Preformed and neoformed extension of shoots and sylleptic branching in relation to shoot length in Tsuga canadensis

Summary Shoot systems developed over 3 successive years were investigated on 55 understorey Tsuga canadensis (L.) Carr. trees. Paired comparisons of preformed-leaf content of terminal buds and numbers of leaves produced on new shoots showed that neoformed leaves were produced in large numbers. Parent-shoot character was not useful in predicting numbers of preformed leaves, was better related to total leaves produced, but left the majority of the variation unexplained. This reflected the capacity of any terminal bud to produce a shoot with more or less neoformation, depending on conditions for growth. All shoots over 6 cm long produced sylleptic shoots that bore from two to many leaves and were arranged in a mesitonic pattern along the parent. Some of the longer sylleptic shoots produced lateral buds or second-order sylleptic shoots. Monopodial second-year extensions of sylleptic-shoot axes followed an acrotonic pattern, as did proleptic shoots from the few lateral buds borne on the parent shoots. Such lateral buds were more frequent on shorter parent shoots: they typically occurred near the proximal and distal ends. Duration of shoot extension was positively correlated with shoot length: terminal buds became evident as shoot extension neared cessation.     

Periods of organogenesis in mono- and bicyclic annual shoots of Juglans regia L. (Juglandaceae)

The organogenetic cycle of shoots on main branches of 4-year-old Juglans regia trees was studied. Mono- and bicyclic floriferous and vegetative annual shoots were analysed. Five parent annual shoot types were sampled between October 1992 and August 1993. Organogenesis of summer growth units was monitored between 16 Jun. and 3 Aug. 1993. Variations over time in the number of nodes, cataphylls and embryonic green leaves of terminal buds were studied. The number of nodes of parent shoot buds was compared with the number of nodes of shoots derived from parent shoot buds. The spring growth units of mono- and bicyclic shoots consist exclusively of preformed leaves which were differentiated, respectively, during the spring flush of growth (mid-April until mid-May) or the summer flush of growth (mid-June until early August) in the previous growing season. Thus, winter buds may consist of flower and leaf primordia differentiated in two different periods during annual shoot extension. The summer growth units of bicyclic shoots consist of preformed leaves that were differentiated in spring buds during the spring flush of growth in the current growing season. Bud morphology is compared between spring and summer shoots.     

Periods of organogenesis in shoots of Nothofagus dombeyi (Mirb.) oersted (Nothofagaceae)

The organogenetic cycle of main-branch shoots of Nothofagus dombeyi (Nothofagaceae) was studied. Twelve samples of 52-59 parent shoots were collected from a roadside population between September 1999 and October 2000. Variations over time in the number of nodes of terminal and axillary buds, and the length, diameter and number of leaves of shoots derived from these buds (sibling shoots) were analysed. The number of nodes of buds developed by parent shoots was compared with the number of nodes of buds developed, I year later, by sibling shoots. The length, diameter and number of leaves of sibling shoots increased from October 1999 to February 2000 in those shoots with a terminal bud. However, extension of most sibling shoots, including the first five most distal leaf primordia, ceased before February due to abscission of the shoot apex. Axillary buds located most distally on a shoot had more nodes than both terminal buds and more proximal axillary buds. The longest shoots included a preformed part and a neoformed part. The organogenetic event which initiated the neoformed organs continued until early autumn, giving rise to the following year's preformation. The absence of cataphylls in terminal buds could indicate a low intensity of shoot rest. The naked terminal bud of Nothofagus spp. could be interpreted as a structure less specialized than the scaled bud found in genera of Fagaceae and Betulaceae.     

The role of the apple spray programme in the protection of fruit buds from powdery mildew

Comparisons of programmes, which differed in the dates on which high-volume sprays of dinocap (0.019 %) were omitted, indicated that applications in the period from green cluster to early fruitlet protected the bourse buds of apple cv. Cox's Orange Pippin from infection, and so controlled the number of mildewed blossom trusses in the following year. This role of the sprays in preventing primary mildew declined after the early fruitlet stage. The existence of two phases of the disease was confirmed, and although dinocap at late blossom and early fruitlet contributed to the control of preliminary infections of secondary mildew on vegetative shoots, the latter phase did not become severe until after the peak invasion of bourse buds. Fewer than 20% of the fruit buds which produced primary-mildewed blossom in 1968 were succeeded by healthy fruit buds in the next year; this poor recovery was not improved by dinocap applied during the flowering period in 1968.
Fruit-set and crop of James Grieve and Cox's Orange Pippin were reduced in 1968 by dinocap applied at green cluster and pink bud, or at full blossom and petal-fall, but no deleterious effects were produced by equivalent sprays in 1969. The need for mildew control during flowering is discussed in relation to phytotoxic sensitivity.     

Evidence that cytokinin controls bud size and branch form in Norway spruce

Shoot elongation in many coniferous species is predetermined during bud formation the year before the shoot extends. This implies that formation of the primordial shoot within the bud is the primary event in annual shoot growth. Hormonal factors regulating bud formation are consequently of utmost importance. We followed the levels of the endogenous cytokinins zeatin riboside (ZR) and isopentenyladenosine (iPA) in terminal buds, whorl buds and lower lateral buds of the uppermost current-year whorl shoots of 15- to 20-year-old trees of Norway spruce [ Picea abies (L.) Karst.] from June to September. Cytokinins were isolated with affinity chromatography columns, purified by high performance liquid chromatography, and quantified by ELISA. The level of ZR was low in June but increased gradually in all buds until September. Throughout the measurement period, the ZR level was highest in terminal buds and lowest in the scattered lateral, buds, with the whorl buds intermediate. The level of iPA peaked in July and decreased later without any consistent differences among the three classes of buds. The development of different kinds of buds was followed by scanning electron microscopy. We found that bud growth was greatest during August and September. The final size of primordial shoots within the buds varied considerably and the weight of the terminal bud was three times that of the whorl buds and more than five times that of the other lateral buds.
We conclude that the increase in ZR level during the period of active bud development is indicative of the importance of cytokinin for this process. Furthermore, the positive correlation between the level of ZR and bud growth during the period of predetermination of next year's branch growth suggests that this hormone indirectly controls the form of single branches in the spruce tree.     

Evidence that cytokinin controls bud size and branch form in Norway spruce

Shoot elongation in many coniferous species is predetermined during bud formation the year before the shoot extends. This implies that formation of the primordial shoot within the bud is the primary event in annual shoot growth. Hormonal factors regulating bud formation are consequently of utmost importance. We followed the levels of the endogenous cytokinins zeatin riboside (ZR) and isopentenyladenosine (iPA) in terminal buds, whorl buds and lower lateral buds of the uppermost current-year whorl shoots of 15- to 20-year-old trees of Norway spruce [ Picea abies (L.) Karst.] from June to September. Cytokinins were isolated with affinity chromatography columns, purified by high performance liquid chromatography, and quantified by ELISA. The level of ZR was low in June but increased gradually in all buds until September. Throughout the measurement period, the ZR level was highest in terminal buds and lowest in the scattered lateral, buds, with the whorl buds intermediate. The level of iPA peaked in July and decreased later without any consistent differences among the three classes of buds. The development of different kinds of buds was followed by scanning electron microscopy. We found that bud growth was greatest during August and September. The final size of primordial shoots within the buds varied considerably and the weight of the terminal bud was three times that of the whorl buds and more than five times that of the other lateral buds.
We conclude that the increase in ZR level during the period of active bud development is indicative of the importance of cytokinin for this process. Furthermore, the positive correlation between the level of ZR and bud growth during the period of predetermination of next year's branch growth suggests that this hormone indirectly controls the form of single branches in the spruce tree.     

LEAF DIMORPHISM IN POPULUS TRICHOCARPA

Critchfield , William B. (Pacific SW Forest & Range Expt. Sta., Berkeley, Calif.) Leaf dimorphism in Populus trichocarpa. Amer. Jour. Bot. 47 (8) : 699–711. Illus. 1960.—In Populus trichocarpa and other species of Populus, each tree bears 2 kinds of leaves, referred to here as “early” and “late” leaves. Both leaf types are present on all long shoots. They differ in many features of external morphology, including petiole length, size and occurrence of marginal glands, venation, and stomatal distribution. This type of foliar dimorphism has its origins in a pronounced difference in leaf ontogeny. The early leaves originate in the developing bud and overwinter as embryonic leaves. The first late leaves are also present in the winter bud, but as arrested primordia, and succeeding late leaves are initiated at the tip of the growing shoot and develop uninterruptedly to maturity during the growing season. A similar correlation between leaf form and the circumstances of leaf ontogeny appears to be a common feature of many other instances of heterophylly. The expansion of the pre-formed early leaves is almost completed by late spring, when the first late leaves begin to grow rapidly. The formation of late leaves may then continue until late in the season. The rapid elongation of the stem does not begin until the first late leaves expand. Elongation is restricted to shoots producing late leaves. Consequently, the early leaves are confined to short shoots and the base of long shoots; adventitious shoots and the upper part of long shoots bear only late leaves. Certain other woody plants with long and short shoots also exhibit a restriction of elongation to those shoots on which a second set of leaves is produced.     

Changes of morphogenic competence in mature Pinus sylvestris L. buds in vitro

The effects of season and cold storage on morphogenic competence in mature Pinus sylvestris buds were investigated. Peroxidase and polyphenol oxidase activity were measured as markers of oxidative metabolism. No growth in vitro was observed on explants detached from the end of January until the beginning of March. Brachioblasts, each with a couple of needles, formed on 11% of the buds without macrostrobili that were detached in early April and introduced immediately into culture. Of the explants detached in late July, 15% formed shoots with brachioblasts and needles. The lowest activity of peroxidase and polyphenol oxidase in pine buds was observed from the end of April until the beginning of June when morphogenic competence of tissues started to increase. Development of bud explants detached in January was achieved by cold storage for 5 months. Low polyphenol oxidase and peroxidase activity coincided with increased morphogenic potential. Results suggest that reduced or stable activity of peroxidase and polyphenol oxidase is associated with an increased ability of tissues to start growth in vitro.     

Timing of reproductive and vegetative development in Saxifraga oppositifolia in an alpine and a subnival climate

Morphogenesis of floral structures, dynamics of reproductive development from floral initiation until fruit maturation, and leaf turnover in vegetative short-stem shoots of Saxifraga oppositifolia were studied in three consecutive years at an alpine site (2300 m) and at an early- and late-thawing subnival site (2650 m) in the Austrian Alps. Marked differences in the timing and progression of reproductive and vegetative development occurred: individuals of the alpine population required a four-month growing season to complete reproductive development and initiate new flower buds, whereas later thawing individuals from the subnival sites attained the same structural and functional state within only two and a half months. Reproductive and vegetative development were not strictly correlated because timing of flowering, seed development, and shoot growth depended mainly on the date of snowmelt, whereas the initiation of flower primordia was evidently controlled by photoperiod. Floral induction occurred during June and July, from which a critical day length for primary floral induction of about 15 h could be inferred. Preformed flower buds overwinter in a pre-meiotic state and meiosis starts immediately after snowmelt in spring. Vegetative short-stem shoots performed a full leaf turnover within a growing season: 16 (+/-0.8 SE) new leaves per shoot developed in alpine and early-thawing subnival individuals and 12 (+/-1.2 SE) leaves in late-thawing subnival individuals. New leaf primordia emerged continuously from snowmelt until late autumn, even when plants were temporarily covered with snow. Differences in the developmental dynamics between the alpine and subnival population were independent of site temperatures, and are probably the result of ecotypic adaptation to differences in growing season length.     

Flowering Habit and Vegetative Behaviour in Tea (Camellia sinensis L) Seed Trees in North-East India

Apical growth of a tea shoot occurs by a succession of flushesseparated by short periods of rest. This paper describes theexternal morphology of flowering, fruiting, and abscission ofleaves of the tea plant in north-east India in relation to thephasic activity of shoot apices. All shoots on a tree make leafy growth when a new cycle of growthbegins in the spring, but terminal buds apparently become dormantas the season advances. Apparently dormant terminal buds shedbud scales, leaving on the stem a considerable number of scars,representing leafless cataphyllary flushes. These cataphyllaryflushes are produced at the same time as the leafy flushes onother shoots. A flower is formed only in the axil of a bud scale. Flowerswhich appear to develop in leaf axils are in fact inserted inthe axils of bud scales of the axillary buds. A distal leafy flush is without flowers. Flowers appear in itsleaf axils only when the terminal bud starts growth for thenext higher flush. A distal floriferous cataphyllary flush appearsas a terminal cluster of flowers. Thus, there is an acropetalsuccession of flowers, flush by flush on a caulome, determinedby the phasic activity of the apical bud. The main crop of flowers exposes anthers from the end of thethird flush (late September to early October) until the endof the winter period of growth (late January to early February).In some plants a second, minor crop of flowers appears in thespring between the end of the first and beginning of the secondflushes. In spite of considerable time lag between anthesis,the fruits produced by these two crops of flowers mature anddehisce at the same time during October to November. Abscission of leaves is also dependent upon the phasic activityof the apical buds. Only the top two flushes of a shoot possessleaves. Resumption of apical growth for a third flush, leafyor cataphyllary, causes the abscission of leaves on the lowermostof the three flushes. Two cataphyllary flushes therefore resultin the loss of all leaves on a shoot.     

Development of shoot inHydrangea macrophylla II. sequence and timing

The development of axillary buds, terminal buds, and the shoots extended from them was studied inHydrangea macrophylla. The upper and lower parts in a nonflower-bearing shoot are discernible; the preformed part of a shoot develops into the lower part and the neoformed part into the upper part (Zhou and Hare, 1988). These two part are formed by the different degrees of internode elongation at early and late phases during a growth season, respectively. Leaf pairs in the neoformed part of the shoot are initiated successively with a plastochron of 5–20 days after the bud burst in spring. The upper axillary buds are initiated at approximately the same intervals as those of leaf pairs, but 10–30 days later than their subtending leaves. Changes in numbers of leaf pairs and in lengths of successive axillary buds show a pattern similar to the changes in internode lengths of the shoot at the mature stage. The uppermost axillary buds of the flower-bearing shoot often begin extending into new lateral shoots when the flowering phase has ended. The secondary buds in terminal and lower axillary buds are initiated and developed in succession during the late phase of the growth season. Internode elongation seems to be important in determining the degrees of development of the axillary buds. Pattern of shoot elongation is suggested to be relatively primitive. Significances of apical dominance and environmental conditions to shoot development are discussed.     

Influence of intra-tree variation in time of budburst of white spruce on herbivory and the behaviour and survivorship of Zeiraphera canadensis

Abstract.

• 1 Field studies were carried out to determine the effects of intra-tree variation in the time of budburst of white spruce, Picea glauca Moench (Voss.), on the behaviour and survivorship of, and herbivory by, the spruce bud moth, Zeiraphera canadensis Mutt. & Free.
• 2 There was significant variation in the time of budburst among whorls, shoots and buds. Budburst was acropetal, with buds in the interior of the lower crown bursting first and terminal buds on terminal shoots in the upper crown bursting last.
• 3 Bud moths laid the greatest proportion of their eggs in the middle of the crown and egg hatch was usually best synchronized to budburst in this region. Many eggs hatched before terminal buds on terminal shoots in the upper crown had burst and thus intra-tree variation in budburst decreased the probability that first-instar larvae would colonize the most important plant parts for growth.
• 4 However, many later instars dispersed upwards and outwards in the crown and colonized the late bursting buds in the upper crown. Such dispersal reduces the effectiveness of intra-plant variation in budburst to reduce herbivory and permits Z.canadensis to eat young nutritious buds for a longer period of time.
• 5 Intra-tree variance in the date of budburst was greater than that between trees but there were no consistent differences between the intra-tree variance of trees in half-sib families with high or low susceptibility to Z.canadensis. Differences between trees in herbivory, bud moth density and survivorship were not related to the amount of intra-tree variation in budburst.

     

Studies of Growth and Flowering in Picea sitchensis (Bong.) Carr. 2. Initiation and Development of Male, Female and Vegetative Buds

Vegetative shoots from the base of the crown, and from partsof the tree likely to form male or female buds, were collectedfrom 40–years–old trees of Picea sitchensis (Bong.)Carr. throughout the 1973–4 annual growth cycle. The morphologyand growth rates of the terminal buds on these shoots were assessed. Bud scale primordia were formed most quickly in the female position,at an intermediate rate in the male position and most slowlyin the basal vegetative position during April, May and June.In July and early August the apical meristems swelled to formdomes and continued to grow at the same relative rates in themale, female and basal vegetative positions. Reproductive budswere first morphologically distinct in late August and sporangiaappeared in October. Dormancy, defined by the pause in apicalvolume increase, extended from mid-October to mid–March.Young strobili grew much faster than basal vegetative shootsof the same age between mid–March and bud burst in lateApril. Throughout the growth cycle, external changes in budsize reflected changes in size of the apical meristem, youngstrobihis or young vegetative shoot inside the bud. It is proposed that the rate of growth of an apical meristemmay be causally related to the type of bud which subsequentlydevelops from it. Sitka spruce, Picea sitchensis, bud development, morphology, growth of apical dome, flowering     

Consequences of the Elimination of Old Leaves upon Spring Phenological Events and the New Leaves Nutrient Concentration in a Wintergreen Woody Species in the Southern Hemisphere

Previous studies analyzed the importance of old leaves conservancy for wintergreen species plant growth only after early spring old leaves elimination. However, carbon and nutrient resources for growth could have already been translocated from old leaves to shoots during autumn. In this work, the effect of old leaves absence on the leaf mass per area (LMA, g m⁻²) and nutrient concentration of new spring leaves, shoot growth, and flowering was studied in Aristotelia chilensis, an Andean Patagonic woody wintergreen species of Argentina. Plants were studied after autumn defoliation (AD) or late winter defoliation (WD) and results were compared to those of undamaged control plants (CO). The new leaves LMA and mineral nutrient (N, P, K, and Mg) concentration values did not decrease in AD or WD compared to CO plants. Conversely, CO plants showed higher flowering intensity and shoot lengthening compared to AD or WD plants. There were not remarkable differences regarding the defoliation time, though non-flowering shoots grew in a lesser degree than the flowering shoots in WD plants. It was concluded that A. chilensis old leaves cohort is an important source to shoot growth and flowering but their absence does not affect the new leaves structure or nutritional status from early spring in either AD or in WD plants. New leaves formation probably is guaranteed by resources (carbon and nutrients) previously stored in stems or even in the buds containing the preformed leaves since March, by the end of summer. Provided the availability of complete resources for the new leaf flush independently of the old leaves A. chilensis would restore the carbon balance as soon as possible to resume the growth of heterotrophic tissues at normal rates. Endogenous response to counterbalance the old leaves absence on non-flowering shoots was more effective when there was greater lag time between defoliation and shoot growth resume. Flowering and non-flowering shoots compete for the available resources when A. chilensis have not yet expanded leaves and shoots supporting reproductive structures were stronger sinks compared to non-flowering shoots in WD plants.     

Effects of Didymella applanata and Botrytis cinerea on axillary buds, lateral shoots and yield of red raspberry

The viability of axillary buds and the growth and potential yield of lateral shoots at nodes of red raspberry (Rubus idaeus) infected naturally by Didymella applanata or Botrytis cinerea were measured on excised nodes, decapitated nursery canes or on canes from fruiting plantations. In comparison with lesion-free nodes, buds at infected nodes were smaller and fewer of them were capable of growth when excised and ‘forced’, although the difference in growth between infected and uninfected nodes decreased during late winter. After February, those buds at infected nodes which were capable of forced growth did so as early and with a similar growth rate as those at lesion-free nodes. In April, 70% of buds at infected nodes were capable of growth compared with 94% of those at lesion-free nodes. When nursery canes of cv. Mailing Delight were decapitated above infected nodes the emergence of lateral shoots from the terminal infected node did not differ significantly from that at lesion-free nodes. On a range of farm sites in Scotland the emergence of shoots at infected nodes in the cropping region of canes was significantly poorer than from uninfected nodes but substantially better at infected nodes of cv. Glen Clova than at those of cvs Mailing Jewel and Mailing Orion. It is suggested that cv. Glen Clova is relatively tolerant of spur blight and cane botrytis. The length and potential yield of laterals which developed at infected nodes in the cropping region of canes in these three cultivars did not differ significantly from those at lesion-free nodes. In all tests there was no significant difference in growth at nodes infected by D. applanata and B. cinerea which may indicate a common mechanism for suppression of buds.     

THE FATE OF THE DWARF SHOOT APEX IN BRISTLECONE PINE (PINUS LONGAEVA)

The fate of the pine dwarf shoot (DS) apex after needle initiation has been controversial. Dwarf shoot primordia of Pinus longaeva were examined to determine the developmental basis for DS with and without interfoliar buds. Interfoliar buds are microscopic buds derived from the original terminal apex of the DS. In October, all the DS primordia are similar in size and appearance. However, as the needles elongate in the following June the apices of more proximal DS decrease in size, such that by July there is a clear diminishing size gradient of apical domes in going from the most distal to the most proximal positions. The distal DSs start to form bud scales in July and have fully formed interfoliar buds by mid-August. In contrast, those DS apices lacking protective bud scales at needle maturity become suberized and can never proliferate into long shoots. The distal placement of interfoliar buds may be due to a group effect, where each developing DS inhibits the more proximal DSs in the long shoot terminal bud.     

Overwintering of Sphaerotheca mors-uvae on black currant and gooseberry

The ability of Sphaerotheca mors-uvae to perennate as cleistocarps, and as mycelium in buds was examined during the winters of 1965-6, 1966-7 and 1967-8 in relation to its two principal hosts, gooseberry and black currant. Cleistocarps on black currant leaves were examined from August 1965 to April 1966 and from July 1966 to March 1967. In 1965 cleistocarps were first observed on the leaves on 5 August; in 1966 on 11 July. These continued to develop through August and September and by October approximately 70% contained well-defined ascospores. The ascospore content remained generally at this level until February 1966 and November 1966; then the numbers of cleistocarps with ascospores fell and by April 1966 and March 1967 few such cleistocarps remained. From 21 March 1966 and 15 February 1967, but not otherwise, discharge of ascospores from the overwintered cleistocarps was readily obtained in laboratory tests. The viability and infectivity of the ascospores was demonstrated by allowing them to discharge on to leaf discs of black currant in the laboratory and also on to leaf discs and plants in the field. Sporulating colonies of S. mors-uvae developed within 8 days. Cleistocarps from shoots of black currant were examined from 4 August 1966 to 9 March 1967, and from 27 July 1967 to 1 January 1968. They developed in a similar manner to those on black currant leaves and by September in both 1966 and 1967 over 60% contained ascospores. This level was not maintained; the number of cleistocarps with ascospores fell gradually and by 8 December 1966 and 1 January 1968 few remained. Only in one laboratory test (21 November 1967) were ascospores discharged from a sample of these cleistocarps. Cleistocarps from shoots of gooseberry were examined from July 1966 to March 1967, and from August 1967 to January 1968. The pattern of ascospore development and subsequent decline in number of cleistocarps with ascospores was similar to that observed for black currant shoots. No discharge of ascospores could be demonstrated in laboratory tests. Evidence that S. mors-uvae perennates in buds of gooseberry was obtained by dissecting buds and by inducing buds on surface-sterilized shoots to burst under conditions which precluded chance infection. Field observations also suggested that bud infection occurred on gooseberry. Similar experiments failed to demonstrate the fungus in buds of black currant, and there was no indication of bud infection of this host in the field.     

Growth forms of Leymus chinesis (Poaceae) at the different developmental stages of the natural population

The manner in which the density of Leymus chinensis increases from a single plant to a dominant population can be understood by tracing the development of a population from early to late stages. Parent shoot density, above‐ground dry weight, spike density, heading rate and spike dry weight, density of spreading shoots (buds/daughter shoots in apical/axillary rhizomes) and clumping shoots (buds/daughter shoots in axillary parent shoots), and young rhizome length and weight were investigated in the same quadrats for a low density/early stage (LE) population and a high density/late stage (HL) population. Clonal growth (buds/daughter shoots formation) and sexual reproduction (spikes formation) increased while rhizome storage (young rhizome weight) decreased during the transition from LE to HL. In a LE population an outward occupation strategy was employed, with a high proportion of spreading shoots. As the population density gradually increased until HL, an inward consolidation strategy increasing shoot amount in previously occupied areas, was adopted. This was characterized by a high proportion of clumping shoots. Interestingly, the trade‐off between spreading and clumping shoots can be adjusted by the duration of young rhizome elongation during a growth season. In other words, compared with a HL population, a LE population shortened the duration of young rhizome elongation during the growth season, which resulted in more time for the production of axillary spreading shoots along the rhizomes, and high amounts and proportions of total spreading shoots. The special growth patterns, that is, trade‐offs among growth forms, allow L. chinensis to establish dominant populations throughout the eastern Eurasian Steppe.     

羊草克隆种群的螺旋扩张规律

羊草是欧亚大陆草原东部区域的一个优势物种。其优势地位的取得是羊草种群年复一年不断克隆复制过程中进行螺旋扩张的结果,其中,每一个螺旋圆环代表一个生长季,进而每个生长季可再分为不同的种群成长阶段,执行不同的生存发展策略:5月下旬—7月下旬为成熟扩张期,羊草的成熟母株通过水平伸长根茎执行扩张策略;7月下旬—10月下旬为幼苗占领期,通过地下芽输出为地上子株执行占领策略;10月下旬—来年3月下旬为羊草的休眠期;3月下旬—4月下旬为它的生长准备期;4月下旬—5月下旬为生长巩固期,子株发育为成熟的母株,完成了领地的彻底控制,执行的是巩固策略。羊草的游击型分株和密集型分株源自于共同的母株,二者位置相互分开,不但可以避免内部竞争,而且通过根茎之间的连接进行资源共享,从而在对外竞争中始终处于优势地位,这是羊草的克隆区隔避险策略;羊草种群通过产生遗传异质性的种子来避免因外界环境条件巨变所造成的整体灭绝,执行的是一种有性繁殖避险策略。羊草在既定策略基础上也表现出明显的二八规律,即通过80%比例的密集型分株进行旧领地的巩固,以20%比例的游击型分株进行新领地的扩张。总之,羊草通过扩张,占领,巩固,再扩张,再占领,再巩固,这样的年复一年的螺旋扩张,成为了当地的优势物种。其研究结果能为提高羊草生产力和种子产量及恢复受损草原生态系统提供十分重要的科学支持。     

Shoot morphogenesis associated with flowering in Populus deltoides (Salicaceae)

Temporal and spatial formation and differentiation of axillary buds in developing shoots of mature eastern cottonwood (Populus deltoides) were investigated. Shoots sequentially initiate early vegetative, floral, and late vegetative buds. Associated with these buds is the formation of three distinct leaf types. In May of the first growing season, the first type begins forming in terminal buds and overwinters as relatively developed foliar structures. These leaves bear early vegetative buds in their axils. The second type forms late in the first growing season in terminal buds. These leaves form floral buds in their axils the second growing season. The floral bud meristems initiate scale leaves in April and begin forming floral meristems in the axils of the bracts in May. The floral meristems subsequently form floral organs by the end of the second growing season. The floral buds overwinter with floral organs, and anthesis occurs in the third growing season. The third type of leaf forms and develops entirely outside the terminal buds in the second growing season. These leaves bear the late vegetative buds in their axils. On the basis of these and other supporting data, we hypothesize a 3-yr flowering cycle as opposed to the traditional 2-yr cycle in eastern cottonwood.