Plant Physiol.

(1980) 65, 1146-1148

0032-0889/80/65/1 146/03/$00.50/0

Characteristics of Cold Acclimation and Deacclimation in Tuber-

bearing Solanum Species1
Received for publication October 2, 1979 and in revised form February 11, 1980

HWEI-HWANG CHEN AND PAUL H. Li

Laboratory of Plant Hardiness, Department of Horticultural Science and Landscape Architecture, University of
Minnesota, Saint Paul, Minnesota 55108

ABSTRACT capable of cold acclimation? Are any of the frost tolerant species
unable to acclimate? Are there frost-sensitive species other than S.
The effect of temperatures on cold acclimation and deacclimation in tuberosum which are able to acclimate? This information would
foliage tissues was studied in Solan commersonii (Oka 4583), a tuber- be of interest not only to plant physiologists, but also would be
bearing potato. The threshold temperature for cold acclimation was about important in breeding for potato frost resistance.
12 C. In a temperature range of 2 to 12 C, the increase in hardiness was
dependent on the acclimating temperature; the lower the acclimating
temperature, the more hardiness achieved. A day/night temperature of 2 MATERIALS AND METHODS
C, regardless of photoperiod, appeared to the optimum acclimating tem- The tuber-bearing Solanum species were propagated from seed
perature for the SoIuuam species studied. A subfreezing temperature
hardened plants less effectively. The maximum level of hardiness could be
tubers obtained from the Potato Introduction Station, Sturgeon
reached after 15 days of cold acclimation. However, it took only I day to
Bay, Wisconsin. Each tuber was planted in a 6-inch pot containing
deacclimate the hardened plants to a preacclimation level when plants a 3:2:2 by volume mixture of soil, sand and peat. They were grown
were subjected to a warm regime from cold. The degree of deacclimation
in a regime of 14-h light and 20/15 D/N temperature. After 2
was dependent on the temperature of the warm regime.
months, uniform plants were selected and used for the studies of
Based on cold tolerance and the capacity to acclimate to cold, tuber-
cold acclimation and deacclimation studies.
bearing Solawun species could be grouped into five categories. Chilling
Cold Acclimation Treatments. To determine the range of tem-
injury was also observed in some of the tuber-bearing Solanum species. peratures in inducing hardiness, 2 month old Solanum commersonii
'Oka 4583' plants were used. They were grown in chambers in
which a 14-h light and a temperature of 20/15, 14, 12, 10, 5, and
2 D/N were held, respectively. Foliage frost hardiness was deter-
mined after 20 days of growth (2, 18).
When the air temperature was lowered to below -1C, the pot
soil froze and resulted in plant wilting (1). Stem-cultured plants of
Frost is often a major factor in reducing potato production or S. commersonii were prepared (1) and thus used as an alternative
in total crop failure in many growing areas (5, 15). The most way to study the effect of subfreezing temperatures on cold
commonly grown Solanum tuberosum potato possesses no frost acclimation. Two-month-old stem-cultured plants grown in media
tolerance, although a number of noncultivated species are frost were treated in regimes of 14-h light and temperatures of 20/15,
hardy and can survive at -4C or colder (1, 3, 16). Some of the 10, 5, 2, 0, and -2 C D/N. Frost hardiness was evaluated after 20
frost hardy species have the capability to acclimate to cold, but days of treatment (2, 18).
the same acclimating conditions fail to harden S. tuberosum (1, 3). Twenty-four tuber-bearing Solanum species were examined for
It is also known that the lower the acclimating temperature the the relationship between the frost resistance and the capacity for
greater the frost hardiness achieved (1). But it is not known cold acclimation. A regime of 2 C and 14-h light was used for the
whether the potato can be hardened to a greater degree when cold acclimation treatment, and the frost hardiness was measured
subjected to the sub-freezing temperatures as has been found in before and after 15-20 days of treatment (2, 18).
other plant species (11, 12). Also, what is the minimum low Deacclimation Treatments. Two-month-old S. commersonil
temperature requirement for the initiation of the acclimation plants were used for the deacclimation study. They were hardened
process? at 2 C D/N for 20 days and the hardiness of mature leaves was
It has been reported for many species that cold acclimated determined at 5-day intervals during acclimation. After 20 days,
plants rapidly lose their hardiness upon exposure to warm tem- the hardened plants were transferred to chambers with regimes of
perature (7, 8, 10, 13, 14). The rate of deacclimation varies with 14-h light and 20 and 10 C D/N, respectively. Frost hardiness of
the genetic composition of plants, the degree of temperature rise, the foliage was then determined at 3, 6, 15, 24, 48, and 96 h after
and the exposure period at the warm temperature. None of this deacclimation.
kind of information is available for tuber-bearing Solanum species.
Frost-killing temperatures of 60 tuber-bearing Solanum species RESULTS AND DISCUSSION
were reported (16). The characteristics of cold acclimation in a
few species were studied (1, 3). However, there are still questions Several studies of potato cold acclimation with a number of
to be answered. For example, are only the frost tolerant species species were made (1-3). In the present study, S. commersonii was
chosen as a model system for the study of deacclimation as well
' Scientific Journal Series No. 10979 of The Minnesota Agricultural as to determine the range of temperatures in inducing frost har-
Experiment Station. This research was supported in part by a research
2
contract from The International Potato Center, Lima, Peru. Abbreviation: D/N: day/night.
1146
Plant Physiol. Vol. 65, 1980 POTATO FROST HARDINESS 1147
diness. This is because S. commersonii is one of the species, Table I. Classffication of Tuber-Bearing Solanum Species in Terms of
identified so far, that shows the greatest capability to acclimate to Cold Tolerance and Cold Acclimation
cold. The relationship between frost tolerance and cold acclima- Killing Tem-
tion was also examined in 24 tuber-bearing Solanum species. perature
Frost hardiness in S. commersonii was increased at a tempera-
ture of 12 C but not at 14 C (Fig. 1). The upper limit of acclimating Categories Species of Solanum Before After
temperature appeared between 12 and 13 C. The optimum accli- Accli- Accli-
mating temperature was about 2 C. The 2 C was also the optimum ma- ma-
acclimating temperature for other potato species (data not shown, tiona tionh
Table I). A similar temperature range has been found effective in
inducing frost resistance in several plant species (4, 6, 9).
At 0 C, plants were acclimated from -4.5 to -9 C within 20 Group I:
days which was 2.5 less hardy than plants acclimated at 2 C. Frost resistant and acaule (Oka 3885)C -6.0 -9.0
When plants were acclimated at -2 C, leaves showed senescence able to cold commersonii (Oka 5040) -4.5 -11.5
after one week and were dead in about 2 weeks. The frost-killing harden multidissectum (PI 210042) -4.0 -8.5
temperature determined after 10 days at -2 C was -6 C. chomatophilum (PI 266387) -5.0 -8.5
In general, plants of S. commersonii (Fig. 2) and others (Table Group II:
I, groups I and III, data not shown) reached their maximum Frost resistant but bolviense (PI 265860) -4.5 -4.5
hardiness after 15 days of treatment. Thereafter, no further in- unable to cold megistacrolobum (Oka 3914) -5.0 -5.0
crease in frost hardiness was observed. harden sanctae-rosae (Oka 5697) -5.5 -5.5
Low temperature requirements for the development of maxi- Group III:
mum hardiness varies from species to species (1 1). Plants such as Frost sensitive but oplocense (Oka 4500) -3.0 -8.0
winter rape (11) and cabbage (12, 14) have been shown to require able to cold polytrichon (PI 184773) -3.0 -6.0
freezing temperature to attain a maximum hardiness level, whereas harden
others (6) do not, including tuber-bearing Solanum potatoes (Fig. Group IV:
1). We observed that the subfreezing temperature caused damage Frost sensitive and brachistotrichum (PI 320265) -3.5 -3.5
to the potato plants. It has been noted that the threshold temper- unable to cold cardiophyllum (PI 186548) -3.0 -3.0
ature for cold acclimation is difficult to determine and varies with harden fendleri (PI 275163) -3.0 -3.0
plant species (14). A temperature of 12 C for inducing cold jamesii (PI 275163) -3.0 -3.0
hardines was reported for cabbage and Mentha viridis (14). Sakai kurtzianum (Oka 4940) -3.5 -3.5
(17) concluded that any temperature above 13 C led to a loss in microdontum (Oka 5623) -3.0 -3.0
freezing tolerance in some tree tissues. pinnatisectum (PI 275232) -2.5 -2.5
stenotomum (PI 195188) -3.5 -3.5
- p I I I I I stolon!ferum (PI 161770) -3.0 -3.0
sucrense (EBS 1791) -3.0 -3.0
-12 tuberosum (cv. Red Pontiac) -3.0 -3.0
venturii (Oka 498) -3.5 -3.5
i
-11 IAl vernei (Oka 4476) -3.5 -3.5
verrucosum (PI 195170) -3.0 -3.0
-10- /1 '
Group V:
Chilling sensitive trifldum (PI 255541) -3.5 deadd
o0 -99 a Plants were grown in a regime of 20/15 C D/N, 14 h light.
b
W. Plants were grown in a regime of 2 C, 14 h for 20 days.
= -8
A-# c Identification number at the Potato Introduction Station, Sturgeon
CO -7
Bay, Wisconsin.
CL d Plants were dead after 20 days of 2 C treatment.
E-
C1 -6
Results of deacclimation studies are shown in Figure 2. When
I-
= -5 the acclimated S. commersonii plants were subjected to a regime
02 -6 of 20 C, frost hardiness was decreased to that of the plant as the
= -5 preacclimation levels in only 1 day (Fig. 2A). Upon close exami-
nation, it was revealed that deacclimation was initiated as early as
2-3 h after the plant was subjected to warm temperature (Fig. 2B).
6-
LL IF When plants were deacclimated at 10 C, the hardiness decreased
-2 from -1 1.5 to -9 C in one day (Fig. 2 A and B) and maintained
at this level during prolonged treatment. The -9 C level was the
-1
maximum hardiness S. commersonii could attain when plants were
acclimated at 10 C for 15 days (1).
u I l To attain the maximum level of frost hardiness, Solanum po-
20/15 14 12 10 8 6 4 2 0 -2 tatoes required about 15 days of cold acclimation but the deaccli-
mation was rapid (I day). These observations were similar to that
Acclimating Temperature,C observed in cereals (7, 13) and woody perennials (8, 10). The level
FIG. 1. The range of temperature effect in inducing frost hardiness in of hardiness reduced in the potato was dependent on the deaccli-
S. commersonii. Pot-grown plants (El) and agar medium-grown, stem- mating temperature. For instance, fully acclimated S. commersonii
cultured plants (0). Frost killing temperatures were determined after 20 plants deacclimated less at 10 C than at 20 C. The deacclimated
days of cold acclimation. a: Frost killing was determined after 10 days of plants at 10 C still retained a hardiness level which was the
treatment. maximum hardiness to which they could be acclimated at 10 C.
1148 CHEN AND LI Plant Physiol. Vol. 65, 1980
sum, belongs to group IV. From the results summarized in Table
0 I, the characteristic of frost resistance does not appear to be an
-1
absolute requirement for cold acclimation. The mechanisms of
frost resistance and cold acclimation seemed to be independent of
each other.
The potato is a cool season crop and it is usually considered to
-3 be chilling resistant, yet chilling injury was observed in S. tr!fidum
P -4
plants grown at 2 C. Dark necrotic spots appeared on the leaf
surface at first, and later the leaf completely dried out. The whole
=
-
-55 plants died in I or 2 weeks. These symptoms were similar to those
of chilling injury reported in other plants (14).
2 -66
E
LITERATURE CITED
7
_
1. CHEN HH, P GAVINLERTVANTA, PH Li 1979 Cold acclimation of stem-cultured
-I 8 plants and leaf callus of Solanum species. Bot Gaz 140: 142-147
2. CHEN P, MJ BURKE, PH Li 1976 The frost hardiness of several Solanum species
a -99 in relation to the freezing of water, melting point depression and tissue water
0

7L content. Bot Gaz 137: 313-317

-10- 3. CHEN P, PH Li 1976 Effect of photoperiod, temperature, and certain growth
regulators on frost hardiness of Solanum species. Bot Gaz 137: 105-109
4. DEVAY M, E PALDI 1977 Cold-induced rRNA synthesis in wheat cultivars during
the hardening period. Plant Sci Lett 8: 191-195
-12 5. ESTRADA RN 1978 Breeding frost-resistant potatoes for the tropical highlands. In
(PH Li, A Sakai, eds,) Plant Cold Hardiness and Freezing Stress. Academic
Press, New York, pp 333-341
o 24 6 S ~10 12 14 16 16 20 22 24 26 6. GRENIER G, A TREMOLIERs, HP THERRIEN, C WILLEMOT 1972 Changements
TIME (Days) dans les lipides de la luzerne en conditions menant a 1'endurcissement au froid.
Can J Bot 50: 1681-1689
FIG. 2. Cold acclimation and deacclimation of S. commersonii (A). 7. GUSTA LV, DB FOWLER 1976 Effect of temperature on dehardening and rehar-
Plants were treated at 2 C for 20 days and then transferred to two regimes dening of winter cereals. Can J Plant Sci 56: 673-678
8. GUSTA LV, CJ WEISER 1972 Nucleic acid and protein changes in relation to cold
of 10 and 20 C, constant D/N temperature. Arrows indicate the initiation acclimation and freezing injury of Korean boxwood leaves. Plant Physiol 49:
of cold acclimation ('?Q) and deacclimation (9). The insertion (B) dem- 91-96
onstrates the rate of deacclimation at hourly intervals. 9. HANTANO S, H SADAKANE, M TUTUMI, T WANTANABE 1976 Studies on frost
hardiness in Chlorella ellipsoidea. II. Effects of inhibitors of RNA and protein
and surfactants on the process of hardening. Plant Cell Physiol 17: 643-651
However, in winter wheat, when acclimated plants were deaccli- 10. HOWELL GS, CJ WEISER 1970 Fluctuation in the cold resistance of apple twigs
mated for 6 days at either 20 or 10 C, hardiness level was reduced during spring dehardening. J Am Soc Hort Sci 95: 190-192
I 1. KACPERSKA-PALACZ A 1978 Mechanism ofcold acclimation in herbaceous plants.
to the same amount in each (7). Deacclimation in S. commersonii In (PH Li, A Sakai, eds,) Plant Cold Hardiness and Freezing Stress. Academic
was initiated at about 2-3 h after exposure to warm temperature Press, New York, pp 139-52
and plants were completely deacclimated within 24 h. It has also 12. KOHN H, J LEVITr 1965 Frost hardiness studies on cabbage grown under
been observed that deacclimation can be initiated in a few hours controlled conditions. Plant Physiol 40: 476-480
13. LAUDE HH 1937 Cold resistance of winter wheat, rye, barley and oats in transition
in some species (13). Other plants are reported to require several from dormancy to active growth. J Agr Res 54: 899-917
days for complete deacclimation (7, 8, 10). 14. LEVVITT H 1972 Responses of Plants to Environmental Stress. Academic press,
Based on the gathered information, the tuber-bearing Solanum New York, p 697
species could be classified into five groups (Table I.). They are: I: 15. Li PH, JP PALTA 1978 Frost hardening and freezing stress in tuber-bearing
Solanum species, In (PH Li, A Sakai, eds,) Plant Cold Hardiness and Freezing
species which are frost resistant (survive.at -4 C or colder) and Stress. Academic Press, New York, pp 49-71
able to cold harden; II: species which are frost resistant but are 16. Li PH 1977 Frost killing temperatures of 60 tuber-bearing Solanum species. Am
unable to cold harden; III: species which are frost sensitive (survive Potato J 54: 452-456
at -3 C or warmer) but able to cold harden; IV: species which are 17. SAKAi A 1967 Mechanism of frost damage on basal stems in young trees. Low
Temp Sci Ser B25, pp 45-47
frost sensitive and unable to cold harden; and V: species which 18. SUKUMARAN NP, CJ WEISER 1972 An excised leaflet test for evaluating potato
showed chilling injury. The commonly grown potato, S. tubero- frost tolerance. Hortscience 7: 467-468