. Among these compounds, only 20-40 are directly associated with the
characteristic apple like aroma (Dixon and Hewett, 2001).
Characteristic aroma compounds of some fruits.

Product Compound(s)

Apple Ripe Ethyle 2-methyle butyrate

Green Hexanal, 2-hexanal
Banana Green 2-hexanal
Ripe Eugenal
Overripe Isopentanol
Grape fruit Nootaketone
Lemon Citral, Limonene
Orange Valencene
Pear Decadienecate ester
Grape Methyanthranilate, Ethyleacetate.

4. Cell wall degradation and softening of fruits : There is increase in soluble

pectic substances and decrease in insoluble ones and firmness. Two processes
are mainly involved:
(i) Depolymerisation (shortening of chain length) and
(ii) Deesterification.
When fruits become soft, there is change in insoluble pectic substances
into soluble forms. There is hydrolysis of starch or fats e.g in avocado.
Softening is an important part of fruit ripening process and it was documented
that changes in cell walls accompany fruit softening (Brummell and Harpster,
2001). Plant cell walls are highly complex structures consisting of many
types of complex polysaccharides and numerous proteins. Pectins,
hemicelluloses and cellulose comprising the major groups of polysaccharides
in the plant cell walls with more than 50 per cent as pectin. Cell wall
metabolism is tightly controlled through regulation of both gene expression
and cell micro environment. Due to structural complexity of the primary cell
wall, a coordination of a variety of enzymes affecting both covalent and non-
covalent bonds is likely involved in cell wall disassembly and fruit softening.
In ripening fruits, much attention was focused on the depolymerization
of acidic pectins by polygalacturonase. However, experiments with transgenic
tomatoes have shown that even though PG is important for the degradation
of pectins, it is not the sole determinant of tissue softening during ripening.
PG antisense constructs for various tomato lines have little effect on the fruit
288 Fruit Tree Physiology

characteristics viz. reduced susceptibility to cracking and decay and other

damages at the later stages of ripening. Now the focus is on the hydrolysis
of neutral sugar side chains, which may weaken the complex network of cell
wall polymers, thus contributing to textural softening.
De-esterification of polygalacturonic chains by pectin methylesterase
(PME) make the chains more susceptible to PG-degradation, facilitating rapid
loss of cell wall structure. Endo- -mananase has more recently been implicated
in cell wall degradation (Bewley et al; 2000). Koslannund et al. (2005)
classified pawpaw fruit into four categories on the basis of firmness and
respiration along with ethylene production rates at harvesting:
● pre-ripening (no to very less loss of firmness; pre-climacteric),
● early ripening (some softening; increasing rate of ethylene and CO2
production,
● mid ripening soft; at or just past climacteric, and
● late ripening (very soft; post-climacteric).
The activity of cell wall degrading enzymes polygalacturonase (PG),
endo- (1 4) -D- mananase (MAN) low in the pre-ripening and early ripening
stages, increased dramatically by mid-ripening coincident with the respiratory
and ethylene climacterics, and decreased in the late ripening. However, PME
activity were highest at green stage when the fruit firmness was high and
decreased as ripening progressed, whereas both EGase and MAN increased
as ripening proceed. The loss of firmness is associated with low but increasing
activities of EGase and MAN. Therefore, PME may first demethylate to
polygalacturonic acid and make it more easily hydrolysed by the three other
enzymes that may play a subsequent role in softening of fruits during ripening.
Thus, these four enzymes may work in concert to soften pawpaw fruit. In
general the different enzymes involved in cell wall degradation are:
pectinesterase, polygalactourasase, gluconase, and cellulose, B-galactanases
etc. which modify the cell wall polysachrides like starch, pectin, cellulose
and hemicelluloses. They lead to dissolution of pectin rich middle lamella.
5. Changes in organic acids : They impart flavour to the fruits, sweetness along
with acidity is useful for imparting flavor. The organic acids vary in different
fruits e.g mango have citric, malic and ascorbic acid, citrus have citric acid,
grapes have tartaric acid and apple have malic acid. But malate and citrate are
more common among fruits. In many fruits such as citrus, loquat, strawberry
and grape, organic acids usually accumulate at the early stages of fruit
development and are used as respiratory substrate during fruit ripening. There
is decrease in organic acid content during ripening because they are used for
respiration and also act as substitute for synthesis of new compounds. With the
 Fruit Ripening – Biochemistry, Physiology and Regulation 289

onset of ripening, the acid content decreases because acids are used in the
process of respiration, converted into sugars, reduced ability of the fruits to
synthesize acids, reduction in translocation from the leaves and increase in
volume of fruits leading to dilution of acids (Dhillon and Gill, 2011).
6. Change in amino acids and proteins : Proteins generally increase with ripening
in many fruits. Methionine and B-alanine are synthesizes from amino acids.
Methionine is the precursor of ethylene biosynthesis. With the onset of ripening,
the ethylene is utilized. So, it signifies the importance of amino acids in ripening
of fruits. They are reduced towards maturity because of incorporation into
proteins required for synthesis of various enzymes. It is presumed that lowering
of amino acids indicates the advancement of maturity. Aspartic acid, glutamic
acids, serine fractions decreases with ripening in muskmelon. Similarly, in
tomato, leucine and iso-leucine decreases with fruit ripening.
7. Enzymes involved in ripening and senescence of fruits : Many of the
chemical and physical affects during ripening and after ripening processes
are attributed to the enzyme actions.

S.No. Action Enzyme involved

1. Softening of fruits Pectin esterase, Polygalacturonase increased

2. Oxidation, ethylene Catalse and peroxidase (involved in autocatalytic synthesis
biosynthesis. of ethylene) increased.
Amine oxidase, alpha-hydroxydase involved in the ethylene
biosynthesis.
3. Glycolytic action. Glucose- phosphate isomerase, phosphofructokinase,
fructose-1.
4. Hydrolytic action. Amylase, cellulase, beta-amylase, starch phosphorilase,
alpha-1, 6-glucosidase.
5. Pigmentation. Chlorophyllase.

8. Phenolic compounds : They impart astringency to fruits e.g tannins. Ripening

generally decreases phenolic compounds and thereby also decreases
astringency. Besides, astringency phenolic compounds are also involved in
in oxidative browning of fruits which is due to polyphenoloxidase enzyme.
Examples:

Crop Phenol Status during ripening

Grape Flavan-3-ol monomers Decreases

Loquat Neocholrogenic acid, hydrobenzoic acid, ferulic acid Decreases
Citrus Naringins, limonoids Decreases
290 Fruit Tree Physiology

Regulation of Ripening
1. Ethylene regulation : Ethylene induces early symptom of senescence in
climacteric and non-climacteric fruits. The increase in ethylene production
whatever it results from, lead to increased levels in the tissue. If this reaches
a critical level, changes in metabolites are initiated, which is associated with
increased rate of respiration. Such changes occur both in climacteric and
non-climacteric fruits provided a sufficiently high concentration of ethylene
is reached in the fruit like plum, which is catalyzed as in banana (Broughton
and Wu, 1979) and apple (Forsyth et al; 1967). Ethylene production is much
higher in climacteric fruits compared to non-climacteric fruits. In climacteric
fruits, the internal ethylene concentration also varies largely while in non-
climacteric ones the variation is only slight.
The use of ethylene gas in achieving faster and more uniform ripening
of fruits is well documented. Ripening is promoted in many fruits by dipping
in 500 to 2000 ppm ethrel in aqueous solution for 30 minutes. The ethrel has
the disadvantage of having to be applied to fruit in aqueous solution, an extra
step in handling which enhances the spread of diseases. Commodities can be
treated with ethylene gas liberated from ethrel in an alkaline medium. Ethylene
released from ethrel was more effective in triggering fruit ripening than
dipping fruits in aqueous solution of ethrel. Enhanced climacteric peak,
increased skin colour, TSS and decreased fruit firmness indicate the effect of
ethylene on fruit ripening (Haithem et al; 2003). Ehtylene generation can be
reduced through:
1. Use of ethylene absorbents e.g KMnO4, AgNO3 etc.
2. Use of antiethylene compounds e.g AVG, AOA.
3. Use of free radical quenching agent's e.g sodium benzoate, n-pyrogallate,
silver ions etc.
4. Ventilation to remove ethylene.
5. UV lamps activated charcoal, catalytic oxidizers etc.
Temperature is the most important factor that regulates ethylene production.
At low temperature shelf life generally enhances due to inhibition of
autocatalytic ethylene production by suppressing the ACC- synthase gene
expression and post transcriptional modification of ACC oxidase. But it can
also enhance ripening due to ethylene synthesis e.g winter pears. High
temperature inhibits ripening by inhibiting conversion of ACC to ethylene.
2. Regulation of O2 and CO2 :
Modified atmosphere packing : The use of storage conditions containing
high concentrations of carbon dioxide and low concentrations of oxygen
 Fruit Ripening – Biochemistry, Physiology and Regulation 291

coupled with low temperatures has proved successful in delaying ripening

and senescence in many fruits. Low levels of oxygen decrease overall
respiration rate, and also appear to block the ethylene forming system. High
carbon dioxide concentrations also decrease ripening, possibly by acting as
a competitive inhibitor of ethylene. It is a useful technology in maintaining
quality and safety of fresh fruits. It changes O2 and CO2 concentrations
within packages, which reduce water loss, respiration and improved sanitation
firmness and skin colour (Kader et al; 2000). In cherry fruits post harvest life
is increased by reducing the rate of decay organisms, retarding softening and
retaining stem colour of fruits (Kupferman and Sanderson, 2001).
Carbon dioxide concentration of 10 per cent or above has been found to
delay the onset of ripening of plum and lengthen the subsequent ripening
period. However, box liner made of polyethylene film to provide modified
atmosphere have not proved useful for several plum cultivars.
Controlled atmosphere storage: Storage life was greatly extended with
a combination of controlled atmosphere storage (CAS). CAS of tropical
fruits has been reported to reduce respiration, lower ethylene biosynthesis
and action, retard ripening and helps to maintain the nutritional quality of
fruit during long-term storage (Kader et al; 2000). Although CAS has been
applied to extend the shelf life of various mango varieties, consumers are
concerned about the poor aroma of CA stored fruits. Increased CO2 and
decreased O2 concentration in the storage atmosphere has been reported to
alter aroma and flavour of various fruits (Kader, 1986). Alphanso mangoes
stored in CO2 concentration above 15 per cent have been reported to cause
fermented odour in the fruit. The concentration of CO2 above 15 per cent
caused a high level of ethanol in the pulp of Kensington Pride mangoes.
High CO2 and/ or low O2 concentration induce anaerobic metabolism, resulting
accumulation of ethanol and acetaldehyde. Mature green and tree ripe Tomy
Atkin mangoes stored in 25 per cent CO2 in combination with 5 per cent O2
tended to have lower terpene and hexanal concentration than in mangoes
stored in 10 per cent CO2, or in air.
Monoterpene and sesquiterpene hydrocarbons are the major volatile
components in all mango varieties, representing 70-90 per cent of total
volatiles. Recently, Lalel et al. (2005) reported the effects of CAS on fruit
ripening and quality in Delta R2E2 mango, whereas, CAS in 6 per cent CO2
and 3 per cent O2 appear to be the most promising to extend the shelf life
(38 days), allowing the fruit to ripen normally with a higher production of
sesquiterpenes and aromatics and lower concentration of alcohols aldehydes
and ester (Lalel and Singh, 2006).
292 Fruit Tree Physiology

3. Chemical treatments :
Calcium : Postharvest application of CaCl2 or Ca (NO3) play an important
role in maintaining the firmness and quality of fruits by thickening of middle
lamella due to increase in formation and deposition of Ca pectate. Postharvest
application of CaCl2 (2-4%) for 5-10 minutes dip extended the storage life
of pear upto 2 months, apple upto 7-months and plum upto 4 weeks at 0-2°C.
Calcium application delays ripening, reduces postharvest decay, controls
development of many physiological disorders and increases calcium content,
thus maintaining the post harvest life of fruits. Calcium can reduce Co2,
ethylene evolution and endogenous ABA levels.
Postharvest calcium application improves the physiological and physico-
chemical characteristics of cell wall pectin by reducing the solubulisation of
uronic acids in the pectin network, without influencing the quality attributes
of many fruits (Tsantili et al; 2002). While beneficial effects of calcium
infiltration have been shown consistently, some studies emphasize the potential
for surface injury, especially in apple fruits, making it applicable only to
fruits destined for processing (Garcia et al; 1995). Apart from calcium chloride,
which has been used extensively in fresh, processed and fresh cut fruits, calcium
lactate and calcium propionate have been proposed as alternative calcium
sources. Whereas these have significant role in freshly cut and canned fruits
(Manganaris et al; 2005). Few studies have focused on compositional changes
in the cell walls of calcium treated fruits (Saftner et al; 2003) and postharvest
application of calcium salt significantly increased firmness, cell wall calcium
content and shelf life of nectarine fruits (Manganaris et al; 2005).

Physiological role of calcium in postharvest physiology of fruits :

● Determines the quality of fruits that are stored for long period.
● Reduces loss of firmness, slow down ripening process of many fruits
when applied as pre and postharvest spray.
● Retards ripening by altering intra cellular and extra cellular processes
which cause lowering of rates of colour change, softening, CO2 and
ethylene production.
Effects of low Ca:
● Produces physiological disorders (bitter pit & cork spot in apple and
stone cells in pear)
● Fruit senescence is hastened.
● Softening of fruits.
● Regulates respiration.
● Ethylene production increases.
 Fruit Ripening – Biochemistry, Physiology and Regulation 293

Physiological role of calcium :

● Ca enters fruits and leaves through stomata and lenticels.
● Ca moves slowly once it is inside cell.
● Increases apoplastic Ca concentration -
1. Exerts a binding effect in the complex of polysaccharides and proteins
comprising the cell wall.
2. Maintains cell wall strength and separation characteristics and firmness
of fruit tissues.
● Cytoplasmic Ca regulates several enzymes
1. Cell wall degrading enzymes polygalacturonase (PG) and
pectinmethylesterase (PME), synthesized de novo during ripening to
softening the ripe fruits.
2. Low peroxidase activity reduces lignification that decreases stone
cells formation in the flesh of pear.
1-MCP: 1-methylcyclopropene (1-MCP) can inhibit fruit ripening by binding
irreversibly to ethylene binding site. As a result, ethylene is unable to bind
and elicit subsequent signal transduction and translation (Serek et al; 1994).
The inhibiting effects of 1-MCP on fruit ripening have been observed in
various fruits and vary widely with fruit species. Concentrations of 0.1-0.5
μl-MCP can effectively block ethylene action in banana fruit (Jiang et al;
2004). The efficacy of 1-MCP treatment on various fruits may differ greatly.
Some fruits such as banana, apple and avocado respond well to low
concentrations (0.1-1.0 μl). However mango and tomato treated with relatively
high concentrations of 1-MCP (20-100 μ1) in order to prevent fruit from
ripening. It is found that postharvest ripening of mango fruit can be inhibited
effectively by treating fruit with 1-MCP using vacuum infiltration which
enhances efficiency of 1-MCP treatment more than 20 folds (Wang et al;
2006).
4. Use of ionizing radiation : Ionizing radiation causes delay in ripening,
destruction of microbes, minimal insect infestation and changes in several
biochemical processes. Generally low radiation dosage is applied to fresh cut
fruits. Gamma rays in pre climacteric fruits reduce not only ethylene
production but also the sensitivity. Gamma irradiation at 100 and 150 krad
increased the shelf life of Solo papaya under ambient conditions by an
additional three days when compared to non irradiated fruits. Singh and Pal
(2009) were able to reduce the decay incidence and increase the postharvest
life by 3-4 days in guava.
294 Fruit Tree Physiology

5. Bioregulators/ chemical regulators : Growth hormones like auxins,

gibberellins and cytokinins delay ripening while ethylene and ABA enhance
ripening. Exogenous application of auxins like 2,4-D; 2,4,5-T stimulate
ethylene production but endogenous auxins inhibit ethylene evolution. GA
and cytokinin delay senescence by preventing the loss of chlorophyll and
enhancing synthesis of carotenoids.
ABA and ethylene promote ripening by affecting the upregulation of
genes encoding ACC synthase. Both abscisic acid (ABA) and C2H4 are
important promoters of ripening where as auxins, cytokinins and gibberellic
acid (GA) invariably have an opposite response to ethylene. Endogenous
auxins act as ripening inhibitors and modify the tissue response to ethylene.
The role of GA as a senescence delaying agent has been demonstrated in
fruits. Cytokinins also interfer preferentially with certain senescence
phenomena mainly delaying colour changes and is associated with the
maintenance of the endogenous GA level and prevention in increase in ABA-
like compound.

CONCLUSION
Fruit ripening is a highly coordinated, genetically programmed, and an irreversible
phenomenon involving a series of physiological, biochemical, and organoleptic
changes, that finally lead to the development of a soft, edible and ripe fruit with
desirable quality attributes. Starch, pectins, cellulose, and hemicelluloses are the
major classes of cell wall polysaccharides that undergo modifications during
ripening. Depolymerization and solubulization of cell wall material by various
enzymes lead to softening of fruits. Increase in respiration occurs in climacteric
fruits during ripening, while no such rise in respiration is observed in non-
climacteric ones. Various biochemical changes associated with fruit ripening involve
chrophyll degradation, synthesis of anthocyanins, carotenoids etc., decreased
acidity, polyphenols, development of volatiles etc. Ethylene a growth hormone
has been found to regulate fruit ripening. Auxin and GA help in delaying the
ripening while ABA and ethylene accelerates the process.

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Lalel HJD and Singh Z (2006) Controlled atmosphere storage of Delta R2E2 mango
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 12

ABSCISSION IN FRUIT PLANTS

A bscission (from the Latin ab meaning away and scindere meaning to cut) is
the shedding of a body part. It takes place primarily at predestined sites that
are commonly located at the base of an organ such as a leaf, flower, fruit or seed.
The process is highly choreographed both in time and space. Trees loose their
leaves by design. When leaves become inefficient and unable to produce food
and growth regulators, a process of shutting down and sealing off begins. Trees
shed many parts besides leaves, including fruit, flowers, bud scales, trichomes,
abscission twigs, and bark. Leaf abscision is the best example of the process of
tissue shedding in trees.

CHANGES DURING ABSCISSION

During abscession changes occur at morphological, anatomical and biochemical
levels.

Morphological and Anatomical Changes

1. Formation of abscission zone at the junction of abscissing organs (leaf, flower,
fruit) and the parent plant body. It is the base of petiole in case of leaves and
the junction of pulvinus and supporting tissue in case of compound leaves.
2. Swelling or constriction of abscission zone w.r.t other parts of petiole.
3. Abscission zone is 1-2 rows or upto 15 or more cell tier thickness.
4. Cells are parenchymatous, devoid of lignin and suberin, dense protoplasm,
more starch, small intercellular spaces and highly branched plasmodesmata.
5. Stele usually divides into separate bundles before it enters the zone.
6. Dissolution of middle lamella resulting in loosening of cells.
 Abscission in Fruit Plants 299

7. Weakening and breaking of vascular connections between leaves, fruits,

flowers and the parent plant.
8. Development of periderm to protect the exposed surface and cells become
suberised.
9. Formation of tylosses (bladder like protrusions from xylem parenchyma cells)
which block xylem vessels.

Biochemical changes
1. Enhanced production of hydrolytic enzymes- cellulase, pectic enzymes, lignase
which cause dissolution of middle lamella and primary walls of the cells in
abscission zone.
2. Polygalactouranase leading to breakdown of middle lamella.
3. Ca pectate in middle lamella is hydrolysed to pectic acid and pectate.
4. Cell expansion by cellulase in abscission zone.
5. Hydrolysis of lignin by lignases.
6. Auxin destruction by increased peroxidase activity.
7. Active synthesis of proteins and RNA in abscission zone.
Besides the above changes, many other processes take place to prepare the
plant for this event. For instance, the vascular elements that service the organ are
occluded and the exposed fracture surfaces are protected from water loss and
invasion by pathogens.

Abscission zones
Abscission zones occur at the base of leaf petioles and at the base of leaflets.
Abscission zones are designed to allow leaf shedding. Leaves are shed through
a number of biological actions which weaken cell walls and initiate cells tearing
away from one another. Abscission zones in trees can be between 5-40 cells wide.
Within this abscission zone, only 1-3 cells will disconnect from each other. Cells
in the abscission zone are of the same types as found elsewhere in the tree.
Abscission zone cells tend to be smaller, more densely packed, with no inter
cellular spaces, less lignin, and have remained in a cell division phase longer than
surrounding cells. Additional cell divisions in this zone prepare these cells for
later abscission processes. Starch is stored in the abscission zone cells to assist
in generating turgor pressure and enzymes for wall degradation. In most abscission
zones, there is a single fault line which develops and is accentuated by additional
wall degradations. Cells adjacent to fault line cells will have weakened walls also,
300 Fruit Tree Physiology

allowing any fractures to propagate along several paths for short distances. Rarely,
several full fault lines occur leaving the abscission wound ragged looking. Fault
lines follow the path of the middle lamella between cells.
Abscission zones are composed of three critical portions (a) a cell wall
degradation area, (b) a shear force generation area and (c) a tree protection zone.
All three abscission zone portions are required for successful leaf shedding and
effective tree survival. Most abscission zones are pre positioned to facilitate
shedding. Abscission zones may not be needed or used, but they are set up to act
as an established barrier and boundary, if needed.
Wall weakening : The abscission process begins with growth regulator signals
initiating cellular changes. Abscission zone cells secrete pectinase and cellulase
(wall degradation enzymes). These enzymes degrade the strength of the middle
lamella and primary wall between cells. The middle lamella, the glue which holds
cells together, begins to dissolve in the abscission zone. At the same time,
surrounding primary walls begin to swell from changes in chemical components.
Calcium bridges across cell wall materials are removed. All cell wall changes are
caused by enzymes and other materials deposited in the cell walls produced by
surrounding living cells. The cells in the abscission zone are dense with cytoplasm
and organelles. Each cell is actively respiring and using energy to produce
abscission materials. These cells remain alive and active until abscission.
Wall changes : As cell wall inter connections are weakened, water pressure
within thin walled cells (turgor pressure in parenchyma) causes these cells to
expand. As cells expand, they generate shear forces by pushing and pulling on
surrounding weakened walls. Mechanically, fracture lines begin to develop between
cell walls. In addition to internal forces, gravity and wind tugging on leaves help
fracture lines grow. As cell walls pull apart from one another, this open wound
is being closed by deposition of blocking materials and protective compounds. A
strong protective boundary zone is prepared to defend remaining tree tissues from
the environment and pests. Tyloses, suberin, lignin and other protective boundary
setting materials are developed and deposited on the tree side of the abscission
zone.

CONTROL MECHANISM OF ABSCISSION

Auxin is a primary growth regulator produced in the leaf and slowly transported
towards the stem base through living cells. As long as auxin is effectively being
transported across the abscission zone, abscission zone cells remain unreactive.
As auxin production begins to decrease in fall and auxin transport rates begin to
decline due to less auxin availability, damage to living cells transporting auxin,
 Abscission in Fruit Plants 301

Fig. : Two dimensional diagram showing cells in a leaf base abscission zone with
a fracture line between cells. Tree protection zone, wall degradation areas,
and cell expansion zone all disrupting cell-to-cell connections are clearly
shown (www.answers.com/topic/abscission).

(www.answers.com/topic/abscission).

and/ or accelerating infection of living tissues by pests, cell wall changes are
initiated. Cell wall changes increasingly inhibit auxin transport and accelerate
ethylene production. Small amounts of ethylene hasten abscission zone
development. ABA (abscisic acid) is responsible (in part) for dormancy on set in
the leaf stimulates ethylene production and inhibits auxin transport.
Deciduous trees do not loose all their leaves at once or just in the fall. The
larger and stronger any connecting xylem elements through the abscission zone,
the longer leaves may be held on the tree. Some species do not fully set an
abscission layer until early winter. In other species, shear forces are not concentrated
in the abscission zone until the beginning of the spring growth period. Juvenile
trees may not establish effective abscission zones at all and hold dead leaves
throughout the winter. Understory trees may hold leaves because of juvenility or
because they are protected from climatic events which could knock off the leaves.
Some trees may abscise all their leaves except on new late season sprouts.
302 Fruit Tree Physiology

Various hormones such as auxin, gibberellins, cytokinins, ABA and ethylene

play significant role in abscission. Auxins, gibberellins and cytokinins retard
abscission while ethylene and ABA accelerate it. During abscission endogenous
auxin and cytokinin content decreases sharply while ABA and ethylene increase.
Exogenous application of auxin or cytokinin delays abscission. But if auxin
(IAA) is applied proximal to abscission zone, it accelerates abscission while
application to distal end (leaf petiole) retards abscission. However, application of
ABA either at proximal or at distal end accelerates abscission. Ethylene application
also promotes abscission. It is postulated that ethylene accelerates abscission by
increasing respiration as well as permeability of the cell membrane.
Mechanism of differentiation of the abscission zone (AZ) : The site where
abscission takes place comprises a tier, or more commonly a number of layers,
of cells that are anatomically distinct long before organ separation has been
initiated i.e abscission zone (AZ) represents a positionally differentiated group of
target cells. Further evidence to support this concept comes from the demonstration
that when the process is triggered specific molecular, biochemical and ultrastructural
changes are restricted to the AZ cells that cannot be detected in adjacent non
separating cells. Such changes include:
● Up-regulation of abscission related genes,
● Increases in the activity of hydrolytic enzymes,
● Activation of the endomembrane complex, and
● The dissociation of the cell wall.
A number of mutants have been characterized whose organ fail to undergo
abscission in order to unravel the events that lead to AZ differentiation (Leslie
et al; 2006). One of the first to be studied was the jointless (j) mutant of tomato
that fails to shed its flowers at the pedicel abscission zone. Such plants were
selected as having an agronomic advantage as harvested fruits were stemless. A
structural examination indicates that an anatomically distinct AZ fails to
differentiate in the pedicel of mutant flowers and fruit. The jointless gene has
been cloned and found to encode a MADS box domain transcription factor (Mao
et al; 2000). It is not clear how JOINTLESS brings about its effects although
experiments on chimaric plants between grafted J/j plants (Szymkowiak and
Irish, 1999) have revealed that expression of JOINTLESS within the L3 layer is
sufficient for AZ differentiation to take place at the knuckle region of the pedicel.
This intriguing discovery suggests that a signal from the vascular tissues, which
develop from L3, is sufficient to bring about the formation of AZ cells within the
cortical and epidermal layers. Although, Arabidopsis plants do not lose their
leaves, this model species has been used effectively as a subject of forward
 Abscission in Fruit Plants 303

genetic screens in the search for phenotypes that fail to abscise floral organs or
seeds (Patterson, 2001). The first non abscising Arabidopsis mutant reported was
inflorescence deficient in abscission (ida). The ida gene was isolated and found
to encode a small polypeptide. Reporter gene analysis has revealed that ida is
expressed specifically in the AZ cells of floral organs and may act as a novel
ligand (Aalen et al; 2006., Matsubayashi and Sakagami, 2006). While sepals,
petals and stamens of the mutant are retained throughout pod development, break
strength analyses have revealed that the force required to detach petals from the
flower declines initially at a similar rate in both ida and wild type plants (Butenko
et al; 2003). Just prior to organ shedding, however, petal break strength in the
mutant increases and the reproductive organs seem to become firmly attached to
the flower once more. The most plausible explanation to account for these
observations is that ida plays a critical role in one of the final stages of abscission.
If this element of the process fails, then organ dissociation does not take place
and that synthesis of a material to protect the fracture surface cements the sepals,
petals and anther filaments back in to place. While this hypothesis may be correct,
it has recently been shown that over expression of ida leads to the formation of
ectopic abscission at the pedicel:stem junction and the base of the silique in
addition to a proliferation of separating cells at the sites of floral organ shedding
(Stenvik et al; 2006). These observations suggest that ida also plays a role in
delimiting the AZ itself and in determining the number of tiers of cells that
undergo separation. The receptor for ida has yet to be identified, however, down
regulation of a leucine rich repeat class of receptor like kinase (RLK) termed
haesa, using RNAi strategy, has been shown to phenocopy the ida mutant in
Arabidopsis (Jinn et al; 2000). Many additional rlk family members exist within
the Arabidopsis genome and it remains to be seen which, if any, of these might
bind the ida protein.
Other genes that have been proposed to contribute to AZ formation include
the functional homologues blade on petiole (bop) 1and 2. In the absence of both
proteins, organ patterning is disrupted and floral organ shedding does not take
place.

REGULATION OF ABSCISSION
Regulation of timing : The time from the differentiation of the abscission zone
to organ shedding can range from days (or even hours) to many months. Thus,
leaves from deciduous species that have just emerged and expanded in the spring
can be induced to abscise by treatment with the gaseous plant hormone ethylene
or lamina removal even though they will not naturally be shed until the autumn.
This observation tells us that the onset of abscission is dictated by events subsequent
304 Fruit Tree Physiology

(Patterson, 2001)
 Abscission in Fruit Plants 305

to AZ differentiation and that both stimulatory and inhibitory regulators may

influence the timing of organ shedding. It is well documented that the main
abscission accelerating agent is ethylene while the plant hormone indoleacetic
acid (IAA) delays the process. It has been proposed that by altering the balance
between these two naturally occurring stimuli the timing of abscission can be
accurately controlled in plants. The origin of the elevated ethylene that precedes
organ shedding emanates from tissues distal to the AZ such as the diseased or
senescing leaf blade, or ripening fruit in climacteric species. Although application
of ethylene has been shown to promote abscission, floral organs of Arabidopsis
mutants such as etr 1, that are blind or insensitive to the gas, are shed albeit at
a later stage of development than wild type indicating that it is not an absolute
requirement for shedding to be initiated (Patterson, 2001). ABA was originally
thought to be a key regulator, however, in recent years its role has been changed
to one of accelerating the senescence, and associated elevated ethylene production,
of distal tissues. The recent studies suggest that methyle jasmonate might also
have a function in regulating the timing or organ loss based on discovery that the
delayed abscission 4 (dab 4) mutant of Arabidopsis is an additional allele of the
coronatine insensitive mutant 1 (coi 1).
Regulation of cell separation : Abscission leads to a series of events that
result in cell separation at the point of organ detachment. Ultra structural analysis
indicates that this is primarily a consequence of the dissolution of the middle
lamella and that immediately prior to shedding it is only the lignified tissues of
the vascular trace that retain an attachment to the parent plant . These connections
are thought to be finally severed by the hydraulic expansion of the isodiametric
AZ cells once the constraints imposed by the matrix have been loosened.
A number of mechanisms have been suggested that bring about wall breakdown
but most important is by an enzymatic process (Sexton and Roberts, 1982). The
principle constituents involved are endo ∝ 1, 4 glucanases (EGases) and
polygalacturonases (PGs) whose activity has been shown to increase almost
ubiquitously in AZ tissues during organ shedding (Roberts et al; 2000) . These
two enzymes cause solubilization of the cellulosic and pectinaceous moities of
the middle lamella respectively. RNAi technology has been used to prevent a
decline in AZ break strength by silencing their expression. This strategy has been
adopted using RNAi to down regulate EGases in tomato and T-DNA insertion
lines to silence PGs in Arbidopsis. While both approaches have brought about a
delay in abscission but the shedding process has not been prevented. It is possible
that these enzymes act in concert or that other gene family member can contribute
to the process, alternatively other enzymes or proteins might play a key role in
the process.
306 Fruit Tree Physiology

Ultrastructural studies of AZ cells have revealed that dilation of the

endomembrane system takes place prior to cell separation and that vesicles appear
to be discharged from the golgi into the plasma membrane. These vesicles deliver
materials that contribute to wall breakdown and to wall reassembly at the fracture
surface. Although the secretory events have yet to be established, the Arabidopsis
mutant nevershed fails to undergo floral organ abscission and has been proposed
to encode a protein that may play a role in the secretory process. Additionally
down regulation of arp7, a member of a gene family encoding actin related
proteins, using RNAi, has been shown to delay the timing of abscission in
Arabidopsis suggesting certain cytoskeletal components in the secretory process
are involved (Kandasamy et al; 2001). Although a number of putative cell wall
degrading agents have been proposed to contribute to the breakdown of matrix
between AZ cells, none of these has yet been definitively shown to play a crucial
role in bringing about abscission.

PROTECTION OF THE FRACTURED SURFACE

Once cell separation has taken place and the subtending organ has been shed, the
scar remaining provides an ideal site for invasion by bacteria and fungi. But the
remaining cells divide and suberize to protect the wound. The broken xylem
vessels become blocked with gums or tylosses and the phloem sieve plates are
callosed over to prevent pathogen entry. The antimicrobial enzymes chitinase and
∝ 1-3-gluconase are produced in the fracture surface to prevent infection. Molecular
studies reveal that shedding is accompanied by an increase in expression of a
number of pathogenesis related (PR) genes (Coupe et al; 1997). These genes
encode such polypeptides as chitinases, protease inhibitors and wound induced
(win) proteins and their expression is up regulated in the absence of a pathogen
attack, indicating that the AZ cells are pre prograrnmed to generate antimicrobial
proteins once the shedding process has been triggered. The mechanism responsible
for co-ordinating gene expression is unclear, although, it is possible that reactive
oxygen intermediates (ROI) might play a role. Such a mechanism would lead to
the build up of free radicals within the AZ tissues unless metallothionein (MT)
like proteins act as scavengers to prevent their accumulation (Coupe et al; 1995).
Anatomical studies have shown that abscission is preceded by the formation
of tyloses within the vascular tissue proximal to the site of separation. These
structures develop from the xylem parenchyma cells and protrude through the
sides of the vessels to block and finally occlude them. The timing of this process
must be closely co-ordinated so that distal tissues are not deprived of water
otherwise the AZ cells would desicate and cell separation would fail to proceed.
The mechanisms responsible for this process are unknown but may be co-ordinated
 Abscission in Fruit Plants 307

with the synthesis of cuticular waxes to provide a protective layer once shedding
has taken place (Aharoni et al; 2004). The isolation and characterization of the
ida mutant suggests that the synthesis of new cell wall material takes place at the
time of organ loss. The precise nature of this phenomenon is unknown, however,
the recent demonstration that ida promotes the expression of a specific
arabinogalactan protein (AGP24) may provide a possible clue to the events that
take place. AGPs have been shown to promote the activity of xyloglucan
endotransglucosylase/ hydrolases (XTHs) a group of enzymes that are known to
remodel the structural cellulose xyloglucan matrix of the cell wall.

Functions of Abscission
● A plant abscises a part either to discard a member that is no longer necessary,
such as a leaf during autumn, or a flower following fertilization, or for the
purposes of reproduction. Most deciduous plants drop their leaves by
abscission before winter, while evergreen plants continuously abscise their
leaves.
● Fruit drop is another form of abscission. When a plant abscises fruit while
still immature, in order to conserve resources needed to bring the remaining
fruit to maturity.
● A plant also abscises damaged leaves to conserve water or photosynthetic
efficiency.

Significance of Abscission
● Separates senessent and dead plant parts.
● Fruit abscission helps in their dispersal and to repeat their life cycles.
● Conservation of water during summer by reducing transpiration surface.
● Helps in vegetative reproduction in lower plants by separating their vegetative
parts.

Factors Influencing Abscission

1. Light : Low light intensity promotes abscission. Shorter photoperiod enhances
abscission while longer photoperiod retards abscission. Red light represses
abscission than far red.
2. Water : Low soil water enhances abscission.
3. Minerals : Mineral deficiency accelerates abscission.
308 Fruit Tree Physiology

4. Air pollution : Air pollutants like ozone, hydrogen fluoride, nitrogen dioxide
accelerate abscission.
5. Oxygen supply : Low oxygen retards abscission due to reducing the activity
of hydrolytic enzymes.

MANIPULATION OF ABSCISSION
Too little abscission of young fruits result in production of large number of
unmarketable fruits whereas too much abscission results in uneconomical yields.
Hence manipulation of abscission under certain conditions is desirable. There are
different techniques through which abscission can be manipulated. Broadly they
can be categorized as follows:
Understanding physiological basis of abscission : By understanding the
physiology of abscission, flower and fruit drop can be manipulated accordingly.
Genetics of abscission : Breeding of varieties with different abscission
characteristics for example raspberry cultivars. There are two tyes of cultivars
viz.1. hand picking ones that do not drop ripe berries by shaking the bush and
2. mechanical picking types, where fruits easily detach.
rDNA technology : Use of antisence RNA technology for ACC synthase to
block the ethylene synthesis and hence slowing abscission. It is also used for wall
degrading enzymes cellulase and polygalactourasae (PG).
Acceleration of abscission : By increasing ethylene production e.g spraying
ethephon, wounding.
Inhibition of abscission : Reducing ethylene production or interfering with
ethylene action e.g use of antagonists like AVG, AOA, silver thiosulphate, DACP.
Prevention of abscission : Use of auxins and cytokinins.

CONCLUSION
Abscission is a highly co-ordinated process that leads to shedding of various
organs in plants. Various morphological, anatomical and biochemical changes
take place during this process. During abscission there is degradation of middle
lamella of cells and formation of abscission zones, and enzymes such as EGs and
PGs are thought to play important role in these events. There is protection of
fractured surfaces through expression of PR related proteins. Although many
hypothesis have been put forth to explain the process of abscission but the
molecular mechanisms remain still unclear.
 Abscission in Fruit Plants 309

REFERENCES
Aalen RB, Butenko MA, Stenvik GE, Tandstad NM and Patterson SE (2006) Genetic
control of floral abscission. In: da Silva JT (ed.) Floriculture, Ornamental and
Plant Biotechnology: advances and topical issue, Pp. 101-108. London: Global
Science.
Aharoni A, Dixits and Jetter R (2004) The SHINE Clade of AP2 domain transcription
factors activates wax biosynthesis, alters cuticle properties, and confers drought
tolerance when overexpressed in Arabidopsis. Plant Cell 16: 2463-2480.
Coupe SA, Taylor JE and Roberts JA (1995) Characterization of an mRNA encoding a
metallothionein like protein that accumulates during ethylene promoted abscission
of Sambucus nigra L leaflets. Planta 197: 442-447.
Coupe SA, Taylor JE and Roberts TA (1997) Temporal and partia1 expression of mRNAs
encoding pathogenesis related proteins during ethylene promoted leaflet abscission
in Sambucus nigra. Plant cell & Env 20: 1517-1524.
Kandasamy MK, Deal RB, McKinney EC and Meagher RB (2005) Silencing the nuclear
actin related protein at ARP4 in Arabidopsis has multiple effects on plant
development, including early flowering and delayed senescence. Plant J 41:
845-858.
Lesile ME, Lewis MW and Liljigren SJ (2006). Organ abscission. Ann Plant Rev 25:
106-136.
Mao L, Begum D, and Chaung HW (2000). JOINTNTLESS is a MADS-box gene
controlling tomato flower abscission zone development. Nature 406: 910-913.
Matsubayashi Y and Sakagami Y (2006). Peptide hormone in plants. Ann Rev Plant Biol
57: 649-674.
Patterson SE (2001) Cutting loose. Abscission and dehiscence in Arabidopsis. Plant
Physiol 126: 494-500.
Roberts JA, Elliot KA and Gonazalez CZH (2002). Abscission, dehiscence and cell
separation processes. Ann Rev Plant Biol 53:131-158.
Sexton R and Roberts JA (1982). Cell biology of abscission. Ann Rev Plant Physiol 33:
133-162.
Stenvik GE, Butenko MA, Urbanowicz BR, Rose JKC and Aelen B (2006) Overexpression
of inflorescence deficient in abscission activates cell separation in vestigial abscission
zones in Arabidopsis. Plant Cell 18: 1467-1476.
Syzmkowiak EJ and Irish EE (1999). Interactions between jointless and wild type tomato
tissues during development of the pedicel abscission zone and the inflorescence
meristem. Plant Cell 11: 159-175.
 INDEX

Abscission - definitions - 1
- abscission zones - 298 - dormex - 20
- anatomical changes - 298 - endodormancy - 2, 4
- fractured surfaces - 306 - factors affecting - 11
- mechanism - 300 - hormone action - 8, 13
- morphological changes - 298 - hydrogen cyanamide - 20
- regulation of cell separation - 305 - mechanism - 5
- regulation of timing - 303 - nutrient diversion - 16
- wall changes - 300 - paradormancy - 2, 4
- wall weakening - 300 - site of dormancy - 4
Alternate bearing Chilling
- factors affecting - 181 - buttoning - 27
- index (ABI) - 179 - chilling negation - 36
- regular bearing cultivars - 98 - chilling requirement - 27
- regulation of flowering - 195 - commulative chilling hours - 28
- smudging - 196 - dynamic model - 42
Annona - 66 - effective chilling temperature - 34
Apricot - 60 - foliation - 26
Apple - 64, 131, 221 - GDH - 26, 34
Banana - 60 - heat accumulation - 36
Bud dormancy - models - 38
- calcium - 16, 22 - resting stage - 34
- classification - 2 - Utah model - 40
 Index 311

Citrus - 57, 64, 222 - detection of incompatibility - 246

Date palm - 55, 224 - methods to overcome
Dwarfism incompatibility - 247
- alar - 268 - symptoms - 235
- hydraulic conductivity - 257 - types of incompatibility - 236
- onium compounds - 269 Loquat -140, 225
- physiology - 251 Mango - 56, 119, 225
- triazoles - 269 Olive - 61
- ways for achieving dwarfism - 254 Parthenocarpy
Flowering - changes due to parthenocarpy - 206
- evocation - 71 - chromosomal aberration - 214
- factors affecting induction of - development of parthenocarpic
flowering - 88 fruits - 209
- genetics of flowering - 101 - importance - 206
- induction - 71 - induction of parthenocarpy - 220
- initiation - 72 - physiological basis - 207
- mechanism - 74 - self incompatibility - 215
- physiology - 70 - types - 205
Fruit Peach - 59, 225
- factors affecting fruit growth and Pear - 136, 226
development - 116 Pineapple - 61, 65
- key growth stages - 114 Plum - 137
- size and shape - 117 Photoperiodism
- stages of development - 111 - short day plants - 76
Grape - 54, 65, 128, 223 - induction - 78
Heat units - 49 - perception - 78
GDD - 50 - nature of stimulus - 78
Incompatibility - phytochrome - 79
- biochemical reasons of graft - florigen - 83
incompatibility - 243 Raspberry - 59
- causes of incompatibility - 237 Ripening
- graft incompatibility - 234 - aroma volatiles - 286
- localized incompatibility -236 - CAS - 291
312 Fruit Tree Physiology

- cell wall degradation - 287 Unfruitfulness

- climacteric - 279 - heterostyly - 155
- degradation of chlorophyll - 284 - dichogamy - 155
- hydrolysis of starch - 285 - duodichogamy - 157
- MAP - 293 - protogyny - 148, 156, 157
- mechanism - 281 - sterility - 160
- non climacteric - 279 - tristyly - 155
- regulation - 290 Vernalisation
- textural softening - 285 - mechanism - 85
Strawberry - 63, 139 - site of vernalization - 85
Thermoperiodism – 61 - stimulus of vernalization - 85
- day-night temperatures - 62