不同光环境下紫椴幼树树冠结构的可塑性响应

从冠形、侧枝和叶片在树冠中的空间分布角度对天然更新紫椴幼树的树冠结构进行了论述,认为紫椴幼树树冠对光照条件的变化有显著的可塑性响应.强光通过抑制主干的生长促进了侧枝的分化,庇荫则通过抑制1级侧枝的生长促进了侧枝的再分枝.随着光照水平的降低,紫椴幼树的数量叶片密度显著降低,且叶片逐渐集中于冠上层.林冠下的紫椴幼树通过这种侧枝和叶片的分布格局,在形态上提高其对光的截获能力在适度庇荫环境中,紫椴幼树垂直生长采取演替先锋种的"避荫”对策,侧枝生长采取中等耐荫种的"掠光”对策;在弱光环境中,紫椴幼树则采取典型的忍耐适应行为.这种树冠结构的变化是提高紫椴幼树对光的截获能力的一种有益适应.     

Relationships between branch spacing, growth rate and light in tropical forest saplings

1. The spacing of branches along central stems was related to growth rate and light level in forest saplings and trees in tropical moist forest on Barro Colorado Island (BCI), Panama. The study included 14 species with tiers of plagiotropic branches (having planar leaf arrangements) and four species with continuous distributions of plagiotropic branches.
2. All species showed increases in branch spacing with increasing light and growth rate of diameter, similar to the patterns in leaf spacing noted previously in species which initially bear large leaves on unbranched stems.
3. Non-tiered species had shorter internodes than tiered species but because the latter bear more branches per node, both groups had similar numbers of branches per unit stem length, when compared at similar growth rates.
4. Differences in the relationship between internode length and growth rate among tiered species were related to demographic characteristics, suggesting that tree architecture may influence forest composition.
5. The strong correlation observed between branch spacing and growth rate suggests that branch spacing may be used to estimate past growth histories of forest tree species with plagiotropic branches.     

Variation in crown light utilization characteristics among tropical canopy trees

BACKGROUND AND AIMS: Light extinction through crowns of canopy trees determines light availability at lower levels within forests. The goal of this paper is the exploration of foliage distribution and light extinction in crowns of five canopy tree species in relation to their shoot architecture, leaf traits (mean leaf angle, life span, photosynthetic characteristics) and successional status (from pioneers to persistent). METHODS: Light extinction was examined at three hierarchical levels of foliage organization, the whole crown, the outermost canopy and the individual shoots, in a tropical moist forest with direct canopy access with a tower crane. Photon flux density and cumulative leaf area index (LAI) were measured at intervals of 0.25-1 m along multiple vertical transects through three to five mature tree crowns of each species to estimate light extinction coefficients (K). RESULTS: Cecropia longipes, a pioneer species with the shortest leaf life span, had crown LAI <0.5. Among the remaining four species, crown LAI ranged from 2 to 8, and species with orthotropic terminal shoots exhibited lower light extinction coefficients (0.35) than those with plagiotropic shoots (0.53-0.80). Within each type, later successional species exhibited greater maximum LAI and total light extinction. A dense layer of leaves at the outermost crown of a late successional species resulted in an average light extinction of 61% within 0.5 m from the surface. In late successional species, leaf position within individual shoots does not predict the light availability at the individual leaf surface, which may explain their slow decline of photosynthetic capacity with leaf age and weak differentiation of sun and shade leaves. CONCLUSION: Later-successional tree crowns, especially those with orthotropic branches, exhibit lower light extinction coefficients, but greater total LAI and total light extinction, which contribute to their efficient use of light and competitive dominance.     

Influence of leaf size on tree architecture: first branch height and crown dimensions in tropical rain forest trees

 First branch height is an important attribute of sapling architecture, as it defines the height at which prolonged lateral growth is possible. First branch height, measured on saplings of 70 species in tropical rain forests of Australia, Costa Rica, Panama, and Sabah, Malaysia, was highly correlated with leaf blade and petiole length. The observed relationship, first branch height ∝ blade length × (petiole length)^0.5, implies that the ratio of first branch height to blade length increases somewhat with increasing leaf size, among species with a given ratio of petiole to blade length. Orthotropic species, with more or less radially symmetric arrangements of leaves on ascending axes, had a mean first branch height of 7x that observed for plagiotropic species, with planar leaf arrangements. The greater first branch height of orthotropic species was associated with their larger leaves and longer petioles. Plagiotropic species had wider crowns than orthotropic species in the sapling stage, as assessed at the Costa Rican site. Thus, leaf dimensions influence the dynamics of crown construction (or visa versa), as well as affecting leaf energy balance and gas exchange. Received: 5 September 1997 / Accepted: 3 March 1998     

Within-crown variation in the timing of leaf emergence and fall of Malaysian trees in association with crown development patterns

In aseasonal tropics, timing of leaf emergence and leaf fall may differ between the shoots of different crown parts within a tree. This is important for the efficient development of crowns because leaves should be produced as soon as enough carbohydrates are accumulated. This hypothesis was tested by investigating leaf demography over a 44-mo period for 17 Malaysian trees and comparing the timings of leaf emergence and fall between the upper and lower crowns. The timings of leaf emergence were synchronized between the upper and lower crowns, but those of leaf fall were less synchronized in most trees. Greater rates of leaf production in the upper than in the lower crowns were attributable to the differences in the number of leaves that emerged per leaf emergence event, rather than differences in frequency of leaf emergence per year. Timings of leaf emergence and leaf fall were mainly simultaneous in the upper and lower crowns, but unsynchronized leaf production and leaf fall also occurred. Such limited plasticity of leaf demography within crowns may be the result of physiological integration of branches or the compromise between the advantages of satiating herbivores and effective crown development in the trees of aseasonal tropics.     

LEAF DISPLAY,CANOPY STRUCTURE,AND LIGHT INTERCEPTION OF TWO UNDERSTORY PALM SPECIES

The implications of leaf size, leaf display, and crown size for whole-plant light interception were investigated in Geonoma cuneata and Asterogyne martiana, two understory palm species native to Central American rain forests. Adults of A. martiana had longer leaves, more leaves per plant, and greater total leaf area than G. cuneata adults. Geometric measurements within whole crowns were used to calculate light interception efficiency, a leaf-based measure of the proportion of total incident light that is intercepted by a crown. Light interception efficiency was higher in adult G. cuneata than in adult A. martiana; seedlings of the two species did not differ significantly in light interception efficiency. Decreased efficiency of adult A. martiana crowns was largely due to an increased proportion of pendent leaves. Compared to G. cuneata, adults of A. martiana had greater light interception capacity (the product of light interception efficiency and total leaf area), but they also had higher biomass costs of light interception. Lower biomass costs of light interception in adult G. cuneata enable this species to exploit successfully the most deeply shaded microsites in the rain forest understory.     

Crown structure and vertical foliage distribution in 4-year-old plantation-grown Eucalyptus pilularis and Eucalyptus cloeziana

Tree growth and form are both influenced by crown architecture and how it effects leaf distribution and light interception. This study examined the vertical distribution of foliage in 4-year-old plantation-grown Eucalyptus pilularis Sm. and E. cloeziana F. Muell. trees. Leaf area (LA) distribution was determined at two different sites using allometric approaches to determine LA in crown sections and for whole trees. Leaf area was distributed more towards the upper crowns when canopies had been closed for longer. Leaf area was also skewed more towards the upper crowns for Eucalyptus pilularis than E. cloeziana. These species differences were consistent with differences in vertical light availability gradients as determined by point quantum sensors. Leaf area of individual branches was highly correlated with branch cross-sectional area (CSA) and whole-tree LA was closely related to stem CSA. Branch-level allometric relationships were influenced by site and crown position. However, the general allometric equations between stem size and whole-tree leaf area could be applied across sites. Results from this study suggest that pruning of live branches in these species should follow species-specific guides for the timing and height of pruning to optimise the effects on stem growth and form.     

Allometry, root/shoot ratio and root architecture in understory saplings of deciduous dicotyledonous trees in central Japan

Plant allometry that is related to plant architecture and biomass allocation strongly influences a plants ability to grow in shaded forest understory. Some allometric traits can change with plant size. The present study compared crown and trunk allometries, root/shoot biomass allometry, and root architecture among understory saplings (0.25--5m height, except for two trees > 5 < 7 m) of seven deciduous dicotyledonous species in central Japan. Associations of the crown and trunk allometries with several plant morphological attributes were analyzed. Branch morphology (plagiotropyvs orthotropy) and life size were correlated with sapling crown and trunk allometries. Both large leaves and orthotropic branches were associated with a narrow small crown and slender trunk. The root/shoot ratio decreased rapidly with increasing plant height for saplings shorter than about 1.5 m. Less shade-tolerant species tended to have smaller root/shoot ratios for saplings taller than 1.5 m. With an increase in plant height, the branch/trunk biomass ratio decreased for saplings with plagiotropic branches but increased for saplings with orthotropic branches. Four subcanopy species (Acer distylum, Carpinus cordata, Fraxinus lanuginosa and Acanthopanax sciadophylloides) had superficial root systems; a common understory species (Sapium japonica) had a deep tap root system; and a canopy species (Magnolia obovata) and a subcanopy species (Acer tenuifolium) had heart root systems of intermediate depth. The root depth was not related to shade tolerance. Among species of the same height, the difference in fine root length can be 30-fold.     

Comparison of the physiology, morphology, and leaf demography of tropical saplings with different crown shapes

Branch architecture, leaf photosynthetic traits, and leaf demography were investigated in saplings of two woody species, Homolanthus caloneurus and Macaranga rostulata, co-occurring in the understory of a tropical mountain forest. M. rostulata saplings have cylindrical crowns, whereas H. caloneurus saplings have flat crowns. Saplings of the two species were found not to differ in area-based photosynthetic traits and in average light conditions in the understory of the studied site, but they do differ in internode length, leaf emergence rate, leaf lifespan, and total leaf area. Displayed leaf area of H. caloneurus saplings, which have the more rapid leaf emergence, was smaller than that of M. rostulata saplings, which have a longer leaf lifespan and larger total leaf area, although M. rostulata saplings showed a higher degree of leaf overlap. Short leaf lifespan and consequent small total leaf area would be linked to leaf overlap avoidance in the densely packed flat H. caloneurus crown. In contrast, M. rostulata saplings maintained a large total leaf area by producing leaves with a long leaf lifespan. In these understory saplings with a different crown architecture, we observed two contrasting adaptation strategies to shade which are achieved by adjusting a suite of morphological and leaf demographic characters. Each understory species has a suite of morphological traits and leaf demography specific to its architecture, thus attaining leaf overlap avoidance or large total leaf area.     

Irradiance heterogeneity within crown affects photosynthetic capacity and nitrogen distribution of leaves in Cedrela sinensis

Because light conditions in the forest understory are highly heterogeneous, photosynthetic acclimation to spatially variable irradiance within a crown is important for crown‐level carbon assimilation. The effect of variation in irradiance within the crown on leaf nitrogen content and photosynthetic rate was examined for pinnate compound leaves in saplings of Cedrela sinensis, a pioneer deciduous tree. Five shading treatments, in which 0, 25, 50, 75 and 100% of leaves were shaded, were established by artificial heavy shading using shade screen umbrellas with 25% transmittance. Although the nitrogen content of leaves was constant regardless of shading treatment, ribulose 1·5‐bisphosphate carboxylase/oxygenase (Rubisco) content and light‐saturated photosynthetic capacity were lower in shade leaves within partially shaded crowns than within fully shaded crowns. Shade leaves within partially shaded crowns contained higher amount of amino acids. Most shade leaves died in partially shaded crowns, whereas more than half of shade leaves survived in totally shaded crowns. Assumptions on photosynthetic acclimation to local light conditions cannot explain why shade leaves have different photosynthetic capacities and survival rates in between partially and totally shaded crowns. Irradiance heterogeneity within the crown causes a distinct variation in photosynthetic activity between sun and shaded leaves within the crown.     

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.     

Branching responses of a plagiotropic clonal herb to localised incidence of light simulating that reflected from vegetation

Plants sense the presence of neighbouring vegetation through phytochrome photoreceptors perceiving a lowered red to far-red ratio (R:FR) of light reflected from such vegetation. We hypothesised that it would be advantageous for the grassland clonal herb, Trifolium repens, to have an inhibitory branching response to perception by leaves of light reflected from neighbouring vegetation (i.e. light with lowered R:FR ratio) but have no response to interception of such light by the plagiotropic stem. We tested whether photoreception of reflected light by plagiotropic stems resulted in a different branching response to photoreception by leaves and whether leaf ontogeny influenced the response. To simulate light reflected from vegetation, FR light-emitting-diodes were used to supplement controlled environment room light so that the R:FR ratio, but not the photosynthetic photon fluence rate, of light incident at the stem or leaf of a phytomer of T. repens was lowered from 1.20 to 0.25. The plagiotropic stems were unresponsive to light simulating that reflected from vegetation. This response differs from that of stems of orthotropic species, indicating that plagiotropic stems have evolved an organ-specific photobiology. Treatment of the mature leaf with light of lowered FR ratio reduced phytomer production only of the branch in the axil of the treated leaf. Similar treatment of the immature leaf retarded, in addition, branching at basal phytomers on the same side of the primary stem axis. Thus the response to light simulating that reflected from neighbouring vegetation depended upon whether the light was incident at the stem or the leaf and on the stage of leaf development. We argue that such responses improve the performance and fitness of T. repens within grassland habitats by allowing axillary buds on plagiotropic stems to branch freely when stems are in receipt of light reflected from vegetation while leaves are in full light.     

Morphology and growth of deciduous and evergreen broad-leaved saplings under different light conditions in a Chinese beech forest with dense bamboo undergrowth

This study compared the morphological and growth adjustment of saplings from three shade- tolerant canopy species (Castanopsis lamontii, Lithocarpus hancei and Fagus lucida; Fagaceae) under different light conditions in a Chinese beech forest with dense bamboo undergrowth. Only F. lucida is deciduous, and it had the most flexible morphology. In shade, F. lucida had flat or bent topshoots and horizontal branches to maximize light interception, while in conditions of high light intensity, it formed vertical topshoots to promote growth in terms of height, and upright branches to mitigate excessive sun exposure on the leaves. In contrast to F. lucida (beech), the evergreen species always had vertical topshoots regardless of light conditions. In shady conditions, the evergreens had greater annual growth rates in terms of both height and diameter than the beech, and between the evergreens, the species bearing plagiotropic branches grew faster in diameter than the species bearing orthotropic branches. The evergreen trees had thicker leaves and thicker stems compared to the beech. It was concluded that the evergreen saplings have advantages over beech saplings in terms of current growth in the forest understorey; whereas, morphological and growth flexibility in the beech aids in its persistence in the understorey.     

不同光环境下木荷幼苗树冠结构的可塑性响应

从冠形、侧枝和叶片在树冠中的空间分布等方面对天然更新木荷（Schima superba Gardn．et Champ．）幼苗的树冠结构进行了研究,认为木荷幼苗的树冠对光照条件的变化有显著的可塑性响应。随着光照水平的提高,幼苗树冠由阔、松散型向相对紧密、窄冠型发展,表明木荷幼苗对不同光照环境有较强的适应能力。木荷幼苗在强光环境下产生短枝,在适度荫庇条件下侧枝和主枝同时向上方和侧方伸长生长;在强度遮阴条件下,侧枝发生强烈的伸长生长并发生强烈的分枝行为,同时在不同自然环境条件下幼苗的叶片密度由全光、林隙到林冠下逐渐提高。随着光照水平的减弱,1级侧枝密度逐渐降低,分枝（2级侧枝和3级侧枝）强度却逐渐增大,且侧枝在树冠上的分布有向匕集中的趋势。     

Functional correlates of leaf demographic response to gap release in saplings of a shade-tolerant tree,<Emphasis Type="Italic"> Elateriospermum tapos</Emphasis>

For a shade-tolerant SE Asian tropical tree, Elateriospermum tapos (Euphorbiaceae), we studied field-established saplings in gaps and the shaded understory to test the hypothesis that differences in leaf demography and leaf life span under contrasting light regimes should be functionally correlated with architecture, self-shading and nitrogen distribution within the sapling crown. Rates of leaf production and net leaf gain were greater for saplings in gaps than those in the understory. Median leaf life span was approximately 26 months in the gap saplings, while it was estimated to be greater than 38 months in the understory saplings. Consequently, gap saplings had a greater standing leaf number and experienced greater degrees of self-shading than understory saplings. Light availability at individual leaves, estimated by a combination of canopy photos and a three-dimensional architecture model, were negatively correlated with leaf age in gap saplings but not so in understory saplings. Leaf nitrogen content per unit area (N_area) was influenced more by light availability than by leaf age in the gap saplings. In contrast, in understory saplings, N_area was neither correlated with light availability nor with leaf age, and did not decrease significantly before 38 months in leaf age. We conclude that saplings of this shade-tolerant species apparently prolong their leaf life span in the shaded understory through slower rates of leaf production, lower standing number of leaves and lower degrees of self shading than in gap, and that the rate of decline of N_area with leaf age depends on architecture and self-shading regimes that respond to changes in light regimes.     

光照条件、植株冠层结构和枝条寿命的关系——以桂花和水杉为例

根据衰老理论的代谢率假说,生物寿命与其代谢率有关,个体大小相同的生物体,在质量较好的微生境中通常比较差生境中具有更高的代谢速率。因此,生物体在资源供给较差的生境中通常比资源供给较充足的生境中具有更长的寿命。枝条是木本植物植冠构建的基本单元之一,如果枝条遵循代谢率假说,则可推测在光照较好环境下的植物枝条或小枝将比其在遮荫环境下具有更短的寿命,即枝条寿命与光照条件成反比。以常绿物种桂花(Osmanthus fragrans)和落叶物种水杉(Metasequoiaglyptostroboides)为研究对象,通过测量不同光照环境下,植株大小(株高和胸径)、冠层深度、冠层轮廓(冠层深度/冠层宽度)、相对冠层宽度(冠层宽度/植株高度)以及植株凋落枝条寿命等性状,探讨了光照条件对成年植株冠层形态结构和植株枝条寿命的影响。调查发现:1)枝条的寿命在遮荫条件下显著高于全光照条件下,与理论预测吻合;2)随遮荫程度增加,植株冠层深度和冠层轮廓增加,相对冠层宽度减小;3)枝条的平均寿命与植株冠层深度和冠层轮廓成正比,与植株相对冠层宽度成反比。这表明光照条件可能通过改变植株冠层结构来影响枝条寿命。未来需要进一步研究枝条生物量分配、叶片光合能力和呼吸速率在不同生活型物种之间的差异,以便更全面的理解枝条寿命与生境质量之间的关系。     

Influence of light level on the growth and morphology of saplings in a panamanian forest

Leaf spacing and aboveground growth were monitored in saplings of ten species in a range of light environments in a Panamanian lowland humid forest. One- to 2-m-tall individuals of the chosen species had intermediate to large leaves on stems with few or no branches. Saplings in high light environments grew faster in height and produced more biomass per unit leaf area than shaded saplings for all species. These growth responses involved morphological plasticity with greater extension per unit biomass increment increasing the height growth rate of gap-grown saplings and greater biomass allocation to leaves decreasing the whole plant light compensation point in shade. The relative performance of the species also varied across the light gradient and was related to differences in leaf lifespan and specific leaf mass. Light-demanding species grew as rapidly in shade as shade-tolerant species, but the shorter leaf lifespan of the former necessitates higher production rates to maintain a given leaf area, largely excluding light-demanders from shaded understory locations. Height growth rate was positively correlated with leaf spacing for each species, and differences between species in the height growth rate-internode length relationship were related to interspecific differences in specific leaf mass. Thus, sapling growth histories may be inferred from their morphologies.     

干旱胁迫下新疆杨树冠不同高度叶片水分状况与非结构性碳动态北大核心CSCD

水力结构调整及非结构性碳动态对于认知干旱胁迫下植物生存前景和死亡风险至关重要。该研究以新疆杨(Populus bolleana Lauche.)为对象,通过干旱处理分析树冠不同高度叶片水力性状、光合生理特性、不同功能器官间的可溶性糖、淀粉及非结构性碳水化合物(NSC)含量变化,以甄别树冠不同高度叶片对干旱胁迫的生理响应和适应差异。结果表明:(1)在干旱胁迫下,树冠上部的叶水势、叶含水量、枝条含水量普遍低于下部,饱和膨压渗透势及膨压损失点水势在不同树冠高度间差异不显著;(2)干旱处理组植株净光合速率随树冠高度的增加而降低,叶绿素SPAD值在树冠的上部显著低于下部,而水分利用效率在树冠的上部却高于下部;(3)干旱处理组比叶面积在各树冠高度下显著低于对照组,而胡伯尔值在中部及上部高于对照组但差异不显著;(4)干旱处理组,树冠上部叶片淀粉含量显著高于下部,枝条可溶性糖及NSC含量在树冠上部显著高于下部,韧皮部的可溶性糖、淀粉及NSC含量在不同树冠高度间无显著差异,细根的可溶性糖、淀粉及NSC含量在对照组与干旱处理组间的差异不显著。研究发现,干旱处理下,树冠高度的增加会加剧新疆杨枝叶的干旱胁迫,致使树冠上部枝条木质部发生栓塞的风险大于下部,并导致NSC在不同器官间的分配和组分产生差异,但新疆杨植株可通过水分利用效率和形态上的适应性调整减缓干旱胁迫。     

两种亚热带榕属乔木植冠损伤后的位置依赖性枝发生

三峡库区需要大量植株用于三峡工程所涉及的诸多建设(如铁路、公路、建筑)之后的植被恢复和绿化.由于具有优美树冠和耐瘠薄土壤的能力,榕(Ficus microcarpa L.)和黄桷树(Fvirens Ait.var.sublanceolata(Miq.)Cornor) 在三峡库区作为绿化和行道树木大量栽植.在三峡库区,这两种树种苗木的培育主要通过切枝扦插的方式进行.大量切枝损伤植株植冠并且使叶组织数量减少,对植株生长有很大影响.植株生长与植株的枝发生格局有很大关系,为明确植冠损伤对植株生长的可能影响,对榕和黄桷树植冠损伤后的枝发生进行了研究.实验发现,切枝造成的植冠损伤对榕和黄桷树植株侧枝上的枝发生没有影响,不同损伤强度之间和不同损伤发生时间之间都不存在明显的差异.但是,植冠损伤后,榕和黄桷树植株主茎上的枝发生是位置依赖性的.植冠损伤不影响榕和黄桷树植株损伤处理后新生主茎段和具侧枝主茎段上的枝发生,却促进了损伤处理后裸露主茎段上的枝发生,并且裸露主茎段上枝发生的数量和密度随植冠损伤强度的升高而增大.此外,实验结果表明,秋季植冠损伤植株裸露主茎段上的枝发生要高于春季植冠损伤植株裸露主茎段上的枝发生;在榕和黄桷树的枝发生中,仅有裸露主茎段上增强的枝发生会促进植冠损伤植株对叶组织的生物量投资并有利于受损植株的恢复和生长.