Bud and crown architecture of white spruce and black spruce

Needle primordia in buds and branch lengths were assessed in the crown of a plantation-grown white spruce tree. There was a gradation in needle primordia in buds in branches within the crown. The largest number of primordia was in the terminal bud of the leading main stem shoot, with the number in first-order whorl lateral shoot terminal buds decreasing from whorl 1 to whorl 4, below which buds contained a similar small number of primordia (about one-third as many as in the terminal shoot). Previous year's shoot elongation followed a similar pattern (i.e., elongation of whorl branches was greater closer to the top of the tree and elongation in the fourth through ninth whorls was about one-third that of the main stem leader). Higher order branches within whorls had within-branch gradation in shoot elongation and number of needle primordia, with older branches having as few as 16–30 primordia in buds and 3–4 cm elongation for high-order branches on older main stem whorls. There were strong correlations between the number of primordia in branch terminal buds and branch length/diameter and bud length/diameter/volume. In both black spruce and white spruce, there were strong correlations of number of needle primordia in main stem leader terminal buds with number of needle primordia in terminal buds of first and second whorl leaders.     

“Lateral Control”: Phytohormone Relations in the Conifer Treetop and the Short- and Long-Term Effects of Bud Excision in Abies nordmanniana

In a conifer tree, such as Nordmann fir, Abies nordmanniana Spach, the leader bud and its immediate surroundings play a decisive role in crown architecture. As subapical branch buds are segregated from the leader meristem, resource allocation between ortho- and plagiotropic growth is determined. The relationship between treetop buds in young trees was studied in the natural state and after surgical removal in early July of either the leader bud (decapitation) or the subapical whorl branch buds (destipitation). The two bud types showed consistent cytokinin profile differences but similar seasonal dynamics in cytokinins and auxin (IAA). After bud excision, ZRP increased dramatically in the subapical stem within 1 h, followed by ZR within 1 week. Supernormal levels of ZR were maintained through autumn and persisted in spring in the destipitated trees, but had returned to normal in the decapitated trees. The treetop buds remaining after bud excision experienced an immediate decrease in most cytokinins, followed, however, by a large surplus later in the season. The following spring this high level persisted in the leader bud of destipitated trees, but not in whorl buds of decapitated trees. Conspicuous growth pattern changes followed from destipitation, but few from decapitation. Growth reactions suggest that resource allocation to main branch buds inhibits leader growth in normal trees, a kind of “lateral control.” Auxin and ABA content in buds and stems was largely unaffected by treatments. Data suggest that subapical leader tissues beneath the apical bud group are a primary source of cytokinin regulation.     

DETERMINATE BRANCH DEVELOPMENT IN ALSTONIA SCHOLARIS (APOCYNACEAE): THE PLAGIOTROPIC MODULE

The development of the vegetative lateral branches in Alstonia scholaris (L.) R. Br. was examined. The overall architecture conforms to Prévost's model and the branches are sympodial complexes of plagiotropic modules. Each module consists of two whorls of 6–11 foliage leaves and a whorl of four scale leaves. The apex is parenchymatized just distal to the scale leaves. Renewal branches grow from buds in the axils of the scale leaves. These large buds are initiated simultaneously with the scale leaves and “use up” a large portion of the original apex. Parenchymatization of the central region of the apex occurs after a period of lateral growth and development that separates and vascularizes the renewal branches. Branch extension occurs sympodially by substitution. More typical buds develop in the axils of the foliage leaves but grow out only in response to injury or pruning. They are smaller than the renewal buds and, unlike them, are delimited by a shell zone during early development.     

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.     

Suppression of growth and death of meristematic tissues in <Emphasis Type="Italic">Abies sachalinensis</Emphasis> under strong shading: comparisons between the terminal bud,the terminally lateral bud and the stem cambium

The suppression of apical growth and radial trunk growth in trees under shade is a key factor in the competition mechanism among individuals in natural and artificial forests. However, the timing of apical and radial growth suppression after shading and the physiological processes involved have not been evaluated precisely. Twenty-one Abies sachalinensis seedlings of 5-years-old were shaded artificially under a relative light intensity of 5% for 70 days from August 1, and the histological changes of the terminal bud and terminally lateral bud of terminal leader and the cambial zone of the trunk base were analyzed periodically. In shade-grown trees, cell death of the leaf primordia in a terminal bud of terminal leader was observed in one of the three samples after 56 and 70 days of shading, whereas the leaf primordia in a terminal bud of terminal leader in all open-grown trees survived until the end of the experiment. In addition, the leaf primordia of the terminally lateral buds of terminal leader retained their cell nuclei until the end of the experiment. No histological changes were observed in the cambial cells after shading, but the shade-grown trees had less cambial activity than the open-grown trees through the experiment. Strong shading appeared to inhibit the formation and survival of cells in the terminal bud of terminal leader rather than the terminally lateral buds of terminal leader and the cambium. The suppression of the terminal bud growth and elongation of the surviving lateral buds would result in an umbrella-shaped crown under shade.     

The timing of sprays for the protection of terminal buds on apple shoots from powdery mildew

Post-blossom sprays of fungicides, repeated at 10-day intervals until leader (syn. extension) shoots had stopped producing new leaves, provided the best protection of terminal buds against Podosphaera leucotricha on the apple cv. Lane's Prince Albert. Spraying was most effective in early summer, although many of these buds were not invaded until later, when the rate of shoot growth declined; applications from July to September did not compensate for the enhanced infection which followed interruptions of the post-blossom programme between late May and early July. This early period was critical because most leaf infections occurred then, and because this phase of the epidemic on foliage determined the eventual intensity of mildew on terminal leaves, and hence the inoculum available for infecting terminal buds. Also, many lateral shoots ceased growth early and their apices were directly protected by sprays applied in June. Applications after early June were too late to protect newly formed fruit buds on spur branches.     

Effect of Gibberellic Acid, 2,3,5-Triiodobenzoic Acid and Indole Acetic Acid on Growth and Development of Impatiens balsamina during Different Photoperiods

This paper deals with the effect of 100 mg/1 each of GA₃ TIBA and IAA singly and in combination with each other on stem elongation, development of lateral branches and floral bud initiation in Impatiens balsamina plants exposed to 8-, 16- and 24-h photoperiods. GA₃ enhances stem elongation, the enhancing effect decreasing with IAA as well as with TIBA during 8-h but increasing during 16- and 24-h photoperiods. It decreases the number of lateral branches, the decrease being greatest during 16-, less during 8- and the least during 24-h photoperiods. The time taken for floral buds to initiate with and length of branches during 16-h photoperiods. During 8-h photoperiods, IAA delays the initiation of floral buds, while GA₃ hastens it when used together with TIBA or IAA or both. GA₃ increases the number of floral buds on the main axis but decreases it on lateral branches, while TIBA decreases the number on the main axis but increases it on lateral branches. IAA reduces the number of floral buds on the main axis only when used alone, but on both the main axis as well as on lateral branches when used together with GA₃ and TIBA. Floral buds were not produced on lateral branches when plants were treated with GA₃, TIBA and IAA all together. GA₃ and TIBA induced floral buds even under non-inductive photoperiods, the number of buds and reproductive nodes being less in TIBA- than in GA₃-treated plants during 24-h photoperiods. The time taken for floral buds to initiate with GA₃ and TIBA during noninductive photoperiods is much longer than that during 8-h inductive photoperiods with or without GA₃ or TIBA application. IAA completely inhibits the GA₃- and TIBA-caused induction during 24-h, but only delays it and reduces the number of reproductive nodes and floral buds during 16-h photoperiods.     

The possible roles of nutrient deprivation and auxin repression in apical control

Apical control is the suppression of growth in lower branches by a higher dominant branch or leader shoot. We investigated possible mechanisms involved in this developmental response in three widely diverse species (Japanese morning glory, Ipomoea nil, hybrid poplar, Populus trichocarpa, × P. deltoides, and Douglas-fir, Pseudotsuga menziesii). The following two hypotheses were tested: (1) the mineral nutrient-deprivation hypothesis, which is that the continued growth of the lower branches is repressed by the diversion of nutrients to the upper dominating branch or shoot, and (2) the auxin-repression hypothesis, which is that auxin produced in the upper dominating branch or shoot moves down to the lower branches where continued growth is repressed. The results of experiments involving the manipulation of available nutrients by dominant branch removal and fertilization were consistent with the first hypothesis for morning glory, poplar, and for second- or third flushing of lateral branches in Douglas-fir. The results of the experiments involving auxin (NAA, 1-naphthalene acetic acid) replacement treatments on decapitated shoots bearing growing lateral branches were inconsistent with the second hypothesis in morning glory, poplar and in first-flushing Douglas-fir. However, despite concerns about possible NAA toxic effects, there was evidence of auxin repression of second flushing in Douglas-fir. Overall, the data supported a significant role for nutrient availability but not for auxin repression in apical control of morning glory and poplar. In Douglas-fir, apical control in first-flushing lateral branches from over-wintered buds was largely insensitive to both nutrient availability and auxin repression; however, second flushing was sensitive to both.     

The Effect of Bud Position on Branch Growth and Bud Abscission in Quercus petraea (Matt.) Liebl.

The time at which a bud began to expand was related to its positionnot only on an individual shoot but also within the crown. Thedistribution of buds and branches on the shoot was uneven; theshoot tip, where they were densely clustered, was termed the‘whorl; and the remainder of the shoot, where they werewidely spaced, the ‘interwhorl’ stem. In spring,the terminal bud started expanding before the ‘whorl’buds which preceded the ‘interwhorl’ stem buds;completion of the flush of growth, determined by the end ofleaf expansion, occurred in the reverse order, ‘interwhorl’> ‘whorl’ > terminal. Similarly bud expansionstarted at the top of the crown and progressed downwards, andthe first shoots to complete their flush were at the bottomof the crown. Approximately 60% of the buds on each shoot beganexpanding in spring but only about half of these formed branches.Bud abscission began in May and by Sep. 45% of buds originallypresent had abscised. Most of-the buds that did not abscisewere the small buds at the base of the shoot that were not originallyassociated with a leaf. Approximately 42% of ‘whorl’buds and 28% of MnterwhorP stem buds formed branches. ‘Whorl’branches were approx. 60% longer that ‘interwhorl’stem branches; buds on the lower surface of the shoot producedlonger branches than those on the upper surface. The implicationsof the results for the development of crown form and selectionof superior oak are discussed. Quercus petraea, oak, buds, branches, crown form     

A developmentally based categorization of branching in Trifolium repens L.: influence of nodal roots

This study describes the successive stages of development of branches from axillary buds in fully rooted plants of Trifolium repens grown in near optimal conditions, and the way in which this developmental pathway differs when nodal root formation is prevented as plants grow out from a rooted base. Cuttings of a single genotype were established in a glasshouse with nodal root systems on the two basal phytomers and grown on so that nodal rooting was either permitted (+R) or prevented (-R). In +R plants, axillary tissues could be assigned to one of four developmental categories: unemerged buds, emerged buds, unbranched lateral branches or secondarily branched lateral branches. In -R plants, branch development was retarded, with the retardation becoming increasingly pronounced as the number of -R phytomers on the primary stolon increased. Retarded elongation of the internodes of lateral shoots on -R plants resulted in the formation of a distinct fifth developmental category: short shoots (defined as branches with two or more leaves but with mean internode length equal to, or less than, 10% of that of the immediately proximal internode on the parent stolon) which had reduced phytomer appearance rates but retained the potential to develop into lateral branches. Transfer of +R plants to -R conditions, and vice versa, after 66 d demonstrated that subsequent branch development was wholly under the control of the youngest nodal root present, regardless of the age and number of root systems proximal to it.     

Branching in Polysphondylium whorls: Two-dimensional patterning in a three-dimensional system

During morphogenesis in the slime mold Polysphondylium pallidum, spherical masses of cells called whorls pinch off from the slug at regular intervals. Soon afterward, branches form at equidistant positions around the whorl equator. We have quantified the relationship between the number of cells in a whorl, and the number of branches that form. We find that the number of branches produced is proportional to the surface area of the whorl, suggesting that the patterning process is confined to the whorl surface. This observation is consistent with theoretical arguments that mechanisms for pattern formation would likely operate in one or two dimensions, not three.     

APICAL DOMINANCE AND FORM IN WOODY PLANTS: A REAPPRAISAL

The form of woody plants is commonly interpreted in terms of apical dominance. Trees with the decurrent or deliquescent branching habit are said to have weak apical dominance, whereas excurrent branching is associated with strong apical dominance. A close examination of many decurrent species such as the oaks, hickories, and maples reveals that almost all of the lateral buds on the current year's twigs are completely inhibited. This complete inhibition of lateral buds by definition and common usage of the term is an expression of strong apical dominance. In trees possessing the excurrent branching habit, such as most conifers and some angiosperms, many of the lateral buds on the current year's twigs elongate to varying degrees. This is usually interpreted as an expression of weak apical dominance. The relationship between bud inhibition and form in woody perennials is much more complex than bud inhibition in herbaceous plants because of the time sequence in the formation and release of lateral buds. For example, it is only after a period of rest or dormancy in the decurrent forms that one or more of the uppermost lateral buds are released, and these may outgrow the currently elongating terminal shoot resulting in forking. Conversely, in the excurrent forms, it seems that the initial expression of weak apical dominance enables the terminal leader to outgrow the currently elongating lateral branches so that it exerts complete control over their subsequent growth and development in later years. An examination of the levels of diffusible auxin at different points along the twigs of excurrent and decurrent species indicates that the balance of growth factors at any given locus, and not the absolute quantity of auxin, exerts primary control over bud inhibition and shoot elongation.     

Relationships among shoot sinks for resources exported from nodal roots regulate branch development of distal non-rooted portions of Trifolium repens L

Two manipulative experiments tested hypotheses pertaining to the correlative control exerted by nodal roots on branch development of the distal non-rooted portion of Trifolium repens growing clonally under near-optimal conditions. The two experiments, differing in their pattern of excision to manipulate the number of branches formed at the first 9-10 phytomers distal to the youngest nodal root, each found that after 20 phytomers of growth the total number of lateral branches formed on the primary stolon remained between five and seven regardless of where the branches formed along the stolon. Additional treatments established that nodal roots influenced branch development via relationships among shoot sinks for the root-supplied resources rather than through variation in the supply of such resources induced by fluctuations in photosynthate supply to roots from branches. Regression analysis of data pooled from treatments of both experiments confirmed that shoot-sink relationships for root- supplied resources controlled the branching processes on the non-rooted portion of plants. A disbudding treatment, which removed all the apical and axillary buds present on basal branches, but left other branch tissues intact, increased branch development of the apical region in the same way as did complete excision of the basal lateral branches. The apical buds and the elongation processes occurring immediately proximal to the buds were thus identified as strong sinks for the root-supplied resources. Such results suggest that branch development on the non-rooted shoot portion distal to the youngest nodal root is regulated by competition among sinks for root-derived resources, of limited availability, necessary for the processes of elongation of axillary buds and the primary stolon apical bud.     

四川大头茶的分枝率和顶芽动态

以四川大头茶植株的枝和顶芽为基本构件单元,对不同年龄级成熟植株的分枝率和枯芽数动态进行了研究。结果表明：四川大头茶成株任一种分枝率在种群内不同龄级植析间以及在同一植株内不同发育位置的主枝间无显著变化;而同一主枝内各类型分枝率并非按相同的几何及数增减,即枝内各类分枝率间具显著的差异。     

Overexpression of the maize Teosinte Branched1 gene in wheat suppresses tiller development

The number of viable shoots influences the overall architecture and productivity of wheat (Triticum aestivum L.). The development of lateral branches, or tillers, largely determines the resultant canopy. Tillers develop from the outgrowth of axillary buds, which form in leaf axils at the crown of the plant. Tiller number can be reduced if axillary buds are not formed or if the outgrowth of these buds is restricted. The teosinte branched1 (tb1) gene in maize, and homologs in rice and Arabidopsis, genetically regulate vegetative branching. In maize, increased expression of the tb1 gene restricts the outgrowth of axillary buds into lateral branches. In this study, the maize tb1 gene was introduced through transformation into the wheat cultivar "Bobwhite" to determine the effect of tb1 overexpression on wheat shoot architecture. Examination of multiple generations of plants reveals that tb1 overexpression in wheat results in reduced tiller and spike number. In addition, the number of spikelets on the spike and leaf number were significantly greater in tb1-expressing plants, and the height of these plants was also reduced. These data reveal that the function of the tb1 gene and genetic regulation of lateral branching via the tb1 mode of action is conserved between wheat, rice, maize and Arabidopsis. Thus, the tb1 gene can be used to alter plant architecture in agriculturally important crops like wheat.     

青桃利用单、复花芽结果对盛果期生长结实的影响

青桃盛果期,特别中庸偏旺的植株,单、复花芽果枝数大体均等,复花芽果枝以其复芽个体较大,枝条较粗而往往表现为优势。然而笔者于1974—1979年观察了青桃树以单、复花芽结果对盛果初期的影响,结果表明:利用单花芽结果,控制了强旺枝和结果部位,使果枝分布均匀,树势稳定,尤其果枝数、花芽数、单株产量等的变动趋势与年度增加无关,r为:-0.2899、0.0620(单、复花芽果枝数分别占总果枝数的35.1％和16.7％),-0.3830、-0.1804(单、复花芽朵数分别占总花芽朵数的44.5％和13.4％),0.4093(单株年产量为179.4公斤,从而延长了盛果期;利用复花芽结果却相反,促进树势极性生长,旺枝增强,果枝外移,内膛空虚,尤其果枝数、花芽数、单株产量等的变动趋势随年度增加而呈显著与极显著负相关,r为:-0.8938~(**)、-0.9818~(**)(单、复花芽果枝数分别占总果枝数的27.1％和21.1％),-0.953~(**)、-0.8908~(**)(单、复花芽朵数占总花芽朵数的35.3％和6.8％).-0.8853~(**)(单株产量较单花芽结果减产19.4％),从而明显缩短盛果期。     

Bud Content and its Relation to Shoot Size and Structure in Nothofagus pumilio(Poepp. et Endl.) Krasser (Nothofagaceae)

Buds of shoots from the trunk, main branches, secondary branchesand short branches of 10–21 year-old Nothofagus pumiliotrees were dissected and their contents recorded. The numberof differentiated nodes in buds was compared with the numberof nodes of sibling shoots developed at equivalent positionsduring the following growing season. Axillary buds generallyhad four cataphylls, irrespective of bud position in the tree,whereas terminal buds had up to two cataphylls. There were morenodes in terminal buds, and the most distal axillary buds, oftrunk shoots than in more proximal buds of trunk shoots, andin all buds of shoots at all other positions. The highest numberof nodes in the embryonic shoot of a bud varied between 15 and20. All shoots had proximal lateral buds containing an embryonicshoot with seven nodes, four with cataphylls and three withgreen leaf primordia. The largest trunk, and main branch, shootswere made up of a preformed portion and a neoformed portion;all other shoots were entirely preformed. In N. pumilio, theacropetally-increasing size of the sibling shoots derived froma particular parent shoot resulted from differences in: (1)the number of differentiated organs in the buds; (2) the probabilityof differentiation of additional organs during sibling shootextension; (3) sibling shoot length; (4) sibling shoot diameter;and (5) the death of the apex and the most distal leaves ofeach sibling shoot. Copyright 2000 Annals of Botany Company Axis differentiation, branching, bud structure, leaf primordia, neoformation, Nothofagus pumilio, preformation, size gradient     

Photoperiodic studies on growth and development of Impatiens balsamina L.

Summary Seedlings of Impatiens balsamina raised under ND and LD conditions were divided into two sub-groups each when they had reached 5-leaf stage. While one sub-group was left under the same condition (NDND or LDLD), the other was transferred to the other photoperiod (NDLD or LDND). NDND plants were subdivided into 2 lots. One of these was transferred to SD in May. The dates of emergence of individual branches and floral buds were recorded and the vegetative period was calculated in each case.It was found that in NDND plants floral buds were produced from all the nodes except the lowermost which produced a single vegetative branch. In LDND plants the vegetative branches were produced from the lower 9 nodes but floral buds from those above these. Small leafy structures which ultimately dried up were produced from a few top nodes in both these cases. In contrast to this in LDLD plants only vegetative branches were produced from all the nodes. In NDLD plants floral buds were produced from the lower 3–5 nodes prior to transfer to LD condition, but vegetative branches were produced from the upper nodes after this transfer. Even some of the lower floral buds reverted to vegetative state under this condition.The production of floral buds or the vegetative branches as the case may be, occurred in acropetal succession under all the photoperiodic conditions and never in basipetal manner.LDLD and NDLD plants, which did not flower at all, continued to produce lateral branches without showing any sign of senescence, while LDND and NDND ones showed yellowing of the apical growing point which spread downwards and lead ultimately to the death of the plant. The senescence was hastened when these plants were transferred to SD condition towards the end of May. The senescence therefore, appears to be related with reproductive development. The results are discussed in the light of current literature.     

Crown architecture and dynamics in Abies procera as influenced by cutting for greenery

Greenery production from 13 to 17 years old Abies procera was studied in three localities differing in soil and tree provenance. Two alternative clipping strategies for greenery were applied during 8 years. Subsequently, destructive crown analyses were carried out on the trees sampled from these and traditionally cut plots. Regeneration occurred from stubs or bases of cut primary branches. In cases when stubs were removed, proliferation took place around the scar; these branches compensated in numbers but not in biomass for the absence of stub branches. More severe cutting regimes resulted in an overall biomass reduction, most notably with shorter and lighter whorl branches. Interwhorl branches and branches issued from the base of primary branches were able to extend progressively in response to severe cutting. Base branches were apparently recruited along most of the trunk, the latency of growth points thus being up to at least 10 years. Competition among interwhorl branches, as well as more complicated interactions between whorl branches and secondary branches arising from their base, are indicated by the data. Reiteration directly from the stem could not be provoked and the results suggest that optimum greenery yield depends on careful stub management.