匍匐茎草本活血丹(Glechomalongituba)在不同养分条件下的克隆形态

通过人工施肥的方法,对活血丹无性系实验种群在不同养分供应情况下的形态特征进行了研究。研究结果表明：施肥可以显著改变克隆植株的形态,无怀系分株产生能力增强,个体生长旺盛;匍匐茎总长增长,生物量增大,节间短,匍匐茎较粗壮;相对叶面积小,叶片厚,叶柄较粗短。在未施肥条件下,克隆植株形态表现则正好相反。养分供应的差异对叶片的形态以及根重和根量的影响较其对隔离者和叶柄的特征的影响更大。     

土壤养分水平影响绢毛匍匐委陵菜匍匐茎生物量投资

为研究匍匐茎草本植物对基质养分供应水平的生物量分配格局的可塑性,在一盆栽实验中对绢毛匍匐委陵菜(Potentilla reptans var. sericophylla)进行了8种不同的养分处理。绢毛匍匐委陵菜植株生物量、匍匐茎数、分株数以及匍匐茎节间长在中等养分条件下最大。随土壤养分的降低,绢毛匍匐委陵菜对叶片和叶柄的生物量投资减小,而对根系的生物量投资增加。在中等养分条件下,绢毛匍匐委陵菜对匍匐茎的生物量投资倾向于最大,而在更高或更低的养分条件下倾向于减少。此生物量分配格局与de Kroon和Schieving的模型模拟结果相符合,结果表明在中等资源水平下增加对匍匐茎的生物量投资是克隆植物增加资源获取的对策之一。     

林下和林窗内绢毛匍匐委陵菜的克隆生长和克隆形态

 为了验证绢毛匍匐委陵菜(Potentilla reptans var. sericophylla)林窗和林下种群间的行为差异是完全由表型可塑性引起,还是局部分化的结果,将生长在北京东灵山油松（Pinus tabulaeformis）林林窗和林下的绢毛匍匐委陵菜,进行生境间的交互移植-重植野外生态实验。研究结果表明,实验植物的叶片长度、叶片宽度、叶柄长度和匍匐茎节间长度等克隆形态特征在两生境间无差异。两个来源的植株,其基株生物量、基株分株数和基株匍匐茎总长度等克隆生长特征在林下生境中都比在林窗生境中小,表现出显著的可塑性。所研究的克隆形态特征和克隆生长特征及其可塑性在不同生境来源的实验植物间没有差异。绢毛匍匐委陵菜克隆形态特征和克隆生长特征及其可塑性在林下和林窗生境间没有发生局部分化,林窗为其较适生境,克隆生长特征的可塑性对绢毛匍匐委陵菜利用生境异质性可能具有重要意义     

高寒草甸退化对短穗兔耳草克隆生长特征的影响

以调查统计和比较样地法研究了江河源区高寒草甸退化对典型匍匐茎植物短穗兔耳草克隆生长特征的影响。结果表明,退化草甸的植物群落结构、功能以及土壤特征发生了明显地变化,继而对短穗兔耳草无性系的克隆生长行为和形态特性产生了影响。高寒草甸退化后短穗兔耳草的匍匐茎有所增多,分支强度加大。退化草甸内短穗兔耳草的基株高度小于未退化草甸,根长大于未退化草甸,基株的叶片数目间没有明显差别。退化草甸内短穗兔耳草的分株高度显著小于未退化草甸,分株叶数明显多于未退化草甸,而根长尽管大于未退化草甸,但差异不显著。短穗兔耳草匍匐茎长度在未退化草甸内明显大于退化草甸,匍匐茎茎生叶数和匍匐茎粗度也在未退化草甸大于退化草甸。短穗兔耳草在未退化草甸用于克隆繁殖的能量投资比例高于退化草甸,其中未退化草甸内短穗兔耳草基株的干重比例略低于退化草甸,分株和匍匐茎的干重比例高于退化草甸。高寒草甸退化对短穗兔耳草克隆生长特征的这些影响,是其对高寒草甸退化导致的资源和生境差异的反应,也是对资源利用达到的最合理状态,是一种选择适应的结果。     

林缘和荒草坡不同草本层盖度小生境中积雪草的等级可塑性

为研究克隆植物在不同生境中对小生境的利用能力和对不同尺度异质性生境的等级可塑性及其格局差异,比较了林缘和荒草坡不同草本层盖度小生境中积雪草(Centella asiatica)分株种群、克隆片段和叶水平的形态和生理特征。在分株种群水平,林缘和荒草坡积雪草被测的分株种群特征差异均不显著,不同草本层盖度小生境间积雪草根冠比差异显著,即在盖度小的小生境中其根冠比显著大于盖度大的小生境。在克隆片段水平,林缘和荒草坡生境显著影响积雪草的根干重、花果干重、一级匍匐茎节间长和总匍匐茎节间长,而不同草本层盖度小生境则显著影响花果干重、根冠比、一级匍匐茎数、二级匍匐茎节间长和总匍匐茎节间长。荒草坡积雪草拥有更大的根干重和花果干重,但一级匍匐茎节间长和总匍匐茎节间长较短。草本层盖度小的小生境中积雪草的根冠比和一级匍匐茎数显著高于盖度大的小生境,而花果干重、二级匍匐茎节间长和总匍匐茎节间长则相反。在叶水平,荒草坡和草本层盖度大的小生境中,积雪草的叶片长、叶片宽和叶面积都较大,而草本层盖度大的小生境中积雪草叶柄长和比叶柄长都显著大于盖度小的小生境;林缘和荒草坡生境与不同草本层盖度小生境对4个叶绿素指标影响均不显著,林缘和荒草坡生境与不同草本层盖度小生境的交互作用对所有测量特征均无显著影响。上述结果表明,积雪草对不同生境和有草本层盖度差异的小生境的反应存在等级可塑性,但是二者的等级差异格局不同;等级反应格局的差异反映出积雪草对不同生境条件有不同的生态适应对策,且对生境小尺度异质性的可塑性强于对大尺度异质性的可塑性。     

匍匐茎草本金戴戴对盐分梯度的表型可塑性

 研究了匍匐茎型克隆草本金戴戴(Halerpestes ruthenica) 4种基株（基因型）对不同盐分处理（0,85.5, 171.0, 256.5和342.0 mM NaCl）的表型可塑性。随着盐分浓度的增加,实验植物与生长相关的性状指标 (如植株干重、总叶面积、分株数和总匍匐茎长度) 显著减小。植株干重、总叶面积和总匍匐茎长度具有显著的基株间差异。实验植物与形态相关的性状指标 (如平均叶柄长和根冠比) 对盐分梯度具有可塑性并具有显著的基株间差异;而其它形态指标 (如平均节间长、比节间长和比叶柄长)     

匍匐茎草本金戴戴对盐分梯度的表型可塑性

研究了匍匐茎型克隆草本金戴戴(Halerpestes ruthenica) 4种基株（基因型）对不同盐分处理（0,85.5, 171.0, 256.5和342.0 mM NaCl）的表型可塑性。随着盐分浓度的增加,实验植物与生长相关的性状指标 (如植株干重、总叶面积、分株数和总匍匐茎长度) 显著减小。植株干重、总叶面积和总匍匐茎长度具有显著的基株间差异。实验植物与形态相关的性状指标 (如平均叶柄长和根冠比) 对盐分梯度具有可塑性并具有显著的基株间差异;而其它形态指标 (如平均节间长、比节间长和比叶柄长)     

光照强度和基质养分对匍匐茎草本金戴戴克隆生长和克隆形态的影响

在深度遮光 (光照强度为高光条件的 6 .2 5% ,约为自然光照的 5.3% )或低养分条件下 ,金戴戴 (HalerpestesruthenicaOvcz.)生物量、初级分株叶面积、分株总数、匍匐茎总数和总长度均显著减小 ,而比节间长和比叶柄长显著增加。在低养分条件下 ,金戴戴匍匐茎平均节间长显著增加 ,而匍匐茎分枝强度和分株数显著减小。这些结果与克隆植物觅食模型相符合 ,表明当生长于异质性生境中 ,金戴戴可能通过以克隆生长和克隆形态的可塑性实现的觅养行为来增加对养分资源的摄取。在深度遮光条件下 ,金戴戴平均间隔子长度 (即平均节间长和平均叶柄长 )均显著减小。这一结果与以往实验中匍匐茎草本间隔子对中度和轻度遮光 (光照强度为高光条件的 1 3%～ 75% ,>1 0 %的自然光照 )的反应不同。这表明 ,在深度遮光条件下匍匐茎克隆植物可能不发生通过间隔子可塑性实现的觅光行为。光照强度和基质养分条件的交互作用对许多性状如总生物量、匍匐茎总数和总长度、二级和三级分株数、分株总数、初级分株叶面积以及分枝强度均有十分显著的效应。在高光条件下 ,基质养分对这些性状有十分显著的影响 ;而在低光条件下 ,基质养分条件对这些性状不产生影响或影响较小。这表明 ,光照强度影响金戴戴对基质养分的可塑性反应。在深度遮光     

光照强度和基质养分对匍匐茎草本金戴戴克隆生长和克隆形态的影响

在深度遮光(光照强度为高光条件的6.25%,约为自然光照的5.3%)或低养分条件下,金戴戴(Halerpestes ruthenica Ovcz.)生物量、初级分株叶面积、分株总数、匍匐茎总数和总长度均显著减小,而比节间长和比叶柄长显著增加.在低养分条件下,金戴戴匍匐茎平均节间长显著增加,而匍匐茎分枝强度和分株数显著减小.这些结果与克隆植物觅食模型相符合,表明当生长于异质性生境中,金戴戴可能通过以克隆生长和克隆形态的可塑性实现的觅养行为来增加对养分资源的摄取.在深度遮光条件下,金戴戴平均间隔子长度(即平均节间长和平均叶柄长)均显著减小.这一结果与以往实验中匍匐茎草本间隔子对中度和轻度遮光(光照强度为高光条件的13%～75%,>10%的自然光照)的反应不同.这表明,在深度遮光条件下匍匐茎克隆植物可能不发生通过间隔子可塑性实现的觅光行为.光照强度和基质养分条件的交互作用对许多性状如总生物量、匍匐茎总数和总长度、二级和三级分株数、分株总数、初级分株叶面积以及分枝强度均有十分显著的效应.在高光条件下,基质养分对这些性状有十分显著的影响;而在低光条件下,基质养分条件对这些性状不产生影响或影响较小.这表明,光照强度影响金戴戴对基质养分的可塑性反应.在深度遮光或低养分条件下,金戴戴可能通过减小匍匐茎节间粗度(增加比节间长)来增加或维持其相对长度,从而更有机会逃离资源丰度低的斑块.     

高寒草甸退化对鹅绒委陵菜克隆生长特征的影响

为了揭示高寒草甸典型匍匐茎克隆植物对不同生境的生态适应对策,验证生境适应假说,并为高寒草地的退化演替机理研究提供依据,以调查统计和比较样地法研究了江河源区高寒草甸退化对鹅绒委陵菜克隆生长特征的影响。研究表明,重度退化草甸的植物群落结构、功能以及土壤特征发生了明显的变化,继而对鹅绒委陵菜无性系的克隆生长行为和形态特性产生了影响。高寒草甸退化后鹅绒委陵菜的匍匐茎增多,分支强度加大。退化草甸内鹅绒委陵菜的基株高度小于未退化草甸,根长大于未退化草甸,基株的叶片数目问没有明显差别。退化草甸的分株高度显著小于未退化草甸,而分株叶数大于未退化草甸,根长小于未退化革甸且差异不显著。未退化草甸内鹅绒委陵菜无性系的问隔子长度、粗度和匍匐茎长度大于退化草甸,间隔子平均数目少于退化草甸,差异都不显著。随着鹅绒委陵菜无性系匍匐茎数目的增加,不论重度退化草甸与未退化草甸,用于鹅绒委陵菜克隆繁殖的能量投资也逐渐增加。鹅绒委陵菜在未退化草甸用于克隆繁殖的能量投资比例高于退化草甸,其中未退化草甸内鹅绒委陵菜基株的干重占无性系总生物量的比例略低于退化草甸,分株和匍匐茎的干重占无性系总生物量的比例高于退化草甸。高寒草甸退化对鹅绒委陵菜克隆生长特征的这些影响,与植物群落结构和功能的变化导致生境变异密切相关,是其对高寒草甸退化导致的资源和生境差异的反应,也是对资源利用达到的最合理状态,是一种选择适应的结果,有利于克隆繁殖潜力的发挥。同时为生境适应假说提供了又一例证。     

匍匐茎草本蛇莓对基质养分条件的克隆可塑性

A pot experiment with three levels of nutrient (N, P, K) supply was carried out to investigate clonal plasticity of stoloniferous herb Duchesnea indica Focke in response to nutrient availability. The plants had greater biomass at higher levels of nutrient availability. The root/shoot ratio of the plants changed with the nutrient availability in the following order: low level > high level > medium level. They had the largest biomass allocation to stolon at the medium level of nutrient availability. The biomass allocation to petiole did not respond to the treatments. The plants formed more stolons and ramets at the high and medium levels than the low level of nutrient availability. The petiole length, specific petiole weight（mg/cm）and stolon internode length of the plants did not respond to the treatments, while the specific stolon weight (mg/cm）of the plants was greater at the high and medium levels than the low level of nutrient availability. The results have been discussed in the context of adaptation of clonal plants to environmental heterogeneity.     

Plasticity in R/S ratio, morphology and fitness-related traits in response to reciprocal patchiness of light and nutrients in the stoloniferous herb, Glechoma longituba L

Clonal fragments of the stoloniferous herb Glechoma longituba were subjected to a complementary patchiness of light and soil nutrients including two spatially homogeneous treatments (SR–SR and IP–IP) and two spatially heterogeneous treatments (IP–SR and SR–IP). SR and IP indicate patches (shaded, rich) with low light intensity (shaded, S), high nutrient availability (rich, R) and patches (illuminated, poor) with high light intensity (illuminated, I) and low nutrient availability (poor, P), respectively. Plasticity of the species in root–shoot ratio, fitness-related traits (biomass, number of ramets and dry weight per ramet) and clonal morphological traits (length and specific length of stolon internodes, area and specific area of laminae, length and specific length of petioles) were experimentally examined. The aim is to understand adaptation of G. longituba to the environment with reciprocal patches of light and soil nutrients by plasticities both in root–shoot ratio and in (clonal) morphology. Our experiment revealed performance of the clonal fragments growing from patches with high light intensity and low soil nutrient availability into the adjacent opposite patches was increased in terms of the fitness-related characters. R/S ratio and clonal morphology were plastic. Meanwhile, the capture of light resource from the light-rich patches was enhanced while the capture of soil nutrients from either the nutrient-rich or the nutrient-poor patches was not. Analysis of cost and benefit disclosed positive effects of clonal integration on biomass production of ramets in the patches with low light intensity and high soil nutrient availability. These results suggest an existence of reciprocal translocation of assimilates and nutrients between the interconnected ramets. The reinforced performance of the clonal fragments seems to be related with specialization of clonal morphology in the species.     

高寒退化与未退化草甸土壤下匍匐茎鹅绒萎陵菜的克隆生长特征的比较

通过鹅绒萎陵菜的野外移栽试验,分析了在矮嵩草草甸土壤和退化草甸土壤下该种植物的克隆繁殖特征．结果表明,在母株产生的匍匈茎数量、母株高度和叶片大小在两类不同土壤下没有明显差异．母株叶片数、匍匈茎的粗度、匍匈茎长度以及间隔子的长度有较明显的差异．在退化草甸土壤中母株叶片较多,匍匈茎长度和间隔子长度明显较长,匍匈茎直径也明显较大．在退化草甸环境中,由于土壤养分相对较缺乏,鹅绒萎陵菜可能通过增加母株叶片的数量,尽可能多地积累光合产物,来保证匍匈茎的生长,匍匈茎也以增加粗度和增加长度来尽量增强其觅食能力．鹅绒萎陵菜在生物量投资分配上也表现出一定的差异．与退化草甸土壤环境相比,在未退化草甸土壤中,鹅绒委陵菜克隆母株和分株的生物量均明显偏向地下部分(根系)的投资,以期从土壤中吸收更多的养分,从而最终提高子株的成活率．     

Phenotypic Responses of a Stoloniferous Clonal Plant Buchloe dactyloides to Scale-Dependent Nutrient Heterogeneity

Clonal plants could modify phenotypic responses to nutrients heterogeneously distributed both in space and time by physiological integration. It will take times to do phenotypic responses to modifications which are various in different growth periods. An optimal phenotype is reached when there is a match between nutrient conditions and foraging ability. A single plantlet of Buchloe dactyloides with two stolons was transplanted into heterogeneous nutrient conditions. One stolon grew in homogeneous nutrient patch, while the other cultured in different scales of heterogeneous nutrient patches. As compared to the other nutrient treatment, heterogeneous nutrient treatments with small scale of 25×25 cm resulted in a higher biomass, and larger number of ramets, clumps and stolons in B. dactyloides at both genet and clonal fragment levels. Significant differences of number of ramets, clumps and stolons were detected at the rapid growth stage, but not in the early stage of the experiment. Foraging ability was more efficient in heterogeneous than in homogeneous nutrient conditions as assessed by higher root mass and root to shoot ratio. Different nutrient treatments did not prompt significant differences in internode and root length. Physiological integration significantly increased biomass, but did not influence other growth or morphological characters. These results suggest that physiological integration modifies phenotypic plasticity of B. dactyloides for efficient foraging of nutrients in heterogeneous nutrient conditions. These effects are more pronounced at genet and clonal fragment levels when the patch scale is 25×25 cm. Time is a key factor when phenotypic plasticity of B. dactyloides in heterogeneous nutrient conditions is examined.     

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.     

不同光照条件下聚花过路黄的克隆构型和分株种群特征

以都江堰灵岩山常绿阔叶林林下、林缘和林缘旷地3种不同光照环境中匍匐茎草本聚花过路黄(Lysimachla congestiflora)为对象,对其匍匐茎节间长、分枝强度、分枝角度、分株种群密度、分株种群生物量等指标进行了测定和分析．结果表明,聚花过路黄的匍匐茎节间长、分枝强度、分株种群密度和分株种群生物量在3种生境间差异显著．Kruskal Wallis检验表明,匍匐茎节间长度和分枝角度的频次分布在3种生境间差异显著．3种生境中匍匐茎节间长度分布偏斜度(skewness)的大小分别为：林缘旷地>林缘>林下．林缘旷地与林缘和林下生境在分枝角度、分株种群高度和分株种群根冠比差异显著,而且林缘旷地生境中分枝角度分布偏斜度最小．林缘和林下生境在分枝角度、分株种群高度和分株种群根冠比差异不显著．从林缘旷地、林缘到林下,聚花过路黄的克隆构型和分株种群特征发生了相应改变．结合克隆植物对资源的利用对策,讨论了不同生境中聚花过路黄克隆构型和分株种群特征可塑性的生态适应意义．     

Differential effects of water depth and sediment type on clonal growth of the submersed macrophyte <Emphasis Type="Italic">Vallisneria natans</Emphasis>

The response of clonal growth and ramet morphology to water depth (from 60 to 260 cm) and sediment type (sand versus organic clay) was investigated for the stoloniferous submersed macrophyte Vallisneria natans in an outdoor pond experiment. Results showed that water depth significantly affected clonal growth of V. natans in terms of clone weight, number of ramets, number of generations, clonal radius and stolon length. V. natans showed an optimal clonal growth at water depths of 110–160 cm, but at greater depths clonal growth was severely retarded. A high allometric effect was exhibited in ramet morphology. Along the sequentially produced ramet generations, ramet weight and plant height decreased while stolon length and the root:leaf weight ratio increased. When using ramet generations as covariate, sediment type rather than water depth more strongly affected the ramet characteristics. For plants grown in clay, ramet weight, ramet height and stolon length were greater, and plants exhibited lower root:leaf weight ratio. These data suggest that water depth and sediment type have differential effects on clonal growth of V. natans: Water depth appears primarily to affect numerical increase in ramets and spatial spread, whereas sediment type mainly affects biomass accumulation and biomass allocation. Handling editor: S. M. Thomaz