亚高山暗针叶林不同林冠环境下华西箭竹的克隆可塑性

以亚高山暗针叶林3种林冠环境中以及暗针叶林林缘的华西箭竹（Fargesia nitida）为对象,对其无性系数量特征、无性系根茎特征、分株生物量以及分株形态特征进行了对比研究。结果表明：（1）林冠环境的差异导致了不同种群的基株密度和每基株分株数的显著差异,但林冠环境差异不影响分株密度。林冠郁闭度愈大,每基株分株数愈少,分株分布愈均匀。（2）不同林冠环境间。分株生物量、分株构件生物量和分株构件的生物量分配百分率均有显著差异。开敞的林冠环境有利于华西箭竹的生长和生物量积累。（3）随着林冠郁闭度的增加,华西箭竹通过增大分枝角度、叶生物量分配百分率、比叶面积和叶面积率以提高光能利用效率,有效适应弱光环境。（4）隔离者长度、隔离者直径和分枝强度在林缘和林窗环境中要显著大于林内环境;同级隔离者分枝角度随林冠郁闭度的增加而最大,其值在林下显著大于林窗和林缘,而异级隔离者分枝角度的变化则正好相反。研究表明,华西箭竹种群在不同的林冠环境中发生了明显的可塑性变化,这些可塑性变化是种群对林冠郁闭度差异的适应性反应的结果,有利于增强种群对环境中有限光资源的利用。     

Response of clonal plasticity of Fargesia nitida to different canopy conditions of subalpine coniferous forest

The aim of this study is to explore the effects of canopy conditions on clump and culm numbers, and the morphological plasticity and biomass distribution patterns of the dwarf bamboo species Fargesia nitida. Specifically, we investigated the effects of canopy conditions on the growth and morphological characteristics of F. nitida, and the adaptive responses of F. nitida to different canopy conditions and its ecological senses. The results indicate that forest canopy had a significant effect on the genet density and culm number per clump, while it did not affect the ramet density. Clumps tended to be few and large in gaps and forest edge plots, and small under forest understory plots. The ramets showed an even distribution under the closed canopy, and cluster distribution under gaps and forest edge plots. The forest canopy had a significant effect on both the ramets’ biomass and biomass allocation. Favourable light conditions promoted ramet growth and biomass accumulation. Greater amounts of biomass in gaps and forest edge plots were shown by the higher number of culms per clump and the diameter of these culms. Under closed canopy, the bamboos increased their branching angle, leaf biomass allocation, specific leaf area and leaf area ratio to exploit more favourable light conditions in these locations. The spacer length, specific spacer length and spacer branching angles all showed significant differences between gaps and closed canopy conditions. The larger specific spacer length and spacer branching angle were beneficial for bamboo growth, scattering the ramets and exploiting more favourable light conditions. In summary, this study shows that to varying degrees, F. nitida exhibits both a wide ecological amplitude and high degree of morphological plasticity in response to differing forest canopy conditions. Moreover, the changes in plasticity enable the plants to optimize their light usage efficiency to promote growth and increase access to resources available in heterogeneous light environments. __________ Translated from Acta Ecologica Sinica, 2006, 26(12): 4019–4026 [译自: 生态学报]     

卧龙亚高山暗针叶林不同林冠环境下华西箭竹分株种群结构特征

详细调查了林下、中林窗、大林窗和林缘旷地等4种亚高山暗针叶林林冠环境下的华西箭竹（Fargesia nitida）分株种群,对其表现结构（株高、基径和生物量）和年龄结构进行了较系统的对比研究.主要研究结果如下：（1）分株水平上,华西箭竹表现结构在4种林冠环境下均有极显著差异（p＜0.01）,且随林冠郁闭度的减小,分株个体的高度、基径和生物量有递增的趋势（林下＜中林窗＜大林窗）;（2）分株各构件在总生物量中所占比例随林冠环境变化而改变：除林缘旷地外,叶的生物量百分比与林冠郁闭度呈正相关.林缘旷地中,地下茎和根系的生物量百分比均高于其余3种环境;（3）不同林冠环境下分株单位叶面积叶重存在显著差异（p＜0.05）,且随林冠郁闭度的减小而增大.单叶生物量和单叶面积均以中林窗最大,林缘旷地次之,二者与大林窗或林下均有显著差异（p＜0.05）.华西箭竹的单株叶片数量以大林窗最大,与其余3种环境有极显著差异（p＜0.01）;（4）分株种群的死亡率以林下最低（P＜0.01）,但各种群平均年龄间无显著差异（p＞0.05）.可见,华西箭竹分株种群对林冠环境变化的反应主要体现在形态和生物量分配上,而非种群的年龄上.     

Pattern and Performance of Understory Bamboos (Chusquea spp.) under Different Canopy Closures in Old-Growth Oak Forests in Costa Rica1

The effect of canopy closure on spatial and morphological patterns of three understory Chusquea species was studied in pristine upper montane oak forests. Plots with either Chusquea talamancenis Widmer & L.G. Clark or C. tomentosa Widmer & L.G. Clark were set out in gaps, under intermediate tree canopy and under closed canopy. The third species Chusquea foliosa L.G. Clark was only found in gap and intermediate canopy sites in the study area. Clumps of all bamboo species tend to be few and large under open conditions and numerous and small under more shady conditions. The larger size of clumps in gaps is reflected in the number of culms per clump, the diameter of the culms and, as a consequence, clump basal area, clump crown area, and biomass (index of plant volume) compared to intermediate and closed canopy. The smaller clump density in gaps compared to closed canopy implies that there is more intraspecific competition and density dependent mortality (self-thinning effect) when a bamboo species is dominant under favourable light conditions. Parameters of performance like culm length, number of nodes per culm, number and length of primary branches, and number of branch nodes seem not to be affected by the light regime, unlike total number of branches, total branch length, and the branching pattern. Species differ in their response to the light environment: C. tomentosa and C. foliosa have a higher degree of morphological plasticity than C. talamancensis, which in turn, appears to be more shade tolerant. No difference between species has been found regarding their contribution to vegetation, parameters reflecting abundance per unit area (site clump area, site crown area and site index of clump volume) were similar for all three species.     

紫耳箭竹克隆形态可塑性对典型冠层结构及光环境的响应

在重庆金佛山国家自然保护内,选择了3种典型群落类型(落叶阔叶林、常绿落叶阔叶混交林和常绿阔叶林),使用Hemiview数字植物冠层分析系统量化群落冠层结构和光环境特征,并对林下紫耳箭竹(Fargesia decurvata)的形态可塑性特征进行调查,分析冠层结构和光环境特征改变下紫耳箭竹形态可塑性的差异,并探讨它们之间的相互关系。结果表明:(1)随着落叶阔叶林"常绿落叶阔叶混交林"常绿阔叶林演替的进行,群落的冠层开度降低,叶面积指数增加,平均叶倾角变小,趋于水平化,冠层对光的截获能力提高,林下光照的强度降低(P0.05)。(2)随着光照强度的降低,紫耳箭竹分株矮小化,叶片变窄,生物量积累降低,但通过增大比茎长、叶面积率和比叶面积提高对光的利用效率,并增大分枝角度和比隔长有效适应弱光环境。(3)在光照条件差的常绿阔叶林下,紫耳箭竹降低对地下茎的投资,将较多的生物量用于秆的增高增长和叶片的生长;而在光照条件好的落叶阔叶林环境下,紫耳箭竹降低对枝、叶生物量的分配,则加大对地下茎的投资,可认为是克隆植物对水分资源所表现的一种觅食行为。研究表明,紫耳箭竹种群随着冠层结构的改变发生了明显的可塑性变化,这些可塑性变化是种群对冠层结构和光环境差异的适应性反应的结果,有利于增强种群对异质生境中光资源的获取和利用;群落内部可以通过调控冠层结构的改变协调和控制小径竹种群的发展。     

Patch structure and ramet demography of the clonal tree, Asimina triloba, under gap and closed-canopy

Clonal understory trees develop into patches of interconnected and genetically identical ramets that have the potential to persist for decades or centuries. These patches develop beneath forest canopies that are structurally heterogeneous in space and time. Canopy heterogeneity, in turn, is responsible for the highly variable understory light environment that is typically associated with deciduous forests. We investigated what aspects of patch structure (density, size structure, and reproductive frequency of ramets) of the clonal understory tree, Asimina triloba, were correlated with forest canopy conditions. Specifically, we compared A. triloba patches located beneath closed canopies and canopy gaps. We also conducted a three-year demographic study of individual ramets within patches distributed across a light gradient. The closed canopy-gap comparison demonstrated that the patches of A. triloba had a higher frequency of large and flowering ramets in gaps compared to closed-canopy stands, but total ramet density was lower in gaps than in closed canopy stands. In the demographic study, individual ramet growth was positively correlated with light availability, although the pattern was not consistent for all years. Neither ramet recruitment nor mortality was correlated with light conditions. Our results indicate that the structure of A. triloba patches was influenced by canopy condition, but does not necessarily depend on the responses of ramets to current light conditions. The lack of differences in ramet recruitment and mortality under varying canopy conditions is likely to be a primary reason for the long-term expansion and persistence of the patches. The primary benefit of a positive growth response to increasing light is the transition of relatively small ramets into flowering ramets within a short period of time.     

Ramet Population Structure of Fargesia nitida (Mitford) Keng f. et Yi in Different Successional Stands of the Subalpine Coniferous Forest in Wolong

Forest structure and succession in Wolong Nature Reserve is influenced by the understory dwarf bamboo population. However, less is known about how the forest succession affects the dwarf bamboo population. To examine the bamboo ramet population growth of Fargesia nitida (Mitford) Keng f. et Yi and to determine how ramet population structure varies along the succession of coniferous forest, we sampled ramet populations of F. nitida from the following three successional stages: (i) a deciduous broad-leaved (BL) stand; (ii) a mixed broad-leaved coniferous (MI) stand; and (iii) a coniferous (CF) stand. We investigated the population structure, biomass allocation, and morphological characteristics of the bamboo ramet among the three stand types. Clonal ramets, constituting the bamboo population, tended to become short and small with succession. The ramet changed towards having a greater mass investment in leaves, branches and underground roots and rhizomes rather than in the culm. With respect to leaf traits, individual leaf mass and area in the BL stand were markedly bigger than those in both the MI and CF stands, except for no significant difference in specific leaf area. The age distribution showed that the bamboo population approached an older age with succession. The results demonstrate that the ramet population structure of F. nitida is unstable and its growth performance is inhibited by succession.（Author for correspondence. E-mail: taojianping@163.com）     

卧龙亚高山暗针叶林不同林冠环境下华西箭竹的克隆生长

该文调查了林下、中林窗、大林窗和林缘旷地等4种亚高山暗针叶林林冠环境下的华西箭竹(Fargesia nitida)分株种群,对其当年生立竹和母株的特征进行了比较研究,并探讨了母株年龄及大小对克隆生长的影响。主要研究结果如下:1)华西箭竹当年生立竹的分株密度以林下种群的最低,从林下→中林窗→大林窗,随林冠郁闭度的减小,华西箭竹基株当年生立竹的分株数逐渐增大。2)4种林冠环境下,当年生立竹和母株的分株高度、基径和生物量均有显著差异,且随林冠郁闭度的减小有递增的趋势(林下<中林窗<大林窗)。3)各林冠环境均以二龄母株产生的当年生立竹数量最大,不同林冠环境之间母株的平均年龄,以及处于同一林冠环境的各龄级母株产生的当年生立竹平均分株数和平均基径均无显著差异。4)不同林冠环境下,华西箭竹当年生立竹基径与一级母株、二级母株基径均呈正相关关系,但当年生立竹基径随一级母株增加的速度快于随二级母株的增加。5)当年生立竹的基径与地下茎直径呈显著的正相关;母株的基径与当年生立竹的地下茎直径呈正相关关系,而与其地下茎长度相关性不明显。     

林冠环境对亚高山针叶林下缺苞箭竹生物量分配和克隆形态的影响

通过对亚高山针叶林的林下、小林窗(130 m²)、中林窗(300 m²)和大林窗(500 m²) 4种林冠环境中缺苞箭竹(Fargesia denudata)分株种群特征进行调查, 研究其生物量分配格局和克隆形态可塑性。研究结果表明: (1)分株生物量、基径、高度以及分株各构件生物量随林冠郁闭度减小均表现为先增加后减小的趋势, 在小林窗中达到最大值; (2)大林窗中, 根生物量分配和数量显著高于其他林冠环境, 随林冠郁闭度增大, 分株不断增大叶生物量分配、比叶面积和分枝百分比等地上部分投资, 以适应低光环境; (3)比茎长和比地下茎长随林冠郁闭度增大表现为先减小后增加的趋势, 在小林窗值最低, 分枝强度在小林窗和中林窗中显著高于林下和大林窗环境。结果显示, 缺苞箭竹在不同林冠环境中具有生物量分配和克隆形态的可塑性, 以利于种群对光资源的有效利用。小林窗环境是缺苞箭竹较适宜的生境, 生物量积累最多, 长势最好。     

Impact of Culm Harvest on Seed Production in a Monocarpic Bamboo

Successful regeneration of bamboos from seed is a key issue in the ecology of many tropical regions and the livelihoods of their inhabitants. The gregarious monocarpy of many bamboos may be driven by a need to satiate seed predators by seeding in abundance at infrequent intervals. In long‐lived clonal monocarps, seed production is expected to be positively related to the success of the clone in generating more and larger ramets during its lifetime. Ramification may be constrained by harvesting of culms, but it is unclear whether the reduction in productivity is proportional to the loss of reproductive biomass. We counted the seed produced by 661 culms (ramet stems) sampled from 90 clumps of the gregariously monocarpic bamboo Schizostachyum dullooa that is intensively harvested by villagers in northeastern India. The smallest clumps had fewer culms and few or no culms more than one year old. Seed production was indeed positively related to culm size and the number of culms in a clump. First‐year culms were markedly more productive than older culms after controlling for culm diameter and clump size. There was a negative effect of clump size on productivity per culm which may occur because clumps that had been harvested heavily were able to exploit resources retained in rhizomes from harvested culms. Nevertheless, small clumps produced much less seed than larger clumps, generating a risk of unknown magnitude that heavily harvested stands of monocarpic bamboos may be unable to satiate seed predators during their single opportunity for reproduction.     

冠层结构和光环境的时空变化对紫耳箭竹种群特征的影响

选择金佛山国家自然保护区内落叶阔叶林、常绿落叶阔叶混交林、常绿阔叶林3种典型群落类型,研究冠层结构和光环境特征,以及林下优势种--紫耳箭竹的种群特征.结果表明: 随着落叶阔叶林→常绿落叶阔叶混交林→常绿阔叶林的演替,Shannon多样性指数、Simpson优势度指数和Pielou指数呈增加趋势,表明群落趋于稳定发展的状态;冠层结构特征也发生了显著改变,冠层开度和平均叶倾角减小,叶面积指数增加,冠层的消光能力增强,林下光照水平降低.上层林冠是造成林型郁闭的主要原因,其中冠层厚度和冠层面积是2个主要的影响因素.冠层结构与林下光照指标显著相关,对林下散射光的影响最大.冠层开度、林下光照条件均随着生长季的到来而下降,而叶面积指数呈现增长的趋势,峰值出现在6、7月;平均叶倾角在春季达到最大值,在夏季为最小值.紫耳箭竹的生长与冠层结构和光环境密切相关,其在光照适中的常绿落叶阔叶混交林中生长得最好,分株粗壮、密度大(29.69±1.68株·m^-2),地下茎拓展能力强;落叶阔叶林中的强光环境可能造成土壤水分缺失,从而对其生长产生影响;而在常绿阔叶林的低光环境下紫耳箭竹分株矮小,密度小(5.80±1.16株·m^-2),克隆扩展能力降低.在森林结构演变的过程中,冠层结构发生了明显的改变,显著影响林下光环境,过度的低光环境对紫耳箭竹种群的更新和发展有限制作用.     

How does the longevity of <Emphasis Type="Italic">Sasa kurilensis</Emphasis> ramets respond to a light gradient? An analysis of ontogenetic changes to hydraulic resistance and carbon budget within a ramet

We investigated how Sasa kurilensis ramet longevity differs under three light conditions. Ramet longevity is an important factor affecting the abundance of ramet populations and their biomass. The objectives of this study were to clarify: (1) whether ramet longevity varies spatially along a gradient of light conditions; (2) whether physiological functions, including water transport in culms, change with ontogenesis; (3) whether ramet longevity is associated with the most effective turnover for gaining a carbon profit within a ramet following a cost-benefit model or is affected by other factors such as death caused by a decline in physiological function. We analyzed S. kurilensis ramet longevity, hydraulic resistance, photosynthetic rate, and carbon content. We then estimated the ramet carbon budget. The longevity of S. kurilensis ramets was 2.8–4.5 years in Beneath-patch plots, 5.8–8.7 years in Edge-patch plots, and 1.6–2.2 years in Open-patch plots. Although leaf photosynthetic activity was stable, the instantaneous photosynthetic rate decreased during clear days in the open area. This may have been due to increased ramet hydraulic resistance. The rotation period of the most efficient carbon budget quantified by ecophysiological measurements was consistent with ramet longevity in Open-patch (2 years) and Edge-patch (5 years) sites. Meanwhile, the longevity of ramets grown under canopies was inconsistent with the cost-benefit model rules for a carbon budget because the carbon gain was low throughout the ramet life span.     

Genet dynamics of a regenerating dwarf bamboo population across heterogeneous light environments in a temperate forest understorey

Despite the advantage of plant clonality in patchy environments, studies focusing on genet demography in relation to spatially heterogeneous environments remain scarce. Regeneration of bamboos in forest understoreys after synchronous die‐off provides an opportunity for assessing how they come to proliferate across heterogeneous light environments. In a Japanese forest, we examined genet demography of a population of Sasa kurilensis over a 7‐year period starting 10 years after die‐off, shortly after which some genets began spreading horizontally by rhizomes. The aboveground biomass was estimated, and genets were discriminated in 9‐m² plots placed under both canopy gaps and closed canopies. Overall, the results suggest that the survival and spread of more productive genets and the spatial expansion of genets into closed canopies underlie the proliferation of S. kurilensis. Compared to canopy gaps, the recovery rate of biomass was much slower under closed canopies for the first 10 years after the die‐off, but became accelerated during the next 7 years. Genet survival was greater for more productive genets (with greater initial number of culms), and the spaces occupied by genets that died were often colonized afterward by clonal growth of surviving genets. The number of genets decreased under canopy gaps due to greater mortality, but increased under closed canopies where greater number of genets colonized clonally from outside the plots than genets died. The colonizing genets were more productive (having larger culms) than those originally germinated within the plots, and the contribution of colonizing genets to the biomass was greater under closed canopies. Our study emphasizes the importance of investigating genet dynamics over relevant spatiotemporal scales to reveal processes underlying the success of clonal plants in heterogeneous habitats.     

Morphological and Physiological Acclimation Responses to Contrasting Light and Water Regimes in <Emphasis Type="Italic">Primula sieboldii</Emphasis>

The effects of the availability of light (high, medium and low) and soil water (wet and dry) on morphological and physiological traits responsible for whole plant carbon gain and ramet biomass accumulation were examined in a splitter-type clonal herbaceous species Primula sieboldii, a spring plant inhabiting broad range of light environments including open grassland and oak forest understory. Growth experiments were conducted for three genets originated from natural microhabitats differing in light and soil water availability. Ramets of a genet from high light and wet microhabitat, which were grown in low light (relative photon flux density: R-PPFD of 5%) showed 41% less light-saturated photosynthetic rate, 50% less dark respiration rate and earlier defoliation than the ramets in high light (R-PPFD of 61%). The estimation of daily photosynthesis revealed that the light acclimation response in leaf gas exchange contributes to efficient carbon gain of whole plants, irrespective of experimental light conditions. Water stress increased root weight ratio, decreased ramet leaf area, petiole length and photosynthetic capacity. These morphological effects of water stress were larger in high and medium light regimes than in low light regime. The consequence of the above responses was recognized in the relative growth rate of the ramets. The relative growth rate of the ramets in high light with wet regime was four-fold of that in low light plus wet regime, and was 1.5-fold of that in high light plus dry regime. However, even in low light and/or dry regimes, ramets kept positive relative growth rates and produced gemma successfully. We could not detect significant variation in growth responses among genets. The high photosynthetic plasticity revealed in the present study should enable Primula sieboldii to inhabit in a broad range of light and soil water availability.     

Shade avoidance in the clonal herb Trifolium fragiferum/: a field study with experimentally manipulated vegetation height

In herbaceous canopies light availability can show high degrees of spatial variability in a vertical and also in a horizontal direction. Stoloniferous plants are hence likely to encounter differences in light availability during their ontogenetic development. Different mechanisms, such as petiole elongation, plasticity in internode length and branching, and an enhanced allocation to sexual reproduction have been suggested to represent viable shade-avoidance mechanisms for clonal plants.In a field experiment we tested the response of the stoloniferous herb Trifolium fragiferum L. to experimentally manipulated vegetation heights. Naturally occurring clonal fragments were exposed to four different vegetation heights ranging from 0 cm (high light availability created by clipping the surrounding natural vegetation at ground level) to 20 cm (natural shading in closed canopy). The growth and development of individual clones was followed for two months. At the end of the experiment above-ground plant parts were harvested. Growth-related and morphological parameters (e.g., petiole and internode length) as well as patterns of meristem utilization (i.e., flowering, branching) were recorded.Neither primary stolon growth and biomass accumulation nor branching and flowering were significantly affected by treatments. However, increased vegetation height resulted in a reduced number of secondary ramets and also had strong positive effects on petiole length, leading to marked changes in the architecture of plants growing in canopies of different heights. In addition, the average weight of individual ramets on the primary stolon was markedly higher in plants exposed to taller vegetation as compared to shorter vegetation.The results of this study suggest the occurrence of a trade-off between clonal expansion (i.e., secondary ramet production) and the average size of clonal offspring. If grown under higher vegetation plants invested more into the size of individual ramets, especially into elongating petioles, and less into the growth and development of lateral branches. Placing leaf laminae higher up in the canopy results in an enhanced light interception which has apparently buffered negative effects of increased vegetation height on whole-clone biomass. Plants grown under shorter vegetation invested more into lateral spread by producing more, but smaller ramets.     

不同生境中淫羊藿克隆构型和分株种群特征

为探求淫羊藿资源的保护策略,通过野外样地调查、室内测定和统计分析等方法,对林缘旷地、林缘和林下三种不同生境中淫羊藿(Epimedium brevicornum)克隆构型及其分株种群特征进行了研究,并初步分析环境因子对淫羊藿克隆构型和分株种群特征的影响.结果表明:分枝强度、间隔子长度、分枝高度、根状茎长度及分株种群密度在3种生境中差异显著,且与生境相对光强和土壤含水量有较强的回归关系.群落相对光强、土壤含水量在淫羊藿的克隆生长过程中起着重要作用.淫羊藿以地下根茎为克隆器官进行克隆生长,有着较强的克隆构型可塑性.结合克隆植物对资源的利用策略,讨论不同生境中淫羊藿克隆构型和分株种群特征可塑性的生态适应意义及淫羊藿保护机制.     

STRUCTURE AND DYNAMICS OF BAMBOOS IN THE WOLONG NATURAL RESERVE,CHINA

Culm (ramet) demography of two bamboos, Fargesia robusta and Bashania fangiana, was studied over 7 yr in southwest China to determine if: 1) ramet numbers remain relatively constant; 2) ramet mortality was constant; 3) density response was greater in spreading vs. compact clones; 4) extrinsic factors (e.g., predation, climate) influenced ramet dynamics; and 5) ramets that die are smaller than ones that live. Culm numbers remained relatively constant (±20%) in most plots, but culm numbers increased (>;20%) on some sites. Mortality of culms >;6 mo old over 7 yr was low and constant, but mortality of younger culms (≤6 mo) was high and variable due to predation. Spreading clones had greater recruitment response to density than compact clones, but predation and climate may have diminished or amplified the observed demographic patterns. Predators and perhaps climate influenced culm demography, and culms that died were smaller than remaining live culms. The demography of the two bamboos we studied is compared to other clonal plants, including bamboos, to identify common processes controlling ramet demography.     

辽东栎林内不同小生境下幼树植冠构型分析

以黄土高原黄龙山林区辽东栎林内3个小生境（林下、林隙、林缘）下辽东栎天然更新幼树为研究对象,采用典型抽样法对辽东栎幼树侧枝、叶片和树冠的空间分布状况以及生物量分配状况进行调查分析,探讨微生境与幼树植冠构型特征的关系,明确辽东栎幼树对不同小生境的适应策略,为栎林经营和林分结构优化提供理论依据。结果显示：（1）3种生境下辽东栎幼树构型发生了可塑性变化,林下幼树树冠层次比较单一,林隙与林缘的幼树树冠层次更加丰富。（2）由林下至林缘,幼树的树高、枝下高呈逐渐减小的趋势,而地径变化趋势与之相反;幼树的冠幅、树冠面积、树冠率呈先增加后减小的趋势,并且林下与林隙、林缘的差异显著;幼树的总体分枝率、逐步分枝率、枝径比呈先增加后减小的趋势。（3）3种生境下,幼树的一级枝的枝长、直径与倾角随着树高的增加而呈减小的趋势,但3种生境的差异不显著;林下一级枝主要分布在冠层中上部,而林隙与林缘一级枝主要分布在冠层中下、中上部。（4）由林下至林缘幼树叶长、叶宽、单叶面积和比叶面积逐渐降低,而单株叶数、叶总面积、叶面积指数呈先增大后减小趋势;与其他2种生境相比林下叶片分布趋于冠层上部。（5）幼树地上部分生物量中林下主干生物量占83%,枝和叶生物量只占17%;而林隙与林缘虽然各部位生物量有所差异但比例基本一致,其中主干占66%左右,枝和叶生物量占34%左右。研究表明,林隙生境下幼树的构型优于林缘和林下生境,在今后栎林的经营中,可以通过适当间伐来增加林隙数量,为森林更新和结构的优化提供有利条件。     

不同生境下刺五加种群构件生物量结构与生长规律

从构件水平对针阔混交林、蒙古栎林和次生杂木林3个生境的刺五加种群各功能构件的生物量结构及生物量比率与年龄之间的关系进行了定量分析. 结果表明,刺五加种群的个体生长和各构件生物量动态与环境条件关系密切.不同环境条件下,刺五加种群各功能构件生物量平均值具有相同的规律：茎构件>根茎构件>叶构件.3个生境中刺五加整体种群水平具有一定的相似性.在郁闭度为40%的蒙古栎林中,刺五加种群个体生物量和各构件生物量较针阔混交林和次生杂木林中大.不同生境下刺五加种群分株生物量的差异蕴涵着重要的生长调节和物质分配策略.在分株较小的幼龄个体以及郁闭度较大而不利于分株生长的次生杂木林中,分株优先建造叶器官,以保证充分的物质生产. 3个生境刺五加种群的叶、茎与分株的相对增重均具有相同的幂函数异速生长规律.     

Genet structure and determinants of clonal structure in a temperate deciduous woodland herb, Uvularia perfoliata

1 We used isozyme variation to examine the genet structure of Uvularia perfoliata patches in gap and closed canopy habitats in a temperate deciduous forest in Maryland, USA.
2 A large patch in a gap habitat was composed of a small number of widely spread genets with many ramets, and a large number of genets with more restricted distribution and few ramets. Genets with many ramets were patchily distributed at a metre scale. Analysis of genet structure on a scale of square centimetres, however, revealed that the genets were highly intermingled with no clear boundaries between them. The presence at both scales of sampling of many genets with unique multilocus genotypes indicated continuing genet recruitment within the population.
3 In the closed canopy habitat, the patches examined were each composed of a single unique multilocus genotype, suggesting that each had developed by asexual propagation following the establishment of a single genet.
4 The clonal structure of U. perfoliata patches in both gap and closed canopy habitats therefore appears to depend on recruitment patterns of genets. Populations in closed canopy habitats are characterized by a 'waiting' strategy, in which asexual ramet production maintains populations until genet recruitment by seed production can occur under the more optimal conditions associated with canopy gaps. Extended sampling suggests that the genetic diversity of U. perfoliata populations is primarily controlled by the disturbance regime of the forest canopy.