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'10

Api seedling plant undergo shifts in their "ontogenetic age" as cells in their apical meristem divide and lay down new tis sue. Axillary buds typically are dormant due to apical dominance, but retain lhe ontogenetic age at origino Once lateral buds start to grow they then undergo change in maturation in lhe same manner as the apical bud, but in their own pattem.

FIGURE 16-15

Apical Meri

eovers a mutation (79). This situation also oeeurs in the budding offruit and nut tree cultiv Although the probability for any single mutation to oceur may be low, the probability of ing it is high because of the single bud propagation and the volume of plants involved. T; explains why off-type plants are usually detected in a commercial orchard rather than in -source orehard where the propagation material was obtained. li the eharacter affects flo ing or fruiting, the mutant may go undetected for a number of years after the plant is pl into its permanent location.

MANAGEMENT OF PHASE VARIATION DURING VEGETATIVE PROPAGATION

"Aging" phenotypic changes with age due either to ontogenetic or to chronological change. or both.

Ontogenetic Versus Chronological Aging in Plants

Ontogenetic Aging
"Aging" has two separate meanings in plant development. Ontogenetic aging refers to phases of development that the seedling plant undergoes from embryonic to juvenile to in mediate to mature (adult) as was described in Chapter 2 (Figure 2-17). The transition fr seedling to flowering plant can be very rapid in annual plants (within days) but can take ove20 years for some trees. Although it is eonvenient to express the length of time required make this transition in days or years, phase change may best be associated with the number nodes that have been developed in the apieal meristem (Figure 16-15). Buds produced at a p ticular node retain the epigenetic potential of that node with respeet to its ontogenetie age. F example, buds produeed at the base of the plant tend to be more juvenile because they we~ formed first during the juvenile phase of development. Phase change is best identified by morphologieal characters for a specific phase, such leaf shape (Figure 16-16) or the ability to flower. In some plant speeies, there is very little o vious change in the plant's appearanee as it grows from a seedling to its mature condition. behavior is called homoblastic. For other plant speeies, there is a distinct variation in specific traits that oceur with age, which is ealled heteroblastic. Several genes and their gene produ

Homoblastic phase changes that involve little obvious change in phenotype with development and age.

Heteroblastic phase changes that show distinct change in phenotype during development.

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Phase changes can be considered to be biologically equivalent in animals, humans, and plants in that embryonic, juvenile, adolescent, and adult phases are present in each. The difference is that animals and plants differ in the way they grow and age. Cells in animal (and human) development are more or less determinate (i.e., cells in all parts of the body mature and age more or less together). Plants, on the other hand, grow from consecutive cell divisions in the meristem of apical and lateral shoots laid down in sequence over time. Juvenile to mature change occurs in the apical

Equivalence of Phases in AnimaIs and Plants

meristem as the shoot grows, but cells remaining behind retain their initial ontogenetic age. In perennials, ontogenetic aging continues in altemating cyc1es of grawth and dormancy. ln annuals and biennials it terminates in reproductive structures and the plant dies. The phase of maturation in different locations of the plant is deterrnined by the pattem of bud development in the plant. Juvenile and mature characteristics, such as flowering, leaf shape, or thorniness, can be used as markers of maturation, their location depending on the chronological age at which ontogenetic maturity is attained in particular parts of the plant.

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The comparison of the two types of "aging" produces a paradox in horticultural terminology. That part of a seedling plant nearest to its base is the "oldest" in terms of chronology but actually "youngest" (i.e., more juvenile) in terms of maturity (onto-

Paradox in Terminology

genetic) age. Likewise, the outer peripheries of the stems and branches are the "oldest" in maturity but "youngest" in chronology (15, 35, 74). The age of a vegetatively prapagated plant depends on the ontogenetic age at which the plant was prapagated.

have been identified that are differentially expressed in the juvenile or mature. These can be used as molecular or biochemical markers of phase development (57). Chronological aging continues through the life of an individual plant whether seedling or vegetatively prapagated. This characteristic may best be designated by the number of years that the plant has grown either fram seed or vegetative prapagule. When young and growing under favorable conditions, the plant should be vigorous, healthy, and may become very floriferous, Eventually, as the plant gets older, vigor declines and, although flowering and fruiting may remain abundant, new growth will tend to be reduced. The plant eventually becomes enescent and may eventually die. Plants appraaching the senescent stage can be returned to vigorous and productive state by reprapagation or by applying various horticultural practices, such as pruning, nitragen fertilization, irrigation, controlling pests and diseases, and, in general, applying good management practices.

Determinate growth habit with shoot apex ending in flowering.

Senescence physiological aging assoeiated primarily with reduced vigor.

Phase Variation and Vegetative Propagation

Ontogenetic aging is important in vegetative prapagation because buds (both terminal and lateral), when removed for vegetative propagation, perpetuate their ontogenetic age in the progeny plant. For instance, if the source plant was juvenile (or the prapagule was taken fram a ruvenile part of the plant), the new progeny plant should initially express the juvenile phase f the source plant but shift toward the mature (aduIt) phase in subsequent growth. Eventually the plant stabilizes at the mature reproductive phase of development. Vegetative propagation from mature branches of a seedling plant consequently may not reproduce the seedling phenotype in the SI generation. Prapagules taken fram the base of the lant tend to praduce biologically juvenile plants; those taken from the top or at the periphery

Juvenile phenotype characteristics associated with the juvenile phase in specified species, usual! y upright growth, vigor, sometimes thorniness, and lack of flowering.

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Adult phenotype characteristics associated with the adult phase, typically bushy growth, reduced vigor, no thorns, and profuse f1owering.

of the plant tend to produce progeny that are biologically mature; the entire range may be produced if plants from intermediate locations are included. The recognition of this variabili . phenomenon by foresters led to the coining of the terms ortet (the So plant) and ramet ( plants) (138). These variations are not due to genetic change but to epigenetic variation in maruration that persists during vegetative propagation.

Juvenile Versus Adult Phenotypes

Continuous growth and consecutive generations of vegetative propagation leads to the stabilization (i.e., "fixing") of the mature phase in individual plants of the clone and the disappearance of seedling phase characteristics. Most clonal cultivars grown for their fruits, seeds, flowers exhibit the mature growth phase of the clonal cycle as the typical "true-to-type" forra Most individual plants come into flower and fruit reasonably quickly after a vegetative phase of development and undesirable traits associated with plants in the juvenile phase disappear. is not appropriate to refer to the vegetative phase of the stabilized mature clone as "juvenile," There are many examples of the application of phase changes in horticulture. Seedlings of many fruit and nut species, such as apple (Malus), pear (Pyrus Citrus), an walnut (Juglans), may take a long time to come into flowering (see Table 2-1). Grafted cultivars in the mature phase not only show genetic uniformity, but also flower and produce fruit or nuts at an earlier age (91) and exhibit a more desirable tree formo Similarly, a flowering ornamental plant grown from seed may require many years to flower. For example, seedling Wisteria and Magnolia take 7 and 15 years, respectively. to flower whereas vegetatively propagated plants flower at a younger age (90). The val of seed orchards of specific genetic sources in forestry results from their production at early age as compared with their natural behavior as seedlings (116, 167). Orchids, bulb species, and other herbaceous perennials require five to ten years of annual development from seed before flowering occurs. Once the mature (adult) stage is reached, annual flowering will occur with alternate phases of vegetative growth (bulbs, corms, pseudobulbs) and reproduction (flowering) (see Chapter 15). Some ornamental vines, such as Hedera sp. or strangling fig (Ficus sp.), are primarih vines in the juvenile phase, whereas the mature flowering phase is treelike (Figure 2-18). Both forms may be useful but for different purposes. Eucalyptus and Acacia have strikingly uni que juvenile foliage (Figure 16-16). Some conifer cultivars have juvenile foliage (needle or awl-like) distinctly different from the adult foliage (flat, rounded, or scalelike) (90). Some cu1tivars (Chamaecyparis pisifera 'Boulevard' and Cryptomeriajaponica 'Elegans') are more or less "fixed." Others (Pinus sylvestris, Cryptomeria japonica, junipers (Juniperus), and Lawson cypress (Chamaecyparis lawsonia) may develop both types of foliage. The juvenile phase of most plants inherently has a higher rooting potential than the mature phase. The application of juvenility to cutting propagation was described in dep; in Chapters 9 and 10. The easy-to-root juvenile vine form of ivy (Hedera) has been a model system for studying root initiation (56, 57). The juvenile phases of the conifers listed in the previous paragraph invariably root easily and can be maintained as cultivars.. The rooting potential of many if not most tree species is correlated closely with seedling age and the retention of the juvenile phase (Figure 16--17). A reason previously cited fOI culti vars of fruit and nut crops to be propagated by grafting was their di:fficulty in rooting associated with their mature phase. For clonal rootstocks, selection and maintenance of the juvenile phase are important strategies for propagation in order to improve rooting potential. See the text box on page 611 for a description of a unique forestry breeding and propagation system that involves both seedling selection and clonal propagation. The juvenile apomictic seedlings of Citrus are very different from the typical mature phase of the cultivar propagated for commercial production. The seedlings are very vigorous, thorny (see Figure 2-21), often have different shaped leaves, show lowproduction, and have fruits with very inferior quality. On the other hand, the traits of uniforrnity and vigor make the apomictic seedlings highly desirable rootstocks (21, 50, 147), for which purpose they are used.

Trueness to type corresponding to phenotypic characteristics of the source planto

"Bulking up" in forestry propagation this term refers to using vegetative propagation to multiply the supply of se!ected genotypes.

CHAPTER 16

PRINCIPLES

fi PRACTICES OF CLOl\AL SELECTIOl\

611 FIGURE 16-16

Foliage variation gradients within seedling trees can be markers of juvenile and mature phenotype. Top left: Eucalyptus seedling tree shows the large, broad, sessile (no petiole) leaves aI lhe base of lhe tree; mature leaves from lhe upper part of lhe lree are narrow. Top right: Gradient in leaf shape is shown by a single shoot. Lower left: ln Acacia melanozylon, lhe bipinnate leaves et lhe base are characteristic of lhe juvenile phase. The leaves at lhe 10p of lhe seedling are expanded petioles (phyllodia) and are characteristic of the adult formo Lower right: Needles of lhe basal juvenile part of many conifers lend lO be acicular (sharp pointed), whereas needles on the upper, more mature part tend lo be scalelike.

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(a) Seed is taken from plants of seedling families that have been previously selected for superior growth rates, form, disease resistance, and so on. (b) These seedling blocks (referred to as ortets) are kept in a juvenile/transition phase by hedging and shearing. (c) Cut(ramets) are obtained and can be rooted because of

their juvenile condition. (d) Rooted plants are transplanted to timber production sites for evaluation over time. (e) Superior ortet farnilies are consequently identified. (f) Cuttings, taken from the seedling plants of these selected elite families which have been maintained by hedging and shearing are multiplied into commercial volumes ("bulking up"). (g) Clonally generated ramets are established in the forest planting in a mosaic of different clones to avoid monoculture production and to reinforce genetic diversity.

612 FIGURE 16-17

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VEGETA TIVE PROPAGATION

Graph showing the effect of age on donor tree relating to key tree characteristics that are important in cJonal forestry. Left: Phenotypic trueness-totype is illustrated by size of stem diameter and shoot length of grafts after multiple years. Right: rooting percentages of cuttings from donor trees of different ages. (From Greenwood et aI., in Clonal Forestry 11, 1993 by permission.)

E ...

6 5 100

C1) C1)

.S! ~
c:

:g
c
(.J

.s

c: :;::

CI

....
40

Eastern Larch
... ...---t)6

~ o
..... E (.J

20

30

40

50

Age donor tree

00

20

40

60

Age (years)

"Clonal forestry" depends on the ability to mass-propagate populations of juvenile prapagules that not only exhibit easy rooting (Figure 16-18, right), but also praduce the desirable juvenile "seedling'' phenotype (Figure 16-18, left) (2). The desirable "true-to-type" form of forest trees grown for lumber and wood is straight-tapered trunk, reduced branching, lateness to flower, and other juvenile characteristics. Vegetatively prapagated plants of radiata pine (Pinus radiata), loblolly pine (Pinus), giant sequoia (Sequoiadendron), and eastern larch (Larix) taken fram more mature locations in the tree showed reduced height and diameter, and more branching (53). Success in "clonal forestry" depends on the ability to contrai maturation and to select the proper "true-to-type" stage (2, 116). See the text box on page 611 (24, 116, 129).

Promoting Shifts from Iuvenile to Mature Phases

The long period of time required for seedling plants of many woody species to come ine flower (Table 2-2) has important applications in horticulture. Breeding programs of woodj plants primarily use seed1ings, but pragress can be hampered by their long juvenile peri (61). Breeding can result in shorter breeding and praduction cycles, since the juvenile period is controlled by genes (156). Growth and development can also be managed by environmental control. The most important concept is to keep the plant continuously grawing in order t grow through the juvenile phases as quickly as possible. For example, birch seedlings gro continuously under long days in the greenhouse during winter passed through the juvenile imo the adult phase much faster than under normal short-day winter conditions (1l7). Crab apple (Malus hupehensisi seedlings grown continuously in a greenhouse to 2.5 to 3.0 m (8 to 10 fi started to flower in 13 months from germination instead of four years (166). Restricting growth, girdling, bending, growing on dwarfing rootstocks, and other horticultural practices are often cited to induce flowering (160). However, these are not useful until the plant approaches the transitional ar mature phase. Prior to that phase, growth should be encouraged to aliow the plant to grow through the juvenile stage.

CHAPTER 16

PRINCIPLES AND PRACTICES OF CLONAL SELECTION

613

Epicormic a shoot emerging from a latent bud on the base of a tree; synonymous with watersprout.

o
Cone of juvenility a cone-shaped area comprising the trunk and lower branches of a seedling tree, which tends to remain juvenile.

Ontogenetic juvenile-mature gradients occur in seedling trees from the base of the tree to the topo Left: Seedling conifer. Juvenile root-shoot junction occurs at A; flowering occurs first at the mature apex B. Other letters refer to shoots at different locations. Gradient in juvenility = A> B > E > D > C > F due to differences in ontogenetic age. Right: Seedling deciduous tree. Juvenile root-shoot junction occurs at A; flowering occurs first in the mature part of tree at the apical part (G). Juvenile structures arising from the "cone of juvenility" near the base of the tree include adventitious root "sucker" (B), watersprout (epicormic) (C), and sphaeroblast (D). A stump sprout from pruning (E) and a hedged tree (F) are also shown. (Redrawn with permission from Bonga [16].)

FIGURE 16-18

Selection and Maintenance of the Iuvenile Phase

Retention of the juvenile phase in a propagation source block may be important to increase rooting potential (see Chapter 9). Following are methods to select, delay, or preserve propagation sources in the juvenile phase: 1. Collect material and propagate from plant parts showing juvenile characteristics. This may mean collecting from the base of a plant in the case of a seedling. Some cultivars are more or less "fixed" as juvenile (e.g., various conifer cultivars with juvenile foliage). 2. Collect material from root sprouts (suckers) or from watersprouts (epicormic shoots) arising from near the base of the planto These arise from latent lateral buds that are located in the "cone of juvenility." The shoots may be forced by cutting back the plant to the ground to produce "sturnp sprouts" (47). 3. Some plant species produce sphaeroblasts (i.e., masses of adventitious buds located on the lower portion of the trunk). Examples of species with such characteristics include sequoia (Sequoiadendron), redwood (Sequoia) (17), some quince cultivars (Cydonia oblonga), and some apple (Malus) cultivars (139). Shoots developing from these structures invariably are juvenile, show characteristic morphology, and are easy to root. 4. Establish hedge rows of stock plants produced by propagating juvenile material (see Chapters 10 and 11). This method is perhaps the most important to provi de large populations of juvenile cutting material (84). Plants are severely pruned annually to a constant height to produce a continuous supply of new shoots. The original material may be seedling populations (115), rooted cuttings, or grafted plants on rootstocks. For the latter, care must be taken to eliminate rootstock shoots or choose a rootstock with morphologically distinct leaf characteristics. - Juvenile material can also be maintained by consecutive ("serial") propagations of several generations from seedling material (2). Serial propagation is practiced in the

Stump sprouts vigarous shoots that are produced from the stump when a tree is pruned back severely.

Sphaeroblasts masses of adventitious buds produced on the lower trunk of some tree species.

Hedge row a row of trees or bushes pruned back to a hedge to stimulate shoots for propagation.

Serial propagation propagation of seedling ar clonal material for several consecutive generations in containers, each propagated after pruning.

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Mature shoot

Juvenile shoots

Rejuvenation produced by consecutive (serial) grafting from a ten-year-old mature Eucalyptus tree. Left: Plant produced by cleft graft of scions from the ten-year-old tree onto a seedling rootstock showed mature leaves. Middle: Plant produced from scions after three serial grafis. One of the shoots shows mature characteristics, the other juvenile. Right: Plant produced after grafiing with scions afier six serial grafts. AlI of the shoots produced juvenile (large) leaves. Cuttings from this plant rooted successfully, whereas rooting from the ten-year-old original mature tree was not successful. (From Sinisca1co and Pavolettoni (126) with permission.)

FIGURE 16-19

annual propagation of new generations of cuttings-collected from asexually propagated container-produced woody and herbaceous ornamentals. 6. Establish "stoolshoot" beds by cutting shoots nearly to the ground (90). This is perhap one of the reasons for success in rooting in mound and trench layering (see Chapter 1Rejuvenation shift from the adult phase to the juvenile or transitional phase.

Reversions from Mature to Iuvenile Phase (Rejuvenation)

The mature phase naturally reverts to the juvenile phase during seed reproduction whether se ual ar apomictic (75). However, reversions have also been induced from the mature to the jcvenile phase in vegetative material. Examples inc1ude the following: 1. Grafting consecutive generations of scions in the mature phase to seedling rootstocks (17,47) has produced reversion from mature to juvenile phases (Figure 16-19). Extensive results have been achieved with Sequoia (66), Eucalyptus (126), and other forest tree species (15, 16). 2. Consecutive subculturing of apical meristems in micropropagation has resulted in the reversion of p1ant shoots to the juveni1e phase (see Chapter 17). For instance, Mullins (99) found that cultured somatic embryos from the ancient clone 'Sauvignon Blanc' produced "seedlings" with typical juvenile foliage. Micropropagated cultivars used as stock plants have resulted in increased rooting both in vitro as well as from tis sue- and culture-produced plantlets established as stock plants (see Chapter 17) (64, 109). 3. Some hormone treatments have resulted in reversions under experimental conditions. For example, applications of gibberellic acid have induced juvenile growth in ivy (Hedera), which couId be reversed with abscisic acid (56).

PATHOGENS AND PLANT PROPAGATION

The presence of disease can be the most important limiting factor in both plant production an:. plant propagation. In this section, the unique relationships between specific plant pathoge and vegetative propagation will be exarnined. In nature, plants have coexisted with many oth arganisms, inc1uding insects, mites, fungi, bacteria, and nematodes, to evolve more or 1 compatible interdependent communities. In some cases, such as mycorrhizal fungi, plants anc microorganisms form mutua11y beneficial associations (31, 33,109,134,135,155). When h mans began to cultivate plants the balances changed. Humankind began to propagate specific kinds of crop plants and adopt monocultural systems, which were more vulnerable to atta -