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 condi-tions on the growth and morphological characteristics of F. nitida, and the adaptive responses of F. nitida to dif-ferent 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 clus-ter 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. More-over, the changes in plasticity enable the plants to optimize their light usage efficiency to promote growth and increase access to resources available in heteerogeneous light environoments.     

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。     

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.     

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.     

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.     

Sexual dimorphism in clonal growth forms and ramet distribution patterns in <Emphasis Type="Italic">Rumex acetosella</Emphasis> (Polygonaceae)

We investigated clonal traits in the dioecious herb Rumex acetosella to characterize sexual dimorphism in clonal forms and to correlate below-ground clonal patterns and above-ground ramet distributions. We recorded creeping root length, branching patterns, ramet and clump (caespitose ramets from the same position on the root) sprouting patterns, and biomass allocations in three females and males. We also estimated the patch size of flowering ramets within a quadrat. No sexual dimorphism was detected in the frequencies of branches and flowering ramets per root length. Male plants allocated proportionally more biomass to below-ground organs. Total root length did not differ between the sexes. Females sprouted more clumps with fewer flowering ramets per root length than males, which sprouted fewer clumps with more flowering ramets, which meant that clump sprouting patterns were phalanx-like in females and guerrilla-like in males. Flowering ramets were aggregately distributed in both females and males and patch sizes were similar between sexes, indicating that the spreader propagations were not found in the guerrilla-like males. We assumed that sexual dimorphism occurred in response to physiological integration for higher reproductive effort in females.     

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

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 Fargesla nitida （Mltford） 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 （ill） 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.     

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.     

Flowering in grassland predicted by CO2 and resource effects on species aboveground biomass

Continuing enrichment of atmospheric CO₂ may change plant community composition, in part by altering the availability of other limiting resources including soil water, nutrients, or light. The combined effects of CO₂ enrichment and altered resource availability on species flowering remain poorly understood. We quantified flowering culm and ramet production and biomass allocation to flowering culms/ramets for 10 years in C₄‐dominated grassland communities on contrasting soils along a CO₂ concentration gradient spanning pre‐industrial to expected mid‐21st century levels (250–500 μl/L). CO₂ enrichment explained up to 77% of the variation in flowering culm count across soils for three of the five species, and was correlated with flowering culm count on at least one soil for four of five species. In contrast, allocation to flowering culms was only weakly correlated with CO₂ enrichment for two species. Flowering culm counts were strongly correlated with species aboveground biomass (AGB; R² = .34–.74), a measure of species abundance. CO₂ enrichment also increased soil moisture and decreased light levels within the canopy but did not affect soil inorganic nitrogen availability. Structural equation models fit across the soils suggested species‐specific controls on flowering in two general forms: (1) CO₂ effects on flowering culm count mediated by canopy light level and relative species AGB (species AGB/total AGB) or by soil moisture effects on flowering culm count; (2) effects of canopy light level or soil inorganic nitrogen on flowering and/or relative species AGB, but with no significant CO₂ effect. Understanding the heterogeneity in species responses to CO₂ enrichment in plant communities across soils in edaphically variable landscapes is critical to predict CO₂ effects on flowering and other plant fitness components, and species potential to adapt to future environmental changes.     

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。     

Ecological benefits of integration of Calamagrostis epigejos ramets under field conditions

We established two independent experiments to estimate the ecological consequences of artificial severing on ramets of the competitively strong perennial grass Calamagrostis epigejos. We compared the responses of mature ramets of different size growing in different environments. Finally, we tested whether the response of young ramets to the severing depends on the density of surrounding vegetation.Severing decreased biomass and number of rhizomes of young ramets but did not affect their competitive tolerance. It decreased survival of mature ramets (probably due to traumata caused by cutting attached rhizomes) but did not influence total biomass of survived ramets. The response of total biomass of mature ramets to severing changed with size of the ramets. Further, biomass allocation to rhizomes changed differently after ramet severing in the two environments.The results suggest that field grown young ramets of Calamagrostis benefit from physiological integration. In contrast, mature ramets seem to be independent units according to the resource economy. Ecological benefits of integration depended on size of ramet clumps: ramets in clump had larger survival probability than control ramets. They also depended on environmental conditions: severing increased formation of new rhizomes at a sand dune subsrate, but it had a negative effect on plants in the forest experimental site. This intra-specific variation should be taken into account when trying to explain ecological patterns of integration benefits of clonal plants.     

Internal heterogeneity of ramet and flower densities of <Emphasis Type="Italic">Typha angustifolia</Emphasis> near the boundary of the stand

Fifteen relatively narrow stands of Typha angustifolia, including both stands bounded to open water and ridged boundary, were investigated in terms of the distributions of ramet density as well as the fraction of flowering ramets. The ramet density, fraction of flowering ramet and morphological characteristics of ramets were measured continuously from one side of the stand to the other, together with depth and the nutrient concentration of pore water. The rhizome biomass was also measured at the periphery and at the center of the stand. The results indicated that the shoot density declined gradually from the edge to the center, however, the fraction of the flowering ramets were significantly higher at the edge while it rapidly declined away from the edge. Statistical analyses indicate that the ramet density depends on either depth or nutrient level of the pore water; however, the distance from the edge was the most probable factor to reduce the fraction of the flowering ramets. The rhizome biomass was mostly larger at the periphery than at the center, likely because the accumulation of the horizontal rhizomes as extension was blocked by the boundary. High flowering fraction near the periphery was probably induced by the hormonal change caused by the blockage. However, a large fraction of ramets flowered simultaneously with large rhizome biomass at the periphery, likely because of the preferable light climates compared with inside the stand.     

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.     

Bamboo,fire and flood: consequences of disturbance for the vegetative growth of a clumping,clonal plant

The clumping growth form of tropical bamboos suggests a consolidator strategy in the face of intense competition rather than an ability to exploit disturbance. We investigated the annual growth and culm demographic responses to disturbance by fire and flood of vegetatively mature clumps of a riparian stand of Bambusa arnhemica. Linear-mixed models were employed to control for the non-independence of culm fates within clumps and clump growth among years. As the stand was monodominant and the species is gregariously monocarpic, responses can be interpreted independently of interspecific competition and trade-offs with sexual reproduction. Disturbance depressed clump growth but the culm demographic responses to fire and flood were quite different. Few culms were destroyed during the fire but damage depressed their subsequent survival, leading to declines in clump basal area. Fire also triggered the release of ramet buds, but the additional recruits were small and short-lived. Prolonged and early flooding in one of the study years depressed culm recruitment in clumps low on the lagoon bank but there was possible compensatory recruitment in the following year. There was no convincing evidence that sparse clumps recruited culms better than dense clumps, though culm recruitment was negatively correlated with culm survival. Fire and prolonged flooding are inevitable elements of the environment occupied by B. arnhemica, but we interpret the species’ response as persisting in the face of disturbance rather than exploiting the opportunities created by it.     

Spacer elongation and plagiotropic growth are the primary clonal strategies used by Vallisneria spiralis to acclimate to sedimentation

The aim of this study is to identify how submerged macrophyte Vallisneria spiralis acclimate to sedimentation by investigating the growth, biomass allocation and clonal characteristics in a greenhouse experiment of 30 days. Experimental treatments combined two sediment types (mud and sand) with four sedimentation depths (0, 2, 4 and 8 cm) in a factorial design. Biomass accumulation (0.98–1.33 versus 0.36 g per plant) and relative growth rate (RGR, 0.082–0.093 versus 0.046 g g⁻¹ day⁻¹) decreased only in the 8 cm sand treatment. Neither sedimentation depth nor sediment type influenced biomass allocation. The ratio of spacer length to biomass was significantly higher in the 8 cm sand (20.4 cm g⁻¹) than in other treatments (6.0–8.5 cm g⁻¹). Branching angles and the depths between ramet basal and sediment surface were only affected by sedimentation depth. Clonal ramets developed nearly vertical branching angles (ranged from 78° to 101°) in the 0 cm sedimentation treatment, but the angles of treated plants decreased at the initial 3–5 ramets (ranged from 68° to 78° at the first ramet level), then remained a relatively constant value (about 90°) in the following spacers. These data indicate that plagiotropic stolons were formed to project the ramets to sediment surface and to escape sedimentation stress primarily by elongating spacer length and decreasing branching angle, rather than by adjusting biomass allocation.     

扎龙湿地芦苇分株生态可塑性及其对土壤因子的响应

扎龙湿地的芦苇既可形成大面积的单优群落,也可形成不同群落斑块。采用大样本抽样调查与统计分析方法,对湿地内水生生境、湿生生境、旱生生境和盐碱生境芦苇种群分株高度和生物量进行比较。结果表明,6—10月份,4个生境芦苇种群分株高度及生物量均以水生生境最高,盐碱生境最低,水生生境株高为盐碱生境的1.5—2.3倍,分株生物量为2.0—5.1倍,生境间的差异性以及差异序位均相对稳定。4个生境株高生境间变异系数(19.45%—31.56%)均高于生境内变异系数(8.07%—17.61%),分株高度在生境间的可塑性更大;分株生物量中水生生境、湿生生境和盐碱生境3个生境间的变异系数(33.43%—55.61%)均低于生境内变异系数(44.85%—79.82%),分株生物量在生境内的可塑性更大。不同生境条件下芦苇种群分株,在生长和生产上均存在较大的生态可塑性,表现出明显的环境效应,其中土壤含水量是该地区芦苇分株生态可塑性变异的主要驱动因子(R0.80),为正向驱动。     

Impacts of forest gaps on soil properties and processes in old growth northern hardwood-hemlock forests

We examined the influence of treefall gaps on soil properties and processes in old growth northern hardwood-hemlock forests in the upper Great Lakes region, USA. We found significantly greater solar radiation, soil moisture contents and soil temperatures in gaps compared to adjacent closed canopy plots. Gaps had significantly less exchangeable base cations (K, Ca, and Mg) compared to forest plots in the upper mineral soil (0–25 cm). Gaps also had significantly more dissolved organic N and extractable nitrate at depth (25–50 cm), indicating increased nutrient leaching in gaps. In-situ N mineralization was significantly greater in gaps and edge plots compared to forest plots. We found significantly greater potential N mineralization (measured in the laboratory at 25°C and 40% water holding capacity) in forest compared to gap plots. Microbial biomass N was significantly greater (ca. two-fold) in the gap edge compared to both gaps and closed forest. Using principal component analyses we found that edge plots were positively correlated with all principal components, indicating increased in-situ and potential N mineralization, microbial biomass N, soil NO₃⁻ and NH₄⁺, and soil organic matter. The gap edge may be a region of optimal microclimate and substrate to enhance microbial biomass and activity within these forest ecosystems. Responsible Editor: Bernard Nicolardet     

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.