根茎克隆植物生态学研究进展

根茎在植物的无性繁殖、克隆分株间信息交流和物质交换、预测资源斑块的质量等方面具有重要意义,并且根茎克隆植物的研究涉及生物入侵、全球变化等诸多生态学前沿领域。作为一种重要的克隆植物类型,根茎克隆植物在资源异质性生境中表现出特有的适应方式,这种方式可以通过形态可塑性、觅食行为、生理整合以及适合度来具体表征。着眼于根茎克隆植物,总结和分析了国内外近年来的研究案例,并对形态可塑性起源与多样性的限制假说和适应假说、觅食行为中的强度觅食和广度觅食策略、克隆分株间间隔子保持和断裂的利益权衡等热点内容进行了讨论。最后联系生态学学科前沿,提出了本领域在未来需要重视的研究方向。     

克隆植物对异质生境的适应对策研究进展

生境异质性是自然生态系统的基本特征,植物生长的必需资源和环境胁迫因子均存在着复杂的时间和空间异质性。克隆植物是指在自然条件下具有克隆特性的植物,即可通过与母株相连的芽、根茎、分蘖或枝条等繁殖体产生无性繁殖的植物,这些繁殖体一旦定居便可成为潜在的独立个体。克隆植物具有独特的生境适应策略(如形态可塑性、克隆整合、克隆分工、觅食行为、风险分摊等),面对异质性的生境条件,它可以通过调整自身的生理和形态结构来适应异质生境。目前,对于克隆植物在异质生境适应行为的研究已有很多报道,然而系统性的归纳和总结尚有欠缺。综述了克隆植物在不同资源异质生境(光照、养分、水分)和不同胁迫生境(盐碱胁迫、风沙胁迫、重金属胁迫)下独特的适应对策。最后,针对克隆植物对异质生境的适应对策,进行了总结并对未来的重点研究方向提出建议：(1)时间异质性尺度上的考量;(2)异质性生境中生物因子的调控作用;(3)克隆植物入侵机制;(4)克隆植物在生态修复中的应用潜力。     

异质生境中水生植物表型可塑性的研究进展

水生植物是一类以草本植物为主、与水紧密相关的生态类群, 大多数具有克隆性。面对水环境的变化, 水生植物在形态、行为和生理上表现出多样化的表型可塑性, 对异质生境具有很强的适应能力。表型可塑性研究已在陆生植物的多个类群展开, 然而目前对异质生境下水生植物的生态适应对策, 尤其是表型可塑性的研究尚重视不够。本文在阐明克隆植物表型可塑性主要实现方式及其关系、水生环境异质性及其特点的基础上, 重点从形态可塑性、觅食行为、克隆整合、克隆分工和风险分摊等5个方面讨论了水生植物如何通过表型可塑性适应异质性水生环境。在今后的水生植物表型可塑性研究中, 建议着重探讨以下问题: (1)表型可塑性的变化规律及机理; (2)克隆整合对群落和生态系统的影响; (3)克隆整合与克隆片段化的权衡; (4)不同克隆构型的表型可塑性及其内在机制; (5)表型可塑性的适应性进化; (6)水生植物与其他类群/营养级物种的关系; (7)水生生态系统对全球变化的响应。     

竹类植物对异质生境的适应——表型可塑性

竹类植物是一类以木本为主的克隆植物,凭借表型可塑性的优势,对异质生境具有很强适应能力。然而,目前对竹类植物表型可塑性的实现方式及其异质生境适应对策未见系统总结,从而在一定程度上限制了竹类生态学的发展。从形态可塑性、选择性放置、克隆整合和克隆分工等4个方面对竹类植物的表型可塑性研究进行分析和梳理,结果表明:竹类植物在异质生境中具有明显的表型可塑反应,主要采用形态可塑性、选择性放置和克隆整合来适应异质生境,而克隆分工的普遍性仍有待验证;目前侧重于研究构件形态和生物量分配格局,而很少深入探讨形态、生理和行为等可塑性机理。今后竹类植物表型可塑性研究重点在于:1)克隆整合的格局与机理;2)克隆整合对生态系统的影响;3)克隆分工的形成及其与环境关系;4)表型可塑性的等级性及环境影响;5)不同克隆构型的表型可塑性特征及其内在机制。     

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     

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。     

蛇莓克隆构型对光照强度的可塑性反应

克隆植物构型的可塑性反应可能是它在资源斑块性状分布的环境中获取资源对策的重要方面,因而可能具有重要的生态学意义。在一田间实验中,匍匐茎草本蛇毒（Duchesnea indica Focke）经历了不同相对光照强度的处理,以研究光照强度对蛇莓克隆构型的影响。结果表明：随着相对光照强度的增加,间隔子的长度逐渐降低,分枝角度、分枝强度和分株密度呈二次曲线变化。结合植物对环境异质性的利用对策,对的揭示的蛇莓克隆构型可塑性进行了讨论。     

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     

匍匐茎草本蛇莓克隆构型对土壤水分的可塑性反应

克隆植物构型的可塑性有可能促进它对斑块性分布土壤水分资源的利用,因而可能具有生态学意义.在田间实验中,匍匐茎草本蛇莓(Duchesnea indica Focke)经历了不同土壤水分水平(土壤最大含水量的40%、60%、80%、100%等)处理,以研究土壤水分对蛇莓克隆构型的影响.结果表明:间隔子长度、分株密度、分枝角度和分枝强度呈二次曲线变化,土壤含水为最大含水量的80%的生境为最适.在不同土壤水分水平生境中,蛇莓克隆构型相关特征的可塑性变化可用动态Logistic模型进行模拟和预测,拟合效果较好.结合植物对环境异质性的利用对策,对所揭示的蛇莓克隆构型可塑性进行了讨论.     

异质性光照条件下匍匐茎草本野草莓的克隆整合

通过盆栽实验,研究了匍匐茎草本野草莓在异质性光照条件下的克隆整合。结果显示克隆整合显著增强了野草莓胁迫分株段的生长,损—益分析表明未受胁迫分株没有显著损耗,整个克隆片段的生长得到显著提高。在局部遮荫处理,克隆整合对克隆形态可塑性的修饰作用没有观察到。最后,讨论了克隆植物对环境的生态适应意义。     

Phylogenetic Meta-Analysis of the Functional Traits of Clonal Plants Foraging in Changing Environments

Foraging behavior, one of the adaptive strategies of clonal plants, has stimulated a tremendous amount of research. However, it is a matter of debate whether there is any general pattern in the foraging traits (functional traits related to foraging behavior) of clonal plants in response to diverse environments. We collected data from 97 published papers concerning the relationships between foraging traits (e.g., spacer length, specific spacer length, branch intensity and branch angle) of clonal plants and essential resources (e.g., light, nutrients and water) for plant growth and reproduction. We incorporated the phylogenetic information of 85 plant species to examine the universality of foraging hypotheses using phylogenetic meta-analysis. The trends toward forming longer spacers and fewer branches in shaded environments were detected in clonal plants, but no evidence for a relation between foraging traits and nutrient availability was detected, except that there was a positive correlation between branch intensity and nutrient availability in stoloniferous plants. The response of the foraging traits of clonal plants to water availability was also not obvious. Additionally, our results indicated that the foraging traits of stoloniferous plants were more sensitive to resource availability than those of rhizomatous plants. In consideration of plant phylogeny, these results implied that the foraging traits of clonal plants (notably stoloniferous plants) only responded to light intensity in a general pattern but did not respond to nutrient or water availability. In conclusion, our findings on the effects of the environment on the foraging traits of clonal plants avoided the confounding effects of phylogeny because we incorporated phylogeny into the meta-analysis.     

A modular concept of plant foraging behaviour: the interplay between local responses and systemic control

In this paper we examined the notion that plant foraging for resources in heterogeneous environments must involve: (1) plasticity at the level of individual modules in reaction to localized environmental signals; and (2) the potential for modification of these responses either by the signals received from connected modules that may be exposed to different conditions, or by the signals reflecting the overall resource status of the plant. A conceptual model is presented to illustrate how plant foraging behaviour is achieved through these processes acting in concert, from the signal reception through signal transduction to morphological or physiological response. Evidence to support the concept is reviewed, using selective root placement under nutritionally heterogeneous conditions and elongation responses of stems and petioles to shade as examples. We discussed how the adoption of this model can promote understanding of the ecological significance of foraging behaviour. We also identified a need to widen the experimental repertoires of both molecular physiology and ecology in order to increase our insight into both the regulation and functioning of foraging responses, and their relationship with the patterns of environmental heterogeneity under which plants have evolved.     

Root Foraging Increases Performance of the Clonal Plant Potentilla reptans in Heterogeneous Nutrient Environments

Background

Plastic root-foraging responses have been widely recognized as an important strategy for plants to explore heterogeneously distributed resources. However, the benefits and costs of root foraging have received little attention.

Methodology/Principal Findings

In a greenhouse experiment, we grew pairs of connected ramets of 22 genotypes of the stoloniferous plant Potentilla reptans in paired pots, between which the contrast in nutrient availability was set as null, medium and high, but with the total nutrient amount kept the same. We calculated root-foraging intensity of each individual ramet pair as the difference in root mass between paired ramets divided by the total root mass. For each genotype, we then calculated root-foraging ability as the slope of the regression of root-foraging intensity against patch contrast. For all genotypes, root-foraging intensity increased with patch contrast and the total biomass and number of offspring ramets were lowest at high patch contrast. Among genotypes, root-foraging intensity was positively related to production of offspring ramets and biomass in the high patch-contrast treatment, which indicates an evolutionary benefit of root foraging in heterogeneous environments. However, we found no significant evidence that the ability of plastic foraging imposes costs under homogeneous conditions (i.e. when foraging is not needed).

Conclusions/Significance

Our results show that plants of P. reptans adjust their root-foraging intensity according to patch contrast. Moreover, the results show that the root foraging has an evolutionary advantage in heterogeneous environments, while costs of having the ability of plastic root foraging were absent or very small.     

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.     

No evidence for nutrient foraging in root-sprouting clonal plants

Clonality is defined as vegetative reproduction via the production of ramets, which are, at least initially, connected by spacers. In general, there are three types of spacers of two origins. Whereas stolons are aboveground spacers, rhizomes are belowground spacers; however, both of stem origin. The third type of spacers are roots in root-sprouting plants. The possibility of foraging in clonal plants has attracted broad interest among ecologists but has been experimentally documented only for stoloniferous clonal plants foraging for light. Foraging for belowground resources has yet to be demonstrated, perhaps because tests of foraging have focused on clonal plants that spread laterally via stolons or rhizomes, i.e. stem organs. Lateral spread based on sprouting roots has not been considered even though, in addition to functioning as conduits between ramets, root spacers are able to sense and take up nutrients. We therefore hypothesized that root-sprouting clonal plants may be able to directly react to environmental heterogeneity and exhibit nutrient foraging. To test this hypothesis, we conducted two experiments with root-sprouters in nutrient-heterogeneous and -homogeneous environments. We found that plants produced more biomass when growing in a heterogeneous environment than in a homogeneous environment and that root biomass was greater in the nutrient-rich patches than in nutrient-poor patches. However, the number of ramets did not differ between patches in the heterogeneous environment. We conclude that plants whose clonality is based on roots, similarly as plants whose clonality is based on stolons or rhizomes, do not exhibit accumulation of ramets in nutrient-rich patches. Foraging at the organ level, i.e. by roots, seems to be more probable in this clonal group. To analyse how clonal plants with different clonal strategies perceive and react to environmental heterogeneity, researchers must account for the high variability in clonal growth forms and in scales of environmental heterogeneity.     

Soil heterogeneity affects ramet placement of Hydrocotyle vulgaris

Aims Soil heterogeneity is common in natural habitats. It may trigger foraging responses (placing more ramets and/or roots in nutrient-rich patches than in nutrient-poor patches) and further affect the growth of plants. However, the impact of soil heterogeneity on competitive interactions has been little tested.Methods We conducted a greenhouse experiment to investigate the effects of soil heterogeneity on intraspecific competition with a stoloniferous herb Hydrocotyle vulgaris. We grew one (without competition) or nine ramets (with competition) of H. vulgaris under a homogeneous environment and two heterogeneous environments differing in patch size (large or small patches). In the heterogeneous treatment, the soil consisted of the same number of nutrient-rich and nutrient-poor patches arranged in a chessboard manner, and in the homogeneous treatment, the soil was an even mixture of the same amount of the nutrient-rich and the nutrient-poor soil.Important findings Irrespective of intraspecific competition, H. vulgaris showed foraging responses to soil heterogeneity in the large patch treatment, e.g. it produced significantly more biomass, ramets, aboveground mass and root mass in the nutrient-rich patches than in the nutrient-poor patches. In the small patch treatment, foraging responses were observed when intraspecific competition was present, but responses were not observed when there was no competition. However, we find a significant effect of soil heterogeneity on neither overall growth nor competitive intensity of H. vulgaris. Our results suggest that foraging responses to soil heterogeneity may not necessarily be adaptive and intraspecific competition may not be influenced by soil heterogeneity.     

Response of physiological integration in Trifolium repens to heterogeneity of UV-B radiation

Physiological integration has been documented in many clonal plants growing under resource heterogeneity. Little is still known about the response of physiological integration to heterogeneous ultraviolet-B radiation. In this paper, the changes in intensity of physiological integration and of physiological parameters under homogeneous and heterogeneous ultraviolet-B radiation (280-315 nm) were measured in order to test the hypothesis that in addition to resource integration a defensive integration in Trifolium repens might exist as well. For this purpose, homogeneous and heterogeneous ultraviolet-B radiation was applied to pairs of connected and severed ramets of the stoloniferous herb Trifolium repens. Changes in intensity of water and nutrient integration were followed with acid fuchsin dye and ¹⁵N-isotope labeling of the xylem water transport. In order to assess the patterns of physiological integration contents of chlorophyll, ultraviolet-B absorbing compounds, soluble sugar and protein were determined and activities of superoxide dismutase (SOD) and peroxidase (POD) measured. When ramets were connected and exposed to heterogeneous UV-B radiation, the velocity of water transportation from the UV-B treated ramet to its connected sister ramet was markedly lower and the percentage of ¹⁵N left in labelled ramets that suffered from enhanced UV-B radiation was higher and their transfer to unlabelled ramets lower. In comparison with clones under homogeneous ultraviolet-B radiation, the content of chlorophyll, ultraviolet-B absorbing compounds, soluble sugar and activities of SOD and POD increased notably if ultraviolet-B stressed ramets were connected to untreated ramets. Chlorophyll and UV-B absorbing compounds were shared between connected ramets under heterogeneous UV-B radiation. This indicated that physiological connection improved the performance of whole clonal plants under heterogeneous ultraviolet-B radiation. The intensity of physiological integration of T. repens for resources decreased under heterogeneous ultraviolet-B radiation in favor of the stressed ramets. Ultraviolet-B stressed ramets benefited from unstressed ramets by physiological integration, supporting the hypothesis that clonal plants are able to optimize the efficiency of their resistance maintaining their presence also in less favorable sites. The results could be helpful for further understanding of the function of heterogeneous UV-B radiation on growth regulation and microevolution in clonal plants.