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.     

贵州雷公山秃杉林不同林冠环境下箭竹分株种群结构特征

通过对贵州雷公山秃杉林的4种林冠环境,即林下(FU)、中林窗(MG)、大林窗(LG)和林缘旷地(FEW)内的箭竹的分株种群结构（包括株高、基径、生物量、叶和分株数等）和年龄结构进行较系统对比研究。结果显示:（1)从林下→中林窗→大林窗,箭竹分株种群的株高(h)、基径（bd)与生物量显著增加。（2)在4种林窗环境中,箭竹各构件的生物量的百分比发生相应变化,枝和叶片生物量的百分比沿林下→中林窗→大林窗→林缘旷地的顺序显著减小,在林缘旷地,地下茎、粗根（d＞0.5 mm)和细根（d≤0.5 mm)生物量的百分比显著高于其他3种林冠环境。（3)箭竹分株单位叶面积叶重沿林下→中林窗→大林窗→林缘旷地的顺序显著增大,单叶生物量与叶面积以中林窗最大,林缘旷地次之,二者与大林窗或林下差异显著;大林窗的单株叶片数显著高于其余3种林冠环境。（4)在4种林冠环境中,箭竹分株种群死亡率以林下最低,但各种群平均年龄间无显著差异。研究表明,箭竹分株种群对林冠环境变化的响应主要体现在形态与生物量的分配上,而非种群的年龄。     

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。     

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。     

常绿阔叶林林冠环境对栲幼苗建成的影响

植物幼苗建成阶段是决定种群自然更新的关键生活史阶段。研究林冠环境对常绿阔叶林优势种幼苗建成阶段的影响对该类森林的恢复和管理具有重要意义。2014-2016年, 该研究在重庆市缙云山国家级自然保护区的常绿阔叶林的不同林冠环境(大林窗: >150 m ², 中林窗: 100-150 m ², 小林窗: 50-100 m ², 对照: 林下)下进行栲(Castanopsis fargesii)种子野外播种实验, 并对栲幼苗命运和生长情况进行了3年的持续监测。结果表明: (1)栲幼苗出土时间从7月持续到12月, 出苗时间较长, 大林窗对幼苗出土具有延迟作用; (2)栲种子野外平均萌发率为(62.8 ± 2.0)%, 第3个生长季(2016年)末幼苗平均存活率为(65.1 ± 2.2)%, 枯萎是栲幼苗死亡的主要原因; (3)林冠环境对栲种子萌发率及第1个生长季(2014年)末的幼苗存活率无显著影响, 对第2个(2015年)和第3个生长季末的幼苗存活率具有显著影响; (4)林冠环境在第1个生长季对幼苗生长无明显影响, 但在第2个和第3个生长季具有显著影响, 大、中林窗中幼苗总生物量、株高、基径、根长和叶片数显著高于林下, 比叶面积显著低于林下; (5) 3个生长季内, 4类林冠条件下栲幼苗的叶质量比和茎质量比升高, 根质量比和根冠比降低, 并且从第2个生长季开始大林窗中栲幼苗的叶质量比显著高于林下, 根质量比和根冠比显著低于林下。栲幼苗早期的存活和生长依赖种子储存的能量, 受林冠条件影响较弱, 后期则依赖光合作用, 受林冠条件影响较强, 从整个幼苗建成过程看, 大、中林窗更有利于栲幼苗定居。     

常绿阔叶林林冠环境对栲幼苗建成的影响

植物幼苗建成阶段是决定种群自然更新的关键生活史阶段。研究林冠环境对常绿阔叶林优势种幼苗建成阶段的影响对该类森林的恢复和管理具有重要意义。2014–2016年,该研究在重庆市缙云山国家级自然保护区的常绿阔叶林的不同林冠环境(大林窗:150 m~2,中林窗:100–150 m~2,小林窗:50–100 m~2,对照:林下)下进行栲(Castanopsis fargesii)种子野外播种实验,并对栲幼苗命运和生长情况进行了3年的持续监测。结果表明:(1)栲幼苗出土时间从7月持续到12月,出苗时间较长,大林窗对幼苗出土具有延迟作用;(2)栲种子野外平均萌发率为(62.8±2.0)%,第3个生长季(2016年)末幼苗平均存活率为(65.1±2.2)%,枯萎是栲幼苗死亡的主要原因;(3)林冠环境对栲种子萌发率及第1个生长季(2014年)末的幼苗存活率无显著影响,对第2个(2015年)和第3个生长季末的幼苗存活率具有显著影响;(4)林冠环境在第1个生长季对幼苗生长无明显影响,但在第2个和第3个生长季具有显著影响,大、中林窗中幼苗总生物量、株高、基径、根长和叶片数显著高于林下,比叶面积显著低于林下;(5) 3个生长季内, 4类林冠条件下栲幼苗的叶质量比和茎质量比升高,根质量比和根冠比降低,并且从第2个生长季开始大林窗中栲幼苗的叶质量比显著高于林下,根质量比和根冠比显著低于林下。栲幼苗早期的存活和生长依赖种子储存的能量,受林冠条件影响较弱,后期则依赖光合作用,受林冠条件影响较强,从整个幼苗建成过程看,大、中林窗更有利于栲幼苗定居。     

慈竹母株大小对克隆生长的影响

 克隆生长在慈竹种群更新中占有重要地位。本文以基径和生物量两个指标同时衡量慈竹(Neosinocalamus affinis)的母株大小,并分析了母株大小对新竹笋产生、竹笋大小和根茎大小的影响。结果表明：1）慈竹的成竹基径与一级母株基径、二级母株基径以及与2龄和3龄母株平均单株生物量呈正相关,但是成竹基径随一级母株基径增加比随二级母株基径增加而增加的速率快得多;2）慈竹母株大小与各样方中出笋数、活笋数和死笋数无显著的正相关;3）发笋母株的基径显著大于未发笋母株,但发一个笋母株与发两个以上笋母株的基径无显著差异;4）母株基径、2龄和3龄母株平均单株生物量都与根茎的长度和粗度呈显著正相关,且竹笋基径与根茎粗度也呈显著的正相关。母株大小对慈竹竹笋的影响是通过影响根茎实现的。总之,慈竹的克隆生长在个体水平受母株大小制约。     

小径竹对亚高山暗针叶林优势 树种林窗更新的影响

在亚高山岷江冷杉林中选取面积≤50m2,50~150m2,>150m2的林窗,每种类型内均包含3种小径竹盖度(≤20%、20%~50%、>50%),共调查林窗9个,并调查包含这3种小径竹盖度的三块林下对照样地,研究了该类森林林窗更新与小径竹生长的关系。结果表明:(1)无论林窗大小,林窗内的更新幼苗数量都比林下的多,林窗更新是岷江冷杉群落更新的主要途径;(2)所选林窗均为发育早期,林窗对更新树种的种类组成和数量的影响主要表现在幼苗上。糙皮桦幼树及幼苗数量随林窗面积的增加而急速增加,它的更新更需要较大的林窗;(3)不同小径竹盖度下幼苗的密度呈现显著性变化,小径竹的生长明显抑制了森林幼苗的更新及填充的进程;(4)华西箭竹的分散程度随林窗面积的增大而降低,而平均高度和基径则有增加的趋势。     

亚高山暗针叶林林缘华西箭竹对岷江冷杉幼苗更新的影响

大量研究表明在箭竹-冷云杉林下,竹类对冷杉的影响大于其它树种。在卧龙自然保护区岷江冷杉(Abies faxoniana)林下,华西箭竹(Fargesia nitida)为灌木层优势种。为了进一步了解箭竹对岷江冷杉幼苗更新的影响,该文采用样带法在四川省卧龙自然保护区亚高山岷江冷杉林林缘,针对华西箭竹和岷江冷杉幼苗展开调查研究。以竹丛覆盖区域为边界,分别向竹丛外2 m (带1)、1 m (带2),向竹丛内1 m (带3)、2 m (带4)、3 m (带5)做5条长30 m的相邻样带,每条样带划分为30个连续的1 m×1 m小样方。对岷江冷杉幼苗数量动态、种群结构、生长与生物量的研究表明: 1)离竹丛越远,大年龄岷江冷杉幼苗数量越多;岷江冷杉幼苗转化率(下一径级苗木数量与前一径级苗木数量之比)随年龄增加而减小,随离竹丛距离增大而增大。2)苗木高度与径级结构相似,随深入竹丛,幼苗数量峰值的径级和高度级分布从较大径级(Ⅲ)向小径级(Ⅰ)变化。3)华西箭竹的存在抑制了岷江冷杉小径级幼苗(Ⅰ)的高生长,基径与高度的回归也表明离竹丛越远,高生长相对更大;但一定密度华西箭竹(带4)增大了大径级(Ⅲ)岷江冷杉幼苗冠幅的扩展。4)岷江冷杉幼苗总生物量随华西箭竹密度的增高而减小,华西箭竹的大量存在抑制岷江冷杉幼苗对地下生物量的分配。总的看来,一定密度华西箭竹环境下岷江冷杉幼苗的生长和更新较好,华西箭竹大量分布的区域,岷江冷杉幼苗的更新和生长受到抑制,可能会影响岷江冷杉种群的扩散以及群落的发展。     

不同土壤雷竹盆栽苗地下茎分枝数量特征及其分布格局

为了探究竹子分株系统构建过程及其与人工经营的关系,本文对雷竹(Phyllostachys praecox C. D. Chu et C. S. Chao 'Prevernalis’)不同年龄母竹、不同覆盖年限竹林进行土壤盆栽实验,比较了各处理竹苗地下茎分枝生长差异。结果显示：在盆栽竹苗分株系统构建过程中,地下茎以竹鞭分枝为主;2年生母竹盆栽苗地下茎分枝数量普遍高于1年生盆栽苗,且地下茎分枝表现出随竹林土壤覆盖年限增加而减少的趋势;盆栽苗地下茎分枝主要分布于第Ⅱ、Ⅲ、Ⅳ分枝的鞭中位置;1年生母竹盆栽苗地下茎分枝以第Ⅱ分枝级别的鞭中、鞭梢部位为多,而2年生母竹盆栽苗则以第Ⅲ分枝级别的鞭中、鞭梢部位为多;随土壤覆盖年限增加,地下茎分枝偏向分布于较为靠前的分枝级别。研究发现,母竹盆栽苗分株系统的构建主要采取了扩大地下分枝的策略,其2年生母竹与竹鞭中部着生侧芽的分枝生长对分株系统拓展贡献率较大;竹林土壤覆盖时间越久越不利于地下茎分枝。由于竹子分株系统具有时空拓展性,其地下茎分枝生长特征尚需持续观察。     

Clonal plasticity in response to rhizome severing and heterogeneous resource supply in the rhizomatous grass Psammochloa villosa in an Inner Mongolian dune, China

In a field experiment, Psammochloa villosa plants were subjected to rhizome severing. Severing rhizomes reduced growth in the young, detached rhizome segments compared to the controls in terms of all measured clonal growth-related characters, i.e. number of rhizomes and shoots, total rhizome length and total number of rhizome nodes. In a container experiment, the control ramets received uniform water and nutrient supply but in heterogeneous treatments high and low levels of water and nutrient supply, respectively were established. The number of ramets, total rhizome length, dry weight per ramet and biomass allocation to the rhizome had higher values at high water and nutrient supply, while spacer length (length of rhizome between shoots) and rhizome internode length were not affected. The local response of ramets given low water supply was enhanced due to connection to a well watered parent ramet in terms of number of ramets, total rhizome length and dry weight per ramet. A remote effect was not observed in the other treatments or in the other measured characters.     

Classifying clonal growth forms based on vegetative mobility and ramet longevity: a whole community analysis

We measured rhizome branching, clonal mobility, and ramet longevity of 98 meadow plant species. A cluster analysis applied to this dataset revealed nine clonal growth types that differ mainly by the ramet lifespan and vegetative mobility. Then we compared the abundance of these groups of clonal species between the three following plant communities: (1) open, (2) restored and (3) overgrown wooded meadows in the Laelatu-Nehatu-Puhtu Nature Reserve, Estonia. This is the first study where the quantitative values of belowground clonal traits have been measured for all species of a species-rich community. We show that species with annual ramets and with a low vegetative mobility were most abundant in open grasslands. The relative abundance of perennial species with annual ramets was positively correlated with shoot density and species diversity, indicating that high ramet turnover rates combined with a high genet longevity can positively affect species coexistence in meadow communities. Hence, this study provides evidence for the fact that the average values of clonal life-history parameters differ between these communities. Herb communities under forest canopy consist, in average, of species with ramets that live longer and are clonally more mobile than in the communities of open sites.     

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.     

Clonal plant allocation to daughter ramets is a simple function of parent size across species and nutrient levels

The evolution of clonal growth is a widespread phenomenon among plant species, characterized by the production of genetically identical clonal fragments (ramets) via rhizomes or stolons that form an interconnected clonal organism (genet). Clonal plant species are known to differ in their investment into ramet production, and exhibit considerable variation in ramet morphology both within and among species. While patterns of resource allocation are thought to be linked to a number of plant characteristics, many analyses are limited by uncertainty in how clonal plants determine the morphology and resources allocated to new ramets. In this study, we attempted to discern what aspects of parent ramets best predicted resource allocation to new daughter ramets, and the relationship between resource allocation and daughter ramet rhizome morphology. We grew two sedge species, Schoenoplectus tabernaemontani and Eleocharis elliptica, in a greenhouse under two levels of fertilizer addition. By harvesting daughter ramets that had initiated stem production, yet remained aphotosynthetic, we were able to isolate parental investment into non-independent daughter ramets at a point where daughter ramet spacer length became fixed. Our results indicate that parent ramets allocated a non-linear proportion of parent rhizome biomass to the production of daughter ramets. Moreover, this relationship was unaffected by environmental nutrient availability. Daughter ramet biomass, in turn, was strongly correlated with daughter ramet spacer length. These observations shed light on key processes governing clonal growth in plants, and their potential application in unifying allocational and morphological perspectives to explore the fitness implications of variability in clonal growth.     

Clonal integration and its ecological significance in Hedysarum laeve, a rhizomatous shrub in Mu Us Sandland

Hedysarum laeve, a rhizomatous clonal half-shrub, commonly dominates in inland dunes in semiarid areas of northern China. This species propagates vegetatively by the extension of horizontal rhizomes resulting in programmed reiteration of apical and/or axillary meristems. In this study, ¹⁴C labeling and experimental defoliation were employed to test the photosynthate translocation within the interconnected parent–daughter ramet pairs. A proportion of ¹⁴C-photosynthates was transported from the parent ramet into the daughter ramet, the roots of the daughter ramet, and the rhizome; these three components showed more than 70% sink activity after 24-h translocation. On the other hand, the basipetal translocation (from daughter ramet into parent ramet) was relatively small with sink activity of less than 5%, but sink activity of the rhizome exceeded 10%. Defoliation had an influence on the photosynthate translocation between parent and daughter ramets. The intact parent ramets significantly increased their ¹⁴C-photosynthate translocation into defoliated daughter ramets when compared to intact daughter ramets. The daughter ramets transported significantly more ¹⁴C-photosynthates to the defoliated parent ramets than to the intact parent ramets. A portion of ¹⁴C-photosynthates was transported into the rhizome from both parent and daughter ramets, indicating that the rhizome is supported by both ramets for photosynthates. The clonal integration between ramets of the species through rhizome connection may confer benefit both to the ramets and the genet in adverse environments. Received: April 12, 2001 / Accepted: November 26, 2001     

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.     

Clonal integration and its ecological significance in Hedysarum laeve,a rhizomatous shrub in Mu Us Sandland

Hedysarum laeve, a rhizomatous clonal half-shrub, commonly dominates in inland dunes in semiarid areas of northern China. This species propagates vegetatively by the extension of horizontal rhizomes resulting in programmed reiteration of apical and/or axillary meristems. In this study, (14)C labeling and experimental defoliation were employed to test the photosynthate translocation within the interconnected parent-daughter ramet pairs. A proportion of (14)C-photosynthates was transported from the parent ramet into the daughter ramet, the roots of the daughter ramet, and the rhizome; these three components showed more than 70% sink activity after 24-h translocation. On the other hand, the basipetal translocation (from daughter ramet into parent ramet) was relatively small with sink activity of less than 5%, but sink activity of the rhizome exceeded 10%. Defoliation had an influence on the photosynthate translocation between parent and daughter ramets. The intact parent ramets significantly increased their (14)C-photosynthate translocation into defoliated daughter ramets when compared to intact daughter ramets. The daughter ramets transported significantly more (14)C-photosynthates to the defoliated parent ramets than to the intact parent ramets. A portion of (14)C-photosynthates was transported into the rhizome from both parent and daughter ramets, indicating that the rhizome is supported by both ramets for photosynthates. The clonal integration between ramets of the species through rhizome connection may confer benefit both to the ramets and the genet in adverse environments.