Breeding Methods in Tree Improvement

Franklin C. Cech
Southlands Experiment Forest, International Paper Company, Bainbridge, Georgia

The two previous papers have presented a very iance (V p ) and is expressed mathematically by
thorough account of current information on heri-
tabilities for wood and morphological characteris- the equation h² = VG/Vp . Narrow sense
broad sense
tics. A primary purpose for determining herita- heritability is obtained by dividing additive vari-
bility figures is to better orient breeding methods. ance (VA ) by phenotypic variance (V p ) and can be
How, then, can this information be used in the
current tree improvement programs and how will it expressed by the formula h 2 =
Va/Vp
affect silviculture in the long run? narrow sense
Ten years ago the descriptive words "phenotype" If the narrow sense heritability figure is high,
and "genotype," commonly used in genetics, had then the mass selection method of breeding will be
very little meaning to management foresters. These most productive. On the other hand, if the dom-
terms are relatively common today and are em- inance or epistatic variation is very high, the great-
ployed freely in discussions of tree improvement est gain can be accomplished by an intraspecific
programs. They exemplify the new set of terms hybridization approach.
used in advancing techniques in genetics and breed- Figures on inheritance, represented by heritabil-
ing procedures. It will be necessary to define and ity values, are essential to the development of any
develop some other new terms as they are used tree breeding program. If, for example, specific
throughout this paper. gravity were controlled to a considerable extent by
Heritability, a term so freely used in previous environment, the field forester would be the one
papers, is defined in the tree improvement glossary who could most easily increase or decrease the
(Snyder 1959) as a "measure of the relative degree specific gravity of stands by silvicultural manipula-
to which a character (or characteristic) is influ- tions. The tree breeder would be able to add very
enced by heredity as compared to environment." little by genetic or breeding techniques. If, on the
Variation among trees can be expressed by formula other hand, as Mr. McElwee has shown, specific
as follows: gravity is controlled to a considerable extent by
Total Variation = Hereditary Variation + heredity, the silviculturist can do less to affect it,
Environmental Variation and management techniques must be quite drastic
to bring about major changes. However, the tree
A properly designed heritability experiment can breeder can accomplish a great deal.
separate variation due to heredity from the varia-
tion due to environment. Heritability is usually
expressed as a decimal or percent figure. For Initiation of a Tree Improvement Program
example, if a broad sense heritability of 0.7 or 70 Assuming that a tree improvement program is
percent is found for a characteristic such as specific about to be initiated, the ideal procedure is to make
gravity, it means that 70 percent of the total varia- a thorough study of the species to be improved.
tion in specific gravity is due to heredity and 30 Detailed information concerning variation between
percent is due to environmental influences. and within individuals must be gathered and a
What elements are included in the heritability complete study made of this species, with an assess-
portion of the estimate and what do they mean? ment of causes of variation over the entire range.
The term "broad sense heritability," or the total Heritability of important characteristics and the
hereditary variation, refers to a numerical estimate mode of inheritance must be determined. Con-
including three values: namely, additive variance, trolled crosses with related species should be made
dominance variance, and variation due to epistasis. and their progenies studied. Armed with this infor-
Additive variance is due to the average or additive mation the tree breeder could then devise a breed-
effect of a gene or genes. Each gene contributes a ing scheme and proceed with improving the species.
small addition to the overall effect. Dominance Since one rarely approaches ideal conditions,
variance is due to the interaction of alleles. Epista- chances are that a tree breeder will find himself
tic variance is due to the interaction between non- with the situation that confronted forest geneticists
alleles. The term "narrow sense heritability" refers approximately 10 years ago, when the first southern
to ratio of additive to phenotypic variation. pine tree improvement programs were initiated.
Broad sense heritability is determined by divid- Little of the necessary information was available
ing total genetic variance (V G ) by phenotypic var- as a base for an efficient breeding design. There

27
was not time to make the required surveys and Induced chromosomal changes.—Another means
conduct experiments that would provide knowledge for producing artificial genetic variation is to sub-
needed to choose among the several immediate ject seed or plants to X-ray ( Snyder et al. 1961)
avenues of approach available. These had already or other radiation (Beers 1962 ). This causes actual
been explored, developed, and exploited in various physical disturbances in one or several chromo-
crop improvement activities with other plant spe- somes. These can be large changes involving whole
cies. The problem in forestry in the Southeast was sections of chromosomes or minute "gene changes"
to determine which breeding method would result which are reflected in changes in the developing
in the greatest improvement in the shortest time seedlings; such seedlings with artificially induced
so as to provide improved seed for the very large changes or mutations have up to now been very
and expanding planting program then underway. difficult to keep alive, and this technique can also
be considered in the laboratory stage as a practical
Methods to Produce "New Creations" breeding tool.
Polyploidy.—The "go for broke" approach was Interspecific hybridization.—Crossing two species
characterized by polyploidy adherents. In this of pine is another method that sometimes produces
breeding system, the fundamental structure of the seedlings with spectacular characteristics. The re-
cell is manipulated to change its genetic constitu- sulting progeny are compared to each parent to
tion. Each cell in every species has two sets of a determine what improvement, if any, has been
fixed number of heredity units or chromosomes. gained. This phase of forest genetics has been
Pine species have 12; aspen, 19; human beings, 23. extensively pursued at the Institute of Forest Ge-
Normally this number is characteristic to the spe- netics at Placerville, Calif. Control crossing tech-
cies, but occasionally something occurs that upsets niques were developed here in the early 1930's,
the normal condition and then the resulting pro- and the major effort of this station has been di-
geny have an abnormal number of chromosomes. rected into the methodology and usefulness of inter-
There are certain plants which are improved by specific hybridization. Such crosses between species
such a change in chromosome number; some per- have produced some remarkable hybrids ( Calla-
sons have suggested that this might be true for ham 1957 ), but for one reason or another very few
forest trees. of them have been of practical use. Two of the
Probably the most intensive search for polyploid hybrids, the Jeffrey x Coulter and the knobcone
trees has been made in the aspen improvement pro- X Monterey, have been quite successful. Jeffrey
x Coulter hybrids are now being planted in Cali-
gram, where individuals were located that were
fornia forest plantations and the Jeffrey x Coulter
extremely vigorous and had exceptionally large
hybrid backcrossed to Jeffrey is also being planted
leaves. A microscopic examination of cells from
in the Jeffrey pine range where it is resistant to
these selections demonstrated that they had 3 sets of
the pine reproduction weevil (Righter 1960).
19 chromosomes instead of the normal 2 sets. Since
these trees were so very vigorous, it was hoped The most impressive practical use of the hybridi-
that such triploids might be especially desirable. zation technique has been developed in Korea
Geneticists artificially recreated triploid indi- ( Hyun 1961) where in 1 year over 1 million con-
viduals with colchicine, a chemical that interrupts trol-pollinated seedlings of a pitch pine x loblolly
the natural processes of cell division (Inst. Paper pine hybrid were produced. This hybrid has much
Chem. 1955). When a seed is placed in a solution of the growth habit of loblolly pine and some of
of colchicine at the time of germination, a few indi- the frost resistance of the pitch pine parent. The
viduals with twice the normal number of chromo- labor cost of such hand-produced hybrids would be
somes develop. An aspen seed with 2 sets of 19 prohibitive in this country, but a few have been
chromosomes can be forced to develop twice this produced to be used on an experimental basis.
number. This individual is partly fertile and, when Hybridization, as a completely practical method,
crossed to a normal aspen, develops seed which cannot be used until some inexpensive system of
grow into very vigorous seedlings with 3 sets of 19 control pollination is developed.
chromosomes. These triploids are normally sterile Some research has been done to develop practical
or have very low fertility so could not be repro- means of mass producing hybrids. Wakeley and
duced by seed. However, it might be possible to Campbell ( personal communication) have tested
produce triploids in sufficient number by establish- a method of applying slash pine pollen to unbagged
ing seed orchards of alternating rows of normal longleaf strobili, but with indifferent success. Brown
diploid and tetraploid individuals. If so, triploid and Greene (1961) and Hyun (1961) are working
individuals could be used in practical forest man- with chemicals that will cause male sterility.
agement if desired. If Wakeley's or some other simple method can
Although examples of polyploidy in pines have be developed so that successful hybrids are pro-
been reported, (Hyun 1954; Mergen 1958, 1959) duced consistently it would be relatively simple to
in every case the seedlings are malformed and grow make a mass collection of pollen and dust a large
poorly, with undesirable form; thus polyploid seed- number of trees in the seed orchard inexpensively.
lings in the pines are of little value. Polyploidy as On the other hand, if male sterility can be induced,
a breeding method in pines must be relegated to orchards could be so designed that the species to
the laboratory, at least for the foreseeable future. be crossed would be planted in alternating rows.

28
The pollen of the seed parents would be rendered liminary surveys indicated that variation was pres-
sterile so that all of the seed would be hybrid. ent and seed source important. In' - erspecific hybrid-
These techniques are still in the experimental stages ization had not been particularly promising. The
but may hold some promise. problem at hand was to develop a genetically im-
proved source of seed as soon as possible. The
Improving Existing Species decision was: mass selection.
Intraspecific hybridization.—The technique of The data now available testify to the validity of
making controlled crosses between members of the this choice for most characteristics. It must be re-
same species may be used to combine desirable membered that many of the experiments from
characteristics from different individuals. It always which these data have been drawn were designed
must be remembered that some selection is implied for other purposes and that much is from relatively
regardless of the breeding scheme used. However, young material, but the significance cannot be
the difficulty of making selections varies consider- denied. Heritability values made from measure-
ably with the type of breeding method in use, and ments of immature material can be expected to
the size of the population that must be examined. increase as the plantings come closer to maturity,
As the size of the population available for selection at least for some species. Let us now, as the poli-
and the number of characteristics being considered ticians say, examine the record.
increases, selection difficulties are compounded. Barber and McElwee have quoted heritabilities
The problem of selecting usable breeding stock for many characteristics and indicated the amount
is minimized with the intraspecific hybridization of improvement that can be expected based on these
method of plant improvement. Usually in this sys- figures. They have urged the immediate pursuit of
tem individuals with one particular outstanding an aggressive tree improvement program based on
trait are selected. The original selections are crossed the mass selection method of breeding, or some
in an attempt to combine the outstanding features variation thereof.
of each into one hybrid. The system is time-con- A brief review of some of the heritability figures
suming, though, especially with species that have quoted will emphasize the soundness of the recom-
considerable time lapse between seed germination mendation.
and the development of reproductive organs. After Some of the oldest plantations which can be used
the original crosses are made, progeny must be for estimating heritabilities were established by
grown and new selections made which contain the personnel of the Southeastern Forest Experiment
desirable characteristics. These are then cross bred Station at Lake City, Fla. Planting consisting of
to fix the characteristics in a high proportion of grafted clones, open pollinated progeny, and con-
the progeny and to establish a seed source. In order trol pollinated progenies are available for estima-
to prevent inbreeding depression, several selections ting the genetic improvement possible. Squillace
must be made and carried on concurrently. The seed and Bengtson (1961) have reported heritability
source is eventually developed from these selections. figures for several characteristics from these 10- to
Mass selection.—This is a system of breeding for 14-year-old plantations. Narrow sense heritabilities
improvement that promises slow steady gains. of 56 percent for specific gravity were obtained with
Normally, the increase one can expect is limited by control pollinated progeny while the broad sense
the genetic capability of the most outstanding indi- or total heritability was estimated to be 73 percent
vidual in the population. With a large amount of from clonal data. Fairly high heritabilities for
natural variation of the additive type, large in- diameter growth (25 to 58 percent), stem volume
creases can be expected; conversely, with little (18 to 35 percent), crown width (24 to 48 percent),
natural variation, there can be little improvement and bark thickness (33 to 67 percent) were ob-
by this method. tained. A fairly low heritability for height growth
As the number of characteristics to be improved of 5 to 10 percent was obtained. A tentative hypo-
increases, selection difficulty increases. In order thesis advanced by Squillace indicates that this is
to make the greatest possible gain a high intensity probably due to the wide spacing (20 by 20 feet)
of selection must be practiced—and suitable indi- of the plantation and the attendant lack of compe-
viduals are difficult to locate. On the other hand, tition. This is somewhat substantiated by Barber's '
once the selections are made, seed production prob- figure of 27 to 37 percent for material planted at
lems are simplified as a continuous supply of large a spacing of 10 by 10 feet, van Buijtenen's figure
quantities of seed can be produced from vegeta- of 20 percent for material planted at an 8 by 8
tively propagated ramets of the original selections. foot spacing (personal communication), and by re-
cent results obtained by Stonecypher (personal
Practical Tree Improvement communication) from seedlings planted also at an
8 by 8 foot spacing. It is interesting to note that
With these methods available, the pioneer south- the broad sense heritability of oleoresin yield is
ern pine tree breeders took a long, hard look at estimated at 90 percent while the narrow sense
their species. Little or no information was avail- estimates vary from 45 to 90 percent depending on
able from previous data to guide them. A few pre- the methods used.
' Barber, John Clark. An evaluation of the slash pine progeny tests of the Ida Cason Callaway Foundation (Pinu s elliottii
Engelm.). Ph.D. Diss., Univ. Minn. 206 pp., illus. 1961.

29
 van Buijtenen (1962) estimated broad sense ally applied is through seed orchards. Individual
heritability of 64 percent and 84 percent from elite selections are made only after the examination
5-year-old grafts for specific gravity, and narrow of thousands of acres of forest land. These selec-
sense values of 37 and 49 percent for 2-year-old tions are rigidly graded in comparison to a number
control pollinated material. He also reported dia- of surrounding dominant specimens, and included
meter heritability of 20 percent for 6-year-old lob- in the orchard only after they indicate a maximum
lolly pine progenies. He estimated that an improve- amount of advantage. The selections are vegeta-
ment of 10 percent for each of these characteristics tively propagated and planted in a central orchard
may be expective from one cycle of selection and location, or a seed orchard of seedlings from the
that an increase of 25 percent in total wood produc- selected parents is established. The orchard is de-
tion for the several factors combined would not signed to insure a minimum amount of inbreeding.
be out of reason (personal communication). Here the trees are cultivated as intensely as in
Stonecypher (personal communication) found a fruit orchards and will serve as a source of high
low narrow sense heritability for first year height quality seed for the future.
growth, which increased with second year measure- Based upon the figures quoted today and if we
ments. He noted that narrow sense specific gravity take into consideration improvement due to in-
heritability estimates for 1- and 2-year-old pro- creased vigor, finer limbs, straighter boles, and less
genies approach the actual values given by van disease, it seems probable that the yield increase
Buijtenen and that they remain constant for the of 10 percent suggested by Barber can be obtained
2 years. with ease and the suggested figure of 25 percent
advanced by van Buijtenen is within reach. As-
Selection severity can be varied from slight to suming no change in pulpwood stumpage values
intense, with selection of low to medium intensity. from the figure advanced by Perry and Wang
Forestry practices can be guided so that any land- (1958) and figuring a 20 percent increase in yield
owner can achieve a moderate amount of genetic due to tree improvement efforts, we can realize a
improvement in his forest stands. Some gain will gross increased profit of some $600 per pound of
be achieved, if, in a seed tree cutting, selection of seed at a 25-year rotation, or an extra $2.10 per
seed trees from which cones will be collected is acre per year. This profit, they say, justifies the
made as the first step in harvesting. Improvement expenditure of $181.27 per pound of seed, allowing
here is controlled by the nature and number of 5-percent interest on the invested money.
trees in the stand being cut. There is no doubt that
a minimum amount of improvement can be ex- Perry's figures are based on yield alone, and no
pected when the selections represent only 10 to attempt was made to include other advantages
20 percent of the stand, which is normally the case. which cannot be represented easily by monetary
As soon as the area is sufficiently stocked, the seed values—for example, the morphological character-
trees are harvested during a year of heavy cone istics of bole straightness. It is difficult to deter-
production. Such areas are designated as seed mine how much more solid wood content would be
collection areas. Careful planning is necessary as delivered per cord by minimizing the amount of
harvesting of the seed trees must be scheduled to crook, spiral, and sweep. It is difficult to learn
coincide with the optimum period for cone collec- how much more cellulose is in each cord and how
tion. Cost of cone collection from trees which have much less cooking liquor will be needed at the
been cut is relatively inexpensive, especially when mill. One can only estimate the increase in usable
large crops are present. fiber contained in the straight pulpwood stick. Add
to these figures a decrease in knot wood volume, a
The next highest level of selection severity is concomitant decrease in compression wood asso-
represented by the seed production area, where ciated with knots, and an increase in the number
better than average stands are selected, carefully of seedlings growing to maturity by virtue of in-
rogued of undesirable individuals, and managed creased disease resistance. Consider also the pulp
for a continuing supply of genetically improved increase reported earlier by McElwee due to in-
seed. Since mature trees must be climbed for cone creases in specific gravity and the probable im-
collection it would seem at first that the expense provement in paper sheet formation due to having
would be prohibitive, but collection costs as low wood with more uniform fibers. All these advan-
as $3 to $5 per bushel have been attained as com- tages, nebulous as they may be, are to be gained
pared to an average cost of $2.50 per bushel when as a result of the activities now taking place. Esti-
cones are purchased on the open market (Goddard mates of possible improvements made several years
1958; Cole 1962). According to Easley (1963), ago covered the entire scale from a minimum 1, 2,
seedlings from one seed production area had a and 5 percent made by the conservative members
height advantage of 24 percent at the age of 8 of this group to a maximum 100 percent by the
years on sandy soil as compared to nursery run most optimistic. Actual values today indicate an
seed; this same seed production area produced intermediate expected increase of 15 to 25 percent
seedlings with a height advantage of 7 percent on based on yield, plus the additive increment due
heavy clay soil. He concludes that the collection to quality. Perry's figures, which seemed so un-
of seed from a local source of selected parent stock obtainable 5 years ago, are becoming more realistic
is advantageous. as we gain additional knowledge of the inheritance
The most severe selection that can be commerci- patterns in the species we are using.

30