Increase in lodging safety factor of thigmomorphogenically dwarfed shoots of mulberry tree

The lodging safety factor of mechanically stimulated shoots of mulberry tree ( Morus bombycis Koidz. cv. Kenmochi) was measured to test the hypothesis that thigmomorphogenic dwarfing increases mechanical stability of plants. The lodging safety factor, an indicator of mechanical stability of erect shoots, has been defined as the ratio of critical lodging load to leaf fresh weight. Two-year-old shoot cuttings of mulberry tree were rubbed, brushed, and shaken every day. During the experimental period, the lodging safety factor of the control shoots remained at about 5. while the safety factor of the thigmomorphogenically dwarfed ones was ca 1.6 times larger, which supports the hypothesis. The mechanical stimulation reduced the stem elongation rate, but did not greatly change the partitioning of dry matter between leaves and stem. Therefore, it is concluded that the reduction of stem elongation rate was the main cause of the increase in the lodging safety factor of the stimulated shoots.     

Ethylene Evolution Stimulated by Succimc Acid-2,2-Dimethylhydrazide in Apple Shoots

The rate of ethylene production in the shoot tips of the apple cultivar ‘Ingrid Marie’ (Red motant) after succinic acid-2,2-dimethylhydrazide (SADH) treatment was investigated. The shoots were treated with SADH at 12.5, 25 or 50 mM in the months of June, July and August. The samples were analysed 7,15, 30 and 60 days after SADH spray. Ethylene production was considerably higher in the SADH-treated shoots than in the untreated ones. Application of 12.5 or 25mMSADH late in June stimulated ethylene production 3-fold, while 50 mM produced a 5-fold increase in ethylene production 7 days after spraying. There was a continuous decrease in ethylene production at the subsequent samplings. In untreated control shoots there was a slight increase. However, even 60 days after spraying the shoots treated with 50 mM had higher ethylene production than the control shoots. The same relationship was found in the shoots treated in the months of July and August. A parallelism has been established between the rate of ethylene production and growth retardation, A residual effect of SADH treatment in earlier years was demonstrated and is discussed.     

THE LAW CONTROLLING THE QUANTITY OF REGENERATION IN THE STEM OF BRYOPHYLLUM CALYCINUM

1. A method is given which allows us to measure the influence of the mass of a leaf upon the quantity of shoots regenerated in an isolated piece of stem. This method consists in isolating a piece of stem with only two leaves left at the basal node and then splitting the stem lengthwise so that each half has one basal leaf. By leaving one leaf intact while the size of the sister leaf is reduced, the influence of the mass of the leaf upon the quantity of shoots regenerated by the stem can be measured. 2. This method has yielded the result that the mass of shoots regenerated at the apex of such a piece of stem increases under equal conditions and in equal time with the mass of the leaf, and is approximately proportional to the mass of the leaf. 3. Such an influence of the mass of the leaf upon the mass of shoots produced by the stem is only intelligible on the assumption that the growth of the regenerating shoot occurs at the expense of material furnished by the basal leaf. 4. This assumption is supported by two facts: first, that in the dark this influence of the leaf disappears more or less completely; and, second, that a leaf attached to the base of a regenerating stem after some time weighs markedly less than does a sister leaf completely detached from the stem, but otherwise under equal conditions. 5. This latter fact that a leaf when attached to the base of an excised piece of stem wilts more rapidly than when completely isolated is the reason that the proportionality between mass of a basal leaf and mass of shoot regenerated at the apex of an isolated piece of stem cannot always be demonstrated with the same degree of accuracy as the proportionality between the mass of completely isolated leaves and the mass of shoots they produce. 6. The material furnished by the leaf to the stem is not restricted to water but includes also the solutes, since not only the fresh weight but also the dry weight of the shoot regenerated by a piece of stem increases with the mass of the leaf attached to the base of the stem; and since not only the water contents but the dry weight of a leaf attached to the base of an excised piece of stem diminish when compared with the dry weight of a completely detached sister leaf. 7. The mass of shoots produced by an isolated piece of stem without leaf is small and almost negligible compared with the mass of shoots produced by the same piece of stem when a leaf of sufficient mass is attached to the base of the stem.     

Modulated Degradation of Ribulose Bisphosphate Carboxylase in Leaves on Top-Pruned Shoots of the Mulberry Tree (Morus alba L.)

Yamashita, T. 1987. Modulated degradation of ribulose ftisphosphatecarboxylase in leaves on top-pruned shoots of the mulberry tree(Morus alba L.).—J. exp. Bot. 38: 1957–1964. The effects of pruning shoot tops on the synthesis and degradationof ribulose 1,5–Wsphosphate carboxylase (RuBPCase) inleaves on remaining shoots were investigated in mulberry trees.Leucine labelled with ¹⁴C was fed to leaf discs from field-grownmulberry trees and ¹⁴C incorporation into RuBPCase was examined.Proportion of ¹⁴C in RuBPCase to leucine–¹⁴C absorbedby leaf discs was remarkably lowered by top-pruning, thoughoccasionally a slight increase was observed soon after pruning.Yet RuBPCase content in leaves on top-pruned shoots became progressivelyhigher than that in leaves on intact shoots. Changes in ¹⁴Cin Ru1BPCase in leaves of mulberry saplings previously fed ¹⁴CO₂were followed. Following ¹⁴CO₂ feeding, the attainment of themaximal level of ₁₄C in RuBPCase was retarded by top-pruning.The highest level of ₁₄C in RuBPCase was maintained in leaveson top-pruned shoots but decreased in leaves on intact shoots.Specific radioactivity in RuBPCase continued to increase inleaves on top-pruned shoots even after attaining a maximum levelin the control leaves. These facts suggest that the increasein RuBPCase by top-pruning results from a cessation of its degradationfor the remobilization of nitrogen for newly developing leaveson shoot tops. Key words: RuBP carboxylase, shoot pruning, mulberry (Morus alba)     

Size structure of current-year shoots in mature crowns

Characteristics of current-year shoot populations were examined for three mature trees of each of three deciduous broad-leaved species. For first-order branches (branches emerging from the vertical trunk) of the trees examined, lengths or diameters of all current-year shoots were measured. Total leaf mass and total current-year stem mass of first-order branches were estimated using an allometric relationship between leaf or stem mass and length or diameter of current-year stems. For each tree, the number of current-year shoots on a first-order branch was proportional to the basal stem cross-sectional area of the branch. On the other hand, first-order branches had shoot populations with size structures similar to each other. As a result, the leaf mass of a first-order branch was proportional to the basal stem cross-sectional area of the branch, being compatible with the pipe-model relationship. All current-year shoot populations had positively skewed size structures. Because small shoots have a larger ratio of leaf mass to stem mass than large shoots, first-order branches had an extremely large ratio of leaf mass to current-year stem mass. This biased mass allocation will reduce costs for current stem production, respiration and future radial growth, and is beneficial to mature trees with a huge accumulation of non- photosynthetic organs. The allometric relationships between leaf mass and basal stem diameter and that between leaf mass and current-year stem mass of first-order branches were each similar across the trees examined. Characteristics of shoot populations tended to offset inter-species diversity of shoot allometry so that branch allometry shows inter-species convergence.     

Changes in crown development patterns and current-year shoot structure with light environment and tree height in Fagus crenata (Fagaceae)

The relative effects of light and tree height on the architecture of leader crowns (i.e., the leading section of the main trunk, 100 cm in length) and current-year shoots for a canopy species, Fagus crenata, occupying both the ridge top and the valley bottom in a cool-temperate forest in Japan were investigated. For leader crowns, the number of current-year shoots and leaves increased with increasing tree height, whereas the mean length of current-year shoots increased with increasing relative photon flux density (PFD). The leader crown area decreased, and the depth and leaf area index of leader crowns increased, with increasing relative PFD. The mass of current-year shoots increased with relative PFD. However, this total mass was allocated differently between stems and leaves depending on tree height, such that the relative allocation to stems increased with increasing tree height. Furthermore, stem structures within current-year shoots also changed with height, such that taller trees produced thicker and shorter stems of the same volume. In contrast, leaf structure and leaf biomass allocations changed with relative PFD. Specific leaf area decreased with increasing relative PFD. In addition, leaf number increased more rapidly with increasing individual leaf mass for trees exposed to greater relative PFD. Consequently, the total leaf area supported by a stem of a given diameter decreased with increasing tree height and relative PFD. Thus, the architecture of leader crowns and current-year shoots were related differently to light and tree height, which are considered important for efficient light capture and the growth of small and tall trees in different environments.     

Current-year and subsequent-year effects of crop-load manipulation and epicormic-shoot removal on distribution of long, short and epicormic shoot growth in Prunus persica

BACKGROUND AND AIMS: The distribution of canopy growth among different shoot types such as epicormic, long and short shoots is not well understood in the peach tree. In this experiment, the effects of crop load and early epicormic sprout removal on current and subsequent-year distribution of vegetative growth among epicormic, long and short shoots was investigated in Prunus persica. METHODS: Field trials were conducted in Winters, California, in 2003-2004. Crop load was manipulated with fruit thinning in 2003 to produce trees that were de-fruited, commercially thinned or full crop, and half of the trees in each cropping treatment had all current year epicormic sprouts removed at the time of fruit thinning. Yield was recorded and trunk and root carbohydrates were sampled to confirm the effect of 2003 crop load differences on tissue carbohydrate concentration. All current-season vegetative-shoot extension growth was harvested from half of the trees in each treatment in the autumn of 2003 and from the other half in the autumn of 2004. Epicormic, long and short shoots were separately evaluated for dry weight, node number and leaf-stem parameters. KEY RESULTS: In 2003, long-shoot dry weight and node number were significantly affected by crop load; however, short-shoot dry weight and node number were not significantly affected. The 2003 crop-load treatments did not affect 2004 vegetative growth of any shoot type. Some re-growth of epicormic shoots followed early epicormic sprout removal: by the end of the 2003 season, trees in the early shoot-removal treatment had approximately one-third of the epicormic-shoot dry weight as unpruned trees. CONCLUSIONS: Fruit thinning promoted distribution of growth similar to that of de-fruited trees. While thinning was effective in increasing fruit size, it exacerbated the problem of epicormic sprouting. Early epicormic sprout removal did not stimulate the excessive epicormic re-growth in the same or subsequent year relative to previously studied summer pruning methods.     

Biomass Partitioning in Tomato Plants Infected with Meloidogyne incognita

Tomato plants were inoculated with Meloidogyne incognita at initial populations (Pi) of 0, 1, 10, 50, 100, and 200 (x 1,000) eggs per plant and maintained in a growth chamber for 40 days. Total fresh biomass (roots + shoots) at harvest was unchanged by nematode inoculation with Pi of 1 x 10⁵ eggs or less. Reductions in fresh shoot weight with increasing Pi coincided with increases in root weight. Total fresh biomass declined with Pi above 1 x 10⁵ eggs, whereas total dry biomass declined at Pi above 1 x 10⁴ eggs. The greatest reduction percentages in fresh shoot biomass induced by root-knot nematodes occurred in the stem tissue, followed by the petiole + rachis; the least weight loss occurred in the leaflets. Although biomass varied among shoot tissues, the relationship between biomass of various shoot tissues and Pi was described by quadratic equations. The linear and quadratic coefficients of the equations (stem, petiole + rachis, or leaflets on Pi) did not differ among tissues when calculations were based on standardized values. Meloidogyne incognita-infected plants had thinner leaves (leaf area/leaf weight) than did uninfected plants. Reductions in leaf weight and leaf area with nematode inoculation occurred at nodes 5-15 and 4, 6-14, respectively. Losses in plant height and mass due to nematodes reflected shorter internodes with less plant mass at each node.     

Interactive effects of mycorrhization and elevated carbon dioxide on growth of young pedunculate oak (Quercus robur L.) trees

Pedunculate oak (Quercus robur L.) was germinated and grown at ambient CO₂ level and 650 ppmv CO₂ in the presence and absence of the ectomycorrhizal fungus Laccaria laccata for a total of 6 month under nutrient non-limiting conditions. Mycorrhization and elevated atmospheric CO₂ each supported the growth of the trees. Stem height, stem diameter, and dry matter accumulation of pedunculate oak were increased by mycorrhization. Elevated atmospheric CO₂ enhanced stem height, stem diameter, fresh weight and dry weight, as well as lateral root formation of the trees. In combination, mycorrhization and elevated atmospheric CO₂ had a more than additive, positive effect on tree height and biomass accumulation, and further improved lateral root formation of the trees. From these findings it is suggested that the efficiency of the roots in supporting the growth of the shoot is increased in mycorrhized oak trees at elevated atmospheric CO₂.Abbreviations DW dry weight - FW fresh weight - RWC relative water content     

实生和嫁接槐树的枝梢生长与光合特性

测定槐树实生树和嫁接树叶片的光合光响应特性、比叶重和叶绿素含量及枝梢生长的结果表明,嫁接树枝梢基径和高度增长趋势与实生树一致,但嫁接树一次、二次梢基径和高度增长量均大于实生树。嫁接树的最大净光合速率、光饱和点、比叶重和单位重量的叶绿素含量均显著高于实生树,高的光合能力与其单位面积叶片重量增加和单位重量叶绿素含量增高有关。嫁接槐树的类胡萝卜素与叶绿素比值高于实生槐树,叶绿素a,b比值和单位重量类胡萝卜素含量显著高于实生槐树,表明其对强光的适应性强。     

Effect of host-plant manipulation by a gall-inducing insect on abundance of herbivores on chestnut trees

We investigated plant-mediated effects of the stem gall wasp, Dryocosmus kuriphilus Yasumatsu (Hymenoptera: Cynipidae), on other herbivores on the chestnut tree Castanea crenata. In the early season, leaves emerged earlier and in greater numbers on galled shoots than on ungalled shoots. On galled shoots the leaf to shoot biomass ratio was lower and the leaves were physically different. In May and June the concentration of nitrogen in leaves was higher on galled shoots than on ungalled shoots. In July, the water content of leaves was lower on galled shoots. In May and June, the number of aphids, Myzocallis kuricola Matsumura (Homoptera: Aphidoidea), on leaves was higher on galled shoots than on ungalled shoots, but the opposite was true at the end of July. Laboratory experiments showed that aphid fecundity and body weight decrease were higher in May and June when aphids fed on leaves on a galled shoot than when they fed on those on ungalled shoots. In contrast, aphid performance in July was greater on ungalled leaves than on galled leaves. Our findings suggest that gall initiation in a chestnut tree affected aphid performance by affecting host plant quality.     

Differences in Architecture and Shoot Growth during Stagnant and Extension Growth Phases of Acanthopanax sciadophylloides(Araliaceae)

The shoot growth of a deciduous tree, Acanthopanax sciadophylloidesFranch. et Savat. shows inter-annual intermittent repetitionof two distinctive phases, a stagnant growth phase (S-phase)and vigorous extension-growth (E-phase). To help understandthe differentiation mechanism, shoot development was studiedover time in both shoot phases. S-phase and E-phase shoots weredistinguished from each other by morphological traits: S-phaseshoots are characterized by higher allocation to leaves anda shorter period of stem growth, while E-phase shoots show continuousstem extension over the growing season. Specific leaf area didnot differ between the two phases. This shoot differentiationwas similar to the morphological differentiation of shoots betweenlong vs. short shoots found in some temperate trees. Leavesof both phases were well-dispersed through adjustment of petiolelength and leaf-blade size to reduce mutual shading within ashoot. Stem-wood density of current-year shoots was lower inE-phase compared with S-phase shoots. Leaves produced earlyin the season affected the growth phase of the following year.These results suggest that annual shoot differentiation of A.sciadophylloides was determined during the previous season andreflects leaf productivity in a given habitat during that growingseason. Copyright 2001 Annals of Botany Company Acanthopanax sciadophylloides, Araliaceae, biomass allocation, intermittent shoot growth, leaf display, shoot architecture, shoot differentiation     

Ontogenetic changes in the numbers of short- vs. long-shoots account for decreasing specific leaf area in Acer rubrum (Aceraceae) as trees increase in size

Specific leaf area (SLA) is reported to decrease with increasing plant size among dicot tree species despite a strong positive correlation between SLA and relative growth rate. This diminishing returns in SLA may result from changes in the relative numbers of different shoot types bearing leaves with different SLAs as trees increase in overall size. This ontogenetic shift hypothesis was examined for 15 Acer rubrum trees differing in basal stem diameter (0.01 m ≤ D ≤ 0.62 m). Detailed analyses of the largest tree showed that short-shoots produced leaves with significantly smaller SLA than the leaves produced by long-shoots regardless of the location of shoots within the canopy. A combination of random effect and split-plot (main-effect) ANOVA models showed that >94% of the variance observed for SLA was attributable to shoot type rather than to the location of leaves in the canopy. Further, with increasing trunk diameter, the number of short-shoots increased rapidly relative to the number of long-shoots. Although the leaves of short-shoots gain disproportionately more surface area per unit mass investment compared to the leaves produced by long-shoots, our data show that ontogenetic shifts occurring at the shoot and whole plant level account for size-dependent decreases in total canopy SLA.     

Biomass partitioning and leaf N,P - stoichiometry: comparisons between tree and herbaceous current-year shoots

We compare the biomass partitioning patterns and the nitrogen/phosphorus (N,P) stoichiometry of the current-year shoots of tree and herbaceous species and ask whether they scale in the same ways. Our analyses indicate that few statistically significant differences exist between the shoot biomass partitioning patterns of the two functional species-groups. In contrast, statistically significant N,P - stoichiometric differences exist between the two functional groups. Across all species, dry leaf mass scales nearly as the square of basal stem diameter and isometrically with respect to dry stem mass. However, total leaf N scales as the 1.37-power and as the 1.09-power of total leaf P across herbaceous and tree shoots, respectively. Therefore, tree shoots can be viewed as populations of herbs elevated by their older, woody herbaceous cohorts. However, tree leaf stoichiometry cannot be modelled in terms of herbaceous N,P - leaf stoichiometry.     

Effect of Fruiting on Leaf Gas Exchange in Olive (Olea Europaea L.)

The effect on traits of photosynthesis and water relations of assimilate demand was studied in olive tree that has strong alternate bearing. The diurnal and seasonal leaf gas exchanges, area dry mass, and saccharide and chlorophyll (Chl) contents were measured by comparing shoots with fruit of "on-trees" (heavy fruit load) with shoots without fruit on both "on-trees" and "off-trees" (light fruit load). In spite of large seasonal and diurnal differences, leaf net photosynthetic rate (P _N), stomatal conductance (g _s), sub-stomatal CO₂ concentration (C ₁), transpiration rate (E), and respiration rate (R _D) were not significantly influenced by fruit load or by the presence or absence of fruit on the shoot. An only exception was at the beginning of July when the one-year-old leaves on shoots with fruit had slightly higher P _N and E than leaves on shoots without fruit. Water content, Chl and saccharide contents, and area dry mass of the leaf were not substantially influenced by the presence/absence of fruit on the shoot or fruit load. Hence the sink demand, associated with fruit growth, did not improve leaf photosynthetic efficiency in olive. This revised version was published online in August 2006 with corrections to the Cover Date.     

Common allometric response of open-grown leader shoots to tree height in co-occurring deciduous broadleaved trees

Background and Aims

Morphology of crown shoots changes with tree height. The height of forest trees is usually correlated with the light environment and this makes it difficult to separate the effects of tree size and of light conditions on the morphological plasticity of crown shoots. This paper addresses the tree-height dependence of shoot traits under full-light conditions where a tree crown is not shaded by other crowns.

Methods

Focus is given to relationships between tree height and top-shoot traits, which include the shoot''s leaf-blades and non-leafy mass, its total leaf-blade area and the length and basal diameter of the shoot''s stem. We examine the allometric characteristics of open-grown current-year leader shoots at the tops of forest tree crowns up to 24 m high and quantify their responses to tree height in 13 co-occurring deciduous hardwood species in a cool-temperate forest in northern Japan.

Key Results

Dry mass allocated to leaf blades in a leader shoot increased with tree height in all 13 species. Specific leaf area decreased with tree height. Stem basal area was almost proportional to total leaf area in a leader shoot, where the proportionality constant did not depend on tree height, irrespective of species. Stem length for a given stem diameter decreased with tree height.

Conclusions

In the 13 species observed, height-dependent changes in allometry of leader shoots were convergent. This finding suggests that there is a common functional constraint in tree-height development. Under full-light conditions, leader shoots of tall trees naturally experience more severe water stress than those of short trees. We hypothesize that the height dependence of shoot allometry detected reflects an integrated response to height-associated water stress, which contributes to successful crown expansion and height gain.     

Effects of shoot position on shoot and leaf morphology of Avicennia marina in the hyperarid Red Sea coastal region of Egypt

We examined the effects of shoot position on shoot growth and morphology of Avicennia marina (Forssk.) Vierh. in the Red Sea coastal region of Egypt. To determine differences in morphological characteristics, we collected shoots from the upper and lower canopies of A. marina individuals in the wild and compared the morphological characteristics of these shoots. The study plot was established in an A. marina mangrove forest. Heights and diameters of individual trunks (n = 14) in the plot were measured at ground level. Then, five shoots with young but fully expanded leaves were collected from the upper and lower canopies of the individuals. We measured shoot length, and dry weight and also area, dry weight, thickness, and Soil Plant Analysis Development (SPAD) value of collected leaves. Our measurements showed that leaf area, dry weight, specific leaf area, and SPAD value of leaves from the upper canopy were smaller than those of lower-canopy leaves in most individuals. From the differences in traits between upper and lower leaves, we concluded that leaves in the upper canopy are typically adapted to high light levels, whereas leaves in the lower canopy exhibit adaptations to low light conditions. In addition, soil-water salinity at the study site was far higher than the optimum salinity for A. marina. Hence, it is also suggested the salinity level at this site may have influenced the reduced leaf size in the upper canopy.     

Annual and spatial variation in shoot demography associated with masting in Betula grossa: comparison between mature trees and saplings

Backgrounds and Aims

Shoot demography affects the growth of the tree crown and the number of leaves on a tree. Masting may cause inter-annual and spatial variation in shoot demography of mature trees, which may in turn affect the resource budget of the tree. The aim of this study was to evaluate the effect of masting on the temporal and spatial variations in shoot demography of mature Betula grossa.

Methods

The shoot demography was analysed in the upper and lower parts of the tree crown in mature trees and saplings over 7 years. Mature trees and saplings were compared to differentiate the effect of masting from the effect of exogenous environment on shoot demography. The fate of different shoot types (reproductive, vegetative, short, long), shoot length and leaf area were investigated by monitoring and by retrospective survey using morphological markers on branches. The effects of year and branch position on demographic parameters were evaluated.

Key Results

Shoot increase rate, production of long shoots, bud mortality, length of long shoots and leaf area of a branch fluctuated periodically from year to year in mature trees over 7 years, in which two masting events occurred. Branches within a crown showed synchronized annual variation, and the extent of fluctuation was larger in the upper branches than the lower branches. Vegetative shoots varied in their bud differentiation each year and contributed to the dynamic shoot demography as much as did reproductive shoots, suggesting physiological integration in shoot demography through hormonal regulation and resource allocation.

Conclusions

Masting caused periodic annual variation in shoot demography of the mature trees and the effect was spatially variable within a tree crown. Since masting is a common phenomenon among tree species, annual variation in shoot demography and leaf area should be incorporated into resource allocation models of mature masting trees.     

Some Effects of Ringing and Deblossoming Branches of Young Apple Trees on Leaf Composition

Bark rings and blossoms were removed from selected branchesof Y-shaped Sunset/M.7 apple trees at full bloom to give the10 possible combinations of ringed and unringed, deblossomedand cropping branches. Comparisons of leaf areas and dry weightper unit leaf area (specific leaf weight) showed that leavesfrom ringed branches were smaller and that extension shoot leavesfrom ringed branches had higher specific leaf weight. Fruitinghad little influence on area per leaf but gave lower specificleaf weight than deblossoming. Mineral analyses of leaves fromextension shoots in late summer were expressed in terms of unitleaf area and showed significant treatment effects. Ringingboth branches reduced the content of mineral elements but ringingonly one branch showed smaller effects on nitrogen, phosphorus,and potassium. This suggests that these elements are taken upthrough the xylem whenever the roots are adequately suppliedwith nutrients from the branches. Calcium content, on the otherhand, was appreciably reduced by any ringing, suggesting thatat least some of this element is normally transported in thephloem. There was no evidence in this experiment that low mineralnutrient levels in leaves on ringed branches had been limitingto dry matter production.     

Shoot elongation,leaf demography and bud formation in relation to branch position on Larix laricina saplings

Summary Shoot development was investigated on branches of Larix laricina (Du Roi) K. Koch trees growing in their 8th year in two plantations and in a natural stand approximately 12 years old. Expansion of throughout-crown series of short and long shoots was measured weekly, and later colour change and natural fall of leaves were assessed. Similar shoots were collected at intervals and dissected, the long shoots by 25-leaf segments. Leaf area and weight, as well as time of bud formation, were determined. Increasing acropetal trends were evident in time to bud burst: duration of short-shoot leaf-cluster expansion; size of leaf clusters and number, area and weight of leaves per cluster; duration and rate of long-shoot elongation; number, area and weight of leaves on long shoots; time to terminal-bud formation on long shoots. Along each long shoot, stem and leaf elongation and lateral-axis formation progressed acropetally. Lateral axes were most numerous on second to fourth 25-leaf segments. On longer shoots, some axes in middle segments developed as sylleptic short shoots rather than as lateral buds. Leaves of short shoots and basal leaves on long shoots turned yellow and abscissed sooner than axial leaves on long shoots. Colour change and loss among axial leaves were acropetal along shoots and up the crown. Thus, last-formed leaves, in axils of some of which lastformed lateral buds occurred, were held longest.