资源异质性环境中三种匍匐茎草本植物克隆内资源共享和分株功能特化

研究了3种来自中国北方林下、草地和碱化草甸匍匐茎型克隆草本植物绢毛匍匐委陵菜 (Potentilla reptans L. var. sericophylla Franch.)、鹅绒委陵菜 (P. anserina L.) 和金戴戴 (Halerpestes ruthenica (Jacq.) Qvcz.) 对由高光照低养分斑块和低光照高养分斑块组成的资源交互斑块性生境的适应性对策.当生长于高光照低养分条件下分株 (HL分株) 与生长于低光照高养分条件下分株 (LH分株) 之间的匍匐茎连接时, 3种克隆植物HL分株、LH分株以及整个分株对系统 (HL分株 + LH分株) 的生物量均得到显著提高.同时, LH分株根冠比显著增加, 而HL分株根冠比显著下降.这表明, 当互连分株置于由低光照高养分斑块和高光照低养分斑块组成的异质性环境中时, 3种植物克隆分株均发生了环境诱导的功能特化.克隆内资源共享以及克隆内不同分株的功能特化有利于整个分株系统对局部丰富资源的获取, 从而能够缓解资源交互斑块性生境对克隆植物的不利影响.     

资源异质性环境中三种葡匐茎草本植物克隆内资源共享和分株功能特化

研究了3种来自中国北方林下、草地和碱化草甸匍匐茎型克隆草本植物绢毛匍匐委陵菜（Potentilla reptans L.var.sericophylla Franch.)、鹅绒委陵菜（P.anserina L.)和金戴戴（Halerpestes ruthenica (Jacq.)Qvcz.)对由高光照低养分斑块和低光照高养分斑块组成的资源交互斑块性生境的适应性对策。当生长于高照低养分条件下分株（HL分株）与生长于低光照高养分条件下分株（LH分株）之间的匍匐茎连接时,3种克隆植物HL分株、LH分株以及整个分株对系统（HL分株＋LH分株）的生物量均得到显提高,同时,HL分株根冠比显增加,而HL分株根冠比显下降。这表明,当互连分株置于由低光照高养分斑块和高光照低养分斑块组成的异质性环境中时,3种植物克隆分株均发生了环境诱导的功能特化。克隆内资源共享以及克隆内不同分株的功能特化有利于整个分株系统对局部丰富资源的获取,从而能够缓解资源交互斑块性生境对克隆植物的不利影响。     

资源交互斑块性生境中两种不同分枝型匍匐茎植物的克隆内分工

以青藏高原和四川盆地过渡带两种不同分枝型匍匐茎植物野草莓 (Fragaria vesca)和过路黄 (Lysimachia christinae)为对象 ,研究它们在高光照低养分斑块和低光照高养分斑块组成的资源交互斑块性生境中的克隆内分工。结果显示 ,与资源的空间同质性处理 (I)和 (II)相比 ,资源的空间异质性处理 (III)和 (IV)中野草莓和过落黄的近端、远端和整个克隆片段的生物量和分株数均获得显著增加。生长在低光高养条件下的远端分株 ,若与高光低养的近端分株相连 ,相比连接到低光高养的近端分株 ,它们分配更多的生物量到地下部分 ;生长在高光低养条件下的远端分株 ,若与低光高养的近端分株相连 ,相比连接到高光低养的近端分株 ,它们分配更多的生物量到地上部分 ;生长在高光低养条件下的近端分株 ,若与低光高养的远端分株相连 ,相比连接到高光低养的远端分株 ,它们分配更多的生物量到地上部分。实验结果表明 ,资源交互斑块性生境中野草莓和过路黄均发生了克隆内分工。通过克隆内分工 ,克隆植物能有效的利用异质性分布的资源 ,缓解资源交互斑块性分布对克隆植物生长的不利影响     

匍匐茎草本绢毛匍匐委陵菜对局部遮荫的克隆可塑性

采自林窗和林内生境的绢毛匍匐委陵菜 (PotentillareptansL .var.sericophyllaFranch)“分株对”(即由一匍匐茎节间相连着的两个分株 ,其一为“目标分株” ,另一为“相连分株”)在一户外实验中经历了全不遮荫、全部遮荫和局部遮荫处理。该植物的基株生物量、匍匐茎总长度、分株数、匍匐茎比节间重、叶柄长、比叶柄重在遮荫条件下较小。匍匐茎节间长度没有对遮荫处理发生反应。在局部遮荫处理 ,遮荫斑块的分株的叶柄长度由于连着未遮荫斑块中分株而变得更长。这种克隆整合对克隆形态可塑性的修饰作用只在林窗生境来源的实验植物中观察到。其他克隆生长和克隆形态特征的可塑性在不同生境来源的实验植物间没有差异。     

PATCHY CONTRAST OF HABITAT AFFECTS INTRACLONAL DIVISION OF LABOR OF POTENTILLA ANSERINA

 相互连接的克隆植物分株分别处于资源互补性的不同斑块时, 将可能发生形态结构的特化, 以更有效地吸收利用所处斑块中丰富的资源, 形成分株的功能分化, 即克隆内分工。生境的斑块对比度, 作为资源或环境异质性的主要素, 在一定程度上决定克隆内分工的发生状况。该文以鹅绒委陵菜(Potentilla anserina)为材料, 在自然条件下将多组分株对置于不同的斑块对比度处理下, 比较了它们克隆内分工的发生状态, 试图发现分工与斑块对比度的关系, 同时考察在克隆分工过程中分株的可塑性变化及其与分工的关系。该实验的理论假设是: 分株发生分工的程度与分株所处斑块的资源对比度成正相关。研究结果表明, 鹅绒委陵菜分株的高度和叶面积对局部光照环境产生强烈的可塑性反应, 反应的结果是增加了对匮乏的光资源的获取。从分株根冠比和我们提出的分工指数来看, 分工的程度在一定的斑块对比度范围内随斑块对比度的增强而增强, 但到达一个最大值后又迅速降低。鹅绒委陵菜分株之间的分工和结构特化往往滞后于分株对所处局部环境的适应性可塑性变化, 而后者往往在分株之间具有独立性和局部特征。克隆内分工主要依赖于生物量分配的调节而实现, 其发生状态都是分株系统在分工收益、分工代价与分工风险之间权衡的结果, 而这种结果在很大程度上取决于分株所处的斑块对比度。     

克隆整合对异质性土壤养分生境下薇甘菊生长的影响

环境资源的异质性在自然界中普遍存在。克隆植物能通过克隆整合较好地适应异质性生境。本文以海南外来强入侵性植物薇甘菊为材料,通过温室盆栽试验,研究克隆整合对异质性土壤养分生境下薇甘菊克隆片段生长的影响。结果表明:在养分异质下,克隆整合显著提高了低养分斑块分株的生物量,但同时降低了高养分斑块分株的生物量,对克隆片段总体的生长无显著影响,且这一结果不受资源输送方向的影响。克隆整合对薇甘菊克隆分株的光合速率和比叶面积影响较小,但当近端分株处于高养分条件时,其改变了克隆分株的根冠比。这些结果指示:克隆整合有利于异质性土壤养分条件下薇甘菊对低养分斑块的占领。因此,克隆整合可影响薇甘菊对资源异质性生境的入侵能力,它使薇甘菊能够扩展到低养分斑块,从而提高其入侵扩散能力。     

青藏高原东缘过路黄在资源交互斑块性生境中的克隆内资源共享

在青藏高原和四川盆地过渡带,分别于618m和1800m两个海拔高度上研究匍匐茎克隆植物过路黄(Lysimachia christinae)在资源交互斑块性生境中的克隆内资源共享及其对生长的影响.结果显示, 在海拔1800m处,与资源的空间同质性处理(Ⅰ) 和(Ⅱ)相比, 资源的空间异质性处理(Ⅲ)和(Ⅳ)下过路黄整个克隆片段的生物量和分株数均获得显著增加;在海拔618m处,与资源的空间同质性处理(Ⅰ) 和(Ⅱ)相比,资源的空间异质性处理(Ⅲ)和(Ⅳ)下过路黄整个克隆片段生物量显著增加.在海拔618m和1800m处,生长在低光高养条件下的远端分株, 若与高光低养的近端分株相连, 相比连接到低光高养的近端分株, 它们分配更多的生物量到地下部分;在海拔1800m处,生长在高光低养条件下的远端分株, 若与低光高养的近端分株相连, 相比连接到高光低养的近端分株, 它们分配更多的生物量到地上部分.在海拔618m和1800m处,生长在高光低养条件下的近端分株, 若与低光高养的远端分株相连, 相比连接到高光低养的远端分株, 它们分配更多的生物量到地上部分.处于资源交互斑块性生境中的过路黄发生了克隆内分工,依靠相连分株间的功能分化, 克隆植物能有效的利用异质性分布的资源, 缓解资源交互斑块性分布对克隆植物生长的不利影响.通过间隔子(匍匐茎或根状茎),相连分株间能够相互传递和共享由不同分株获得的资源,这种资源共享能够提高克隆植物在异质性生境中的存活与生长.同时,方差分析显示环境异质性和海拔的交互作用显著影响克隆片段的生物量和分株数.相比于海拔618m,在海拔1800m处克隆内资源共享对克隆植物生长表现的影响更大.     

异质性光照生境下克隆整合对外来入侵植物薇甘菊生长的影响

薇甘菊(Mikania micrantha)原产中、南美洲,1919年薇甘菊作为杂草在中国香港出现,目前已广泛侵入中国广东、海南、云南等省区,是世界上最有害的100种外来入侵物种之一。本文以海南重要外来入侵植物薇甘菊(Mikania micrantha)为实验材料,通过温室盆栽实验,研究了克隆整合对异质性光照生境下薇甘菊克隆片段生长的影响。结果表明:在异质性光照下,克隆整合显著提高了低光斑块分株的生物量,但同时降低了高光斑块分株的生物量,对克隆片段总体的生物量积累无显著影响,且这一结果不受资源输送方向的影响;克隆整合降低了异质性光照下克隆分株间根冠比的差异,加大了克隆分株间净光合速率的差异,且倾向于提高生长在高光斑块分株的比叶面积;克隆整合虽然并不能促进薇甘菊匍匐茎克隆片段在异质性光照生境下总体的生长,但可以促进低光斑块下分株的生长,因而,克隆整合可促进薇甘菊从开阔生境向低光生境(森林、杂草群落等)的入侵能力,因此,克隆整合特性是薇甘菊对异质性光照环境的重要适应对策之一。     

异质性生境中匍匐茎草本野草莓(Fragaria vesca)的克隆内资源共享

研究了不同海拔高度(1800和3900m)的匍匐茎克隆植物野草莓(Fragaria vesca)种群对光照和养分资源斑块性分布生境的响应。结果表明:与资源的空间同质性处理(Ⅰ)和(Ⅱ)相比,资源的空间异质性处理(Ⅲ)和(Ⅳ)中2个种群的野草莓的近端、远端分株部分和整个克隆片段的生物量和分株数均明显增加。当近端分株部分经历低光高养,而与其相连的远端分株部分经历高光低养时,相比于整个克隆片段都处于低光高养的同质性生境,来自2个海拔的种群的近端分株部分都会增加对根的生物量分配;当近端分株部分经历高光低养,而与其相连的远端分株部分经历低光高养时,相比于整个克隆片段都处于低光高养的同质性生境,来自2个海拔的种群的近端分株部分都会减少对根的生物量分配,远端分株部分也被观察到类似的生物量分配格局。相比于高光低养的同质性生境,当与低光高养的远端分株部分相连时,经历高光低养的近端分株部分有更大的叶面积;相比于低光高养的同质性生境,当与低光高养近端分株部分相连时,经历高光低养的远端分株部分有更大的叶面积。结果表明,野草莓在资源交互斑块性生境中发生了克隆内分工,克隆内分工有利于克隆植物对异质性资源的利用,对克隆植物在资源斑块性分布生境中的生存和生长具有重要的意义。     

基于根系形态可塑性的空心莲子草克隆分工特征

资源在空间和时间上不均匀分布现象往往形成资源异质性斑块,克隆植物凭借强大的侧向生长能力占据广阔空间,分株间的生理连接促进了其对异质性生境的适应。克隆分株首先通过资源获取结构的功能特化来提高从各种资源富养斑块中的养分获取,然后通过克隆整合作用实现分株间的养分传输,这种功能特化和资源共享模式被称为‘分工’。该文以入侵克隆植物空心莲子草(Alternanthera philoxeroides)为研究对象,研究其根系对资源异质性分布的形态可塑性响应;通过调节光照强度和土壤养分来实现资源的异质性分布,共设置4个处理:1近端分株高光低养—远端分株高光低养(HL-HL),2近端分株低光高养—远端分株低光高养(LH-LH),3近端分株高光低养—远端分株低光高养(HL-LH),4近端分株低光高养—远端分株高光低养(LH-HL);使用WinRHIZO Pro软件分析相关根系指标,SPSS 18.0单因素方差(one-way ANOVA)分析方法分析异质性条件对近、远端分株以及整个克隆片段的影响。结果表明:异质性斑块中经历高光低养的分株分配更多的生物量到地上部分,经历低光高养的分株分配更多的生物量到地下部分,空心莲子草通过调整对地上和地下部分的生物量分配比例实现了克隆分工;异质性斑块中,生长在富养斑块中的空心莲子草分株根系有更高的根生物量、根长、根表面积、根体积以及分枝系数等,表明空心莲子草分株根系通过对异质性斑块的形态可塑性变化提高了土壤养分的吸收能力。由此可见,空心莲子草通过对资源获取结构的功能特化提高了其资源吸收能力,这可能是其具强入侵能力的重要原因。     

Clonal Plasticity in Response to Reciprocal Patchiness of Light and Nutrients in the Stoloniferous Herb Glechoma longituba L.

Interconnected ramets of clonal plants can functionally specialize in the uptake of resources from aboveground and/or underground sources. Ramet pairs of the clonal stoloniferous herb Glechoma Iongltuba L. were grown In spatially heterogeneous environments with complementary availability of light and nutrients. Plasticity with respect to root-shoot ratio, fitness-related traits （biomass, number of ramets and dry weight per ramet）, morphological traits （lamina area, root length） were experimentally examined. The aim was to understand the adaptation of G. Iongltuba to an environment with reciprocal patchiness of light and soil nutrients by plasticity In Its root-shoot ratio and clonal morphology. The results showed that the performance of ramets growing In patches with high light Intensity and low soil nutrients into the adjacent opposite patches was Increased in terms of fitness-related traits. However, the performance of those from patches with low light Intensity and high soil nutrients into the adjacent opposite patches was not changed. The root-shoot ratio and clonal morphology were plastic. Morphological traits such as lamina area and root length were altered In a way that enhanced the capture of light resources and soil nutrients. Apparent reciprocal resource transport between the ramets In an environment of reciprocal patchiness of resources can enhance the growth of ramets with complementary resource deficiencies.     

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.     

Do Clonal Plants Show Greater Division of Labour Morphologically and Physiologically at Higher Patch Contrasts?

Background

When growing in reciprocal patches in terms of availability of different resources, connected ramets of clonal plants will specialize to acquire and exchange locally abundant resources more efficiently. This has been termed division of labour. We asked whether division of labour can occur physiologically as well as morphologically and will increase with patch contrasts.

Methodology/Principal Findings

We subjected connected and disconnected ramet pairs of Potentilla anserina to Control, Low, Medium and High patch contrast by manipulating light and nutrient levels for ramets in each pair. Little net benefit of inter-ramet connection in terms of biomass was detected. Shoot-root ratio did not differ significantly between paired ramets regardless of connection under Control, Low and Medium. Under High, however, disconnected shaded ramets with ample nutrients showed significantly larger shoot-root ratios (2.8∼6.5 fold) than fully-lit but nutrient-deficient ramets, and than their counterparts under any other treatment; conversely, fully-lit but nutrient-deficient ramets, when connected to shaded ramets with ample nutrients, had significantly larger shoot-root ratios (2.0∼4.9 fold) than the latter and than their counterparts under any other treatment. Only under High patch contrast, fully-lit ramets, if connected to shaded ones, had 8.9% higher chlorophyll content than the latter, and 22.4% higher chlorophyll content than their isolated counterparts; the similar pattern held for photosynthetic capacity under all heterogeneous treatments.

Conclusions/Significance

Division of labour in clonal plants can be realized by ramet specialization in morphology and in physiology. However, modest ramet specialization especially in morphology among patch contrasts may suggest that division of labour will occur when the connected ramets grow in reciprocal patches between which the contrast exceeds a threshold. Probably, this threshold patch contrast is the outcome of the clone-wide cost-benefit tradeoff and is significant for risk-avoidance, especially in the disturbance-prone environments.     

Clonal Integration of Fragaria orientalis in Reciprocal and Coincident Patchiness Resources: Cost-Benefit Analysis

Clonal growth allows plants to spread horizontally and to experience different levels of resources. If ramets remain physiologically integrated, clonal plants can reciprocally translocate resources between ramets in heterogeneous environments. But little is known about the interaction between benefits of clonal integration and patterns of resource heterogeneity in different patches, i.e., coincident patchiness or reciprocal patchiness. We hypothesized that clonal integration will show different effects on ramets in different patches and more benefit to ramets under reciprocal patchiness than to those under coincident patchiness, as well as that the benefit from clonal integration is affected by the position of proximal and distal ramets under reciprocal or coincident patchiness. A pot experiment was conducted with clonal fragments consisting of two interconnected ramets (proximal and distal ramet) of Fragaria orientalis. In the experiment, proximal and distal ramets were grown in high or low availability of resources, i.e., light and water. Resource limitation was applied either simultaneously to both ramets of a clonal fragment (coincident resource limitation) or separately to different ramets of the same clonal fragment (reciprocal resource limitation). Half of the clonal fragments were connected while the other half were severed. From the experiment, clonal fragments growing under coincident resource limitation accumulated more biomass than those under reciprocal resource limitation. Based on a cost-benefit analysis, the support from proximal ramets to distal ramets was stronger than that from distal ramets to proximal ramets. Through division of labour, clonal fragments of F. orientalis benefited more in reciprocal patchiness than in coincident patchiness. While considering biomass accumulation and ramets production, coincident patchiness were more favourable to clonal plant F. orientalis.     

Effects of soil heterogeneity and clonal integration on Alternanthera philoxeroides populations with a radial ramet aggregation

Some clonal plants can spread their ramet populations radially, and soil heterogeneity and clonal integration may greatly affect the establishment of these types of populations. We constructed Alternanthera philoxeroides populations with a radial ramet aggregation, allowing old ramets of clonal fragments to concentrate in central pots and younger ramets to root in peripheral pots. The peripheral pots were supplemented either with three different levels (high, medium and low) of soil nutrients to simulate a heterogeneous soil environment, or only one medium level of soil nutrients to simulate a homogeneous environment. Stolon connections between the central older ramets and the peripheral younger ramets were left intact or severed to test the effect of clonal integration. The maintenance of stolon connection could induce the division of labor between different‐aged ramets, by increasing the root investment in central ramets and the above‐ground growth in peripheral ramets. The maintenance of stolon connection could improve the growth of the central and peripheral ramets, clonal fragments and even the whole population. However, the positive consequence in peripheral ramets and whole fragments was only detected in the high‐nutrient patch of heterogeneous treatment. In sum, in the population with the radial ramet aggregation, clonal integration can play a key role in the rapid recruitment of young ramets of A. philoxeroides fragments, as well as the expansion of the whole population. The magnitude of clonal integration also became more obvious in the peripheral young ramets and whole fragments that experienced high‐nutrient patches.     

Environmentally-induced functional specialization for locally abundant resources in the stoloniferous herb Duchesnea indica

Reciprocally patchy environments, where the availability of two resources are patchily distributed and negatively correlated in each patch, are common in many ecosystems. Interconnected ramets of clonal plants can specialize in the uptake of locally abundant resources. Ramet pairs of the stoloniferous herb Duchesnea indica were grown in reciprocally patchy environments i.e., one ramet of a pair was grown in the high light but low water patch (high light patch) and the other in the low light but high water patch (high water patch). Biomass allocation pattern (root-shoot ratio), morphological traits (leaf area and root length) and physiological traits (photosynthetic rate and chlorophyll content) were altered in a way that potentially enables ramets to enhance the capture of the locally abundant resource (i.e., increase the capture of light resource in the high light patch and of water in the high water patch). As a result,biomass and number of ramets in the connected ramet pairs were greatly improved. Functional specialization of ramets, modified by clonal integration, may have contributed greatly to the growth increase of D. indica in the reciprocally patchy environment.     

Evidence for unexpected higher benefits of clonal integration in nutrient-rich conditions

Physiologically integrated clonal plants cope better with spatial heterogeneity due to their ability to share resources among ramets. According to theoretical predictions and experimental evidence, such benefits of resource sharing should increase with higher patch quality of an exporting ramet and lower patch quality of an importing ramet. This study investigated the effect of spatial heterogeneity in nutrient availability on benefits of clonal integration under plausible scenarios of clonal spread, in which more developed ramets give rise to new ones. Pairs of mother and daughter ramets of a stoloniferous grass, Agrostis stolonifera, were grown in various nutrient conditions. Disconnected pairs of ramets were used as controls. Results showed considerable benefits of integration for developmentally younger daughters and no costs for older mothers in all treatments. Surprisingly, benefits of integration were more pronounced in nutrient-rich daughters, and allocation to integrated daughters decreased with increasing nutrient level of mothers. In addition, integration in general increased root-to-shoot ratio of daughters. One possible explanation of the observed patterns may be prevailing translocation of photosynthates rather than nutrients. Daughters also responded to nutrients by changes in clonal architecture. Number of stolons increased, and maximum stolon length decreased in high nutrient levels. Integration increased maximum stolon length in small daughters. The architectural responses are generally in accord with the foraging behaviour concept. Overall, our results suggest that resource translocation within a clonal fragment need not be easily predictable from a gradient of resource availability.     

Environmentally-induced functional specialization for locally abundant resources in the stoloniferous herb Duchesnea indica

Reciprocally patchy environments, where the availability of two resources are patchily distributed and negatively correlated in each patch, are common in many ecosystems. Interconnected ramets of clonal plants can specialize in the uptake of locally abundant resources. Ramet pairs of the stoloniferous herb Duchesnea indica were grown in reciprocally patchy environments i.e., one ramet of a pair was grown in the high light but low water patch (high light patch) and the other in the low light but high water patch (high water patch). Biomass allocation pattern (root-shoot ratio), morphological traits (leaf area and root length) and physiological traits (photosynthetic rate and chlorophyll content) were altered in a way that potentially enables ramets to enhance the capture of the locally abundant resource (i.e., increase the capture of light resource in the high light patch and of water in the high water patch). As a result,biomass and number of ramets in the connected ramet pairs were greatly improved. Functional specialization of ramets, modified by clonal integration, may have contributed greatly to the growth increase of D. indica in the reciprocally patchy environment.