Importance of starting points in heterogeneous environments: interactions between two clonal plants with contrasting spatial architectures

在异质性环境中，植物能够通过将更多的生物量投资到高资源斑块而获利。因此，相对于高资源斑块的初始生长位置可能会对植物的生长和竞争能力产生影响，且这种影响可能与植物的构型有关。然而，以往有关土壤异质性对植物生长和竞争影响的研究都忽略了植 物初始生长位置的潜在效应。在本研究中，我们把密集型克隆植物翼果苔草(Carex neurocarpa)和游击型克隆植物扁秆荆三棱(Bolboschoenus planiculmis)分别单独种植(不存在竞争)或者混合种植(存在竞争)在由高营养斑块和低营养斑块组成的异质性环境中。在不存在竞争处理时，每种植物的一个分株被分别单独种植于高营养或者低营养斑块中；在存在竞争处理时，目标种的一个分株被种植于高营养斑块中，而竞争种被种植在这个高营养斑块中或者邻近的低营养斑块中，或目标种的一个分株被种植于低营养斑块中，而竞争种被种植在这个低营养斑块中或者邻近的高营养斑块中。结果表明，在不存在竞争时，初始生长位置处于高营养斑块的翼果苔草的生物量和分株数均显著高于初始位置处于低营养斑块的翼果苔草，但在存在竞争时，却未发现这种差异。因此，初始位置处于高营养斑块的翼果苔草受到的竞争强度要显著高于初始位置处于低营养斑块的翼果苔草，而且这种效应与其竞争者的初始生长位置无关。相反的，扁秆荆三棱的初始生长位置对其生长和竞争的响应没有影响。因此，在异质性环境中，克隆植物的初始生长位置能够影响其生长和相对竞争能力，并且这些效应可能取决于植物的克隆构型。     

Clonal performance of Scirpus yagara in multiple levels of substrate heterogeneity and submergence

荆三棱在多等级基质异质性与水淹处理下的克隆表现 环境异质性可以影响克隆水生植物的表现。鲜有研究者关注两个层次的环境异质性并将其融入 对克隆植物生态学的研究中。本研究的目的是: (1)检验不同基质异质性与水淹处理是否对植物表现产生相 似效应，(2)探索克隆植物的觅食行为。本研究将荆三棱(Scirpus yagara)置于不同基质异质性与水淹处理之中。基质处理包括1个均质性基质处理(湖泥与沙等体积混合)与3个异质性基质处理(湖泥斑块与沙斑块交错构建的两斑块、四斑块与八斑块基质)。水淹处理包括：0、10和30 cm。本实验测量了克隆分株数、克隆代数、叶数、球茎数、克隆分株高度、茎长、根状茎长、克隆半径、间隔子长、间隔子厚度、总生物量、球茎生物量与单个球茎生物量等性状数据。研究结果表明，水位上升导致克隆分株数、克隆代数、叶数和球茎数显著减少，同时基质异质性造成间隔子长度与间隔子厚度的显著变化。水位与基质异质性两因子对克隆分株数、叶数和间隔子长度产生了显著的交互效应。在两斑块基质与四斑块基质中，荆三棱对湖泥斑块表现出显著的觅食行为，更多的构件被放置于湖泥斑块中。尤其在两斑块基质中，所有的构件被放置于湖泥斑块中。在八斑块基质中，荆三棱表现出双向觅食，这导致构件在不同斑块中的均匀放置。研究结果表明，荆三棱的觅食行为与斑块大小具有相关性。     

根茎型植物扁秆荆三棱对光照强度和养分水平的生长响应及资源分配策略

自然界中光照和养分因子常存在时空变化,对植物造成选择压力。克隆植物可通过克隆生长和生物量分配的可塑性来适应环境变化。尽管一些研究关注了克隆植物对光照和养分因子的生长响应,但尚未深入全面了解克隆植物对光照和养分资源投资的分配策略。以根茎型草本克隆植物扁秆荆三棱(Bolboschoenus planiculmis)为研究对象,在温室实验中,将其独立分株种植于由2种光照强度(光照和遮阴)和4种养分水平(对照、低养分、中养分和高养分)交叉组成的8种处理组合中,研究了光照和养分对其生长繁殖及资源贮存策略的影响。结果表明,扁秆荆三棱的生长、无性繁殖及资源贮存性状均受到光照强度的显著影响,在遮阴条件下各生长繁殖性状指标被抑制。且构件的数目、长度等特征对养分差异的可塑性响应先于其生物量积累特征。在光照条件下,高养分处理的总生物量、叶片数、总根茎分株数、长根茎分株数、总根茎长、芽长度、芽数量等指标大于其他养分处理,而在遮阴条件下,其在不同养分处理间无显著差异,表明光照条件可影响养分对扁秆荆三棱可塑性的作用,且高营养水平不能补偿由于光照不足而导致的生长能力下降。光照强度显著影响了总根茎、总球茎及大、中、小球茎的生物量分配,遮阴条件下,总生物量减少了对地下部分根茎和球茎的分配,并将有限的生物量优先分配给小球茎。总根茎的生物量分配未对养分发生可塑性反应,而随着养分增加,总球茎分配下降,说明在养分受限的环境中球茎的贮存功能可缓冲资源缺乏对植物生长的影响。在相同条件下,根茎生物量对长根茎的分配显著大于短根茎,以保持较高的繁殖能力;而总球茎对有分株球茎的生物量分配小于无分株球茎,表明扁秆荆三棱总球茎对贮存功能的分配优先于繁殖功能。研究为进一步理解根茎型克隆植物对光强及基质养分环境变化的生态适应提供了依据。     

土壤养分斑块对比度改变活血丹克隆整合强度和方向

克隆植物相连片段(或分株)常常生长在不同的土壤养分斑块中。克隆整合使得生长在异质养分斑块中的克隆片段(或分株)产生局部和非局部反应,从而影响相连片段(或分株)的表型可塑性。为了揭示养分斑块对比度对活血丹(Glechoma longituba)克隆整合的影响,在一控制实验中,将活血丹克隆片断种植于4种不同对比度的环境中,即:无对比度(对照)、低对比度、中对比度和高对比度。活血丹在气体交换、水势、荧光、形态、生长与分配方面的克隆整合强度随养分斑块对比度的增强而表现出增强或减弱的变化趋势;养分斑块对比度越强,活血丹气体交换和荧光暗反应的整合强度越小,叶片水势整合强度越大。斑块对比度可改变部分性状的克隆整合方向;超过一定的对比阈值,整合强度随养分对比度的变化趋势会向着相反方向转变。克隆整合对生理特征的修饰幅度小于对生长特征的修饰幅度。这些结果指示:养分斑块对比度可通过修饰克隆整合格局(即强度和方向)而改变克隆植物的表型可塑性。     

土壤养分异质性和种间竞争对互花米草和芦苇生长繁殖特性的影响

土壤养分异质性在自然界中普遍存在。土壤养分异质性对克隆植物生长繁殖有一定影响,但耦合竞争对入侵植物和本土植物生长繁殖影响的研究却相对匮乏。以入侵克隆植物互花米草和本土克隆植物芦苇为研究对象,通过温室控制实验模拟设置了土壤养分同质和养分异质处理(保证两种土壤的总养分含量相同),交叉3种定植模式(实验容器内单植入侵种6株,单植本土种6株以及两物种各3株交替种植),探究土壤养分异质性和种间竞争对入侵种互花米草和本土种芦苇生长繁殖的影响。结果表明:土壤养分异质性显著降低了互花米草的单株叶片数,而增加了其单株节间长以及芦苇的单株根状茎长;种间竞争模式下,互花米草的单株分株数,以及两种植物的单株节间长、根状茎节数和根状茎长都显著高于单植模式,而芦苇的单株地上生物量、叶生物量、茎生物量都显著低于单植模式;土壤养分异质性和种间竞争对互花米草单株的叶片数和茎生物量产生了显著的交互作用,而对芦苇各指标无显著影响。这些结果说明,土壤养分异质性可以影响互花米草与芦苇的种间竞争关系。     

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.     

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。     

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     

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。     

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。     

Effects of Spatial Scale of Soil Heterogeneity on the Growth of a Clonal Plant Producing Both Spreading and Clumping Ramets

Soil nutrients are commonly heterogeneously distributed at different spatial scales. Although numerous studies have tested the effects of soil nutrient heterogeneity on growth of clonal plants producing either spreading ramets or clumping ramets, no study has examined the effects on the growth of clonal plants producing both spreading and clumping ramets and how spatial scale affects such effects. To test these effects, clones of Buchloe dactyloides, a stoloniferous clonal plant that produces both clumping and spreading ramets, were grown in six heterogeneous environments with different patch sizes and one homogeneous environment containing the same quantity of nutrients. Total biomass, total number of ramets, number of clumping ramets, number of spreading ramets, spacer length, or root:shoot ratio of the whole plants did not differ significantly among the seven treatments. However, at the patch level there were significant effects of patch size by nutrient level on biomass, number of ramets, number of spreading ramets, and number of clumping ramets, and these four variables were significantly larger in the nutrient-rich patches than in the nutrient-poor patches in the heterogeneous treatment with the largest patch size, but not in the other five heterogeneous treatments with smaller patch sizes. Neither nutrient level nor patch size significantly affected spacer length or root:shoot ratio. Based on our results, we propose that B. dactyloides can efficiently exploit nutrient-rich patches by a plastic response of clumping ramets and spreading ramets at larger spatial scales of soil heterogeneity but not at smaller ones.     

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.     

Influence of sediment fertility on morphological variability of Vallisneria spiralis L.

One homogeneous and three heterogeneous nutrient enrichment treatments were imposed to investigate the growth responses of Vallisneria spiralis L. Morphological features of V. spiralis differed significantly between different nutrient patches. Roots elongated in nutrient-poor patches, and the specific root length (SRL) also increased significantly. Stolon length, diameter and leaf length and width increased significantly in nutrient-rich patches. Total plant biomass of V. spiralis grown in the homogeneous and three heterogeneous treatments on average were 2.9, 3.0, 3.9 and 2.3 fold higher than that grown in the control treatment. Number of ramets per clone was significantly higher in the heterogeneous treatments than in the homogeneous treatment. In three varying heterogeneous treatments, ramet biomass in nutrient-rich patches was 2.7, 4.3 and 3.0 fold higher than in nutrient-poor patches; however, ramet number was not affected by sediment nutrients, resulting in bigger ramets in nutrient-rich patches. The biomass allocation established adaptive plasticity to heterogeneous environments. The maximum value of biomass allocation to underground parts reached 16% in nutrient-rich patches, whereas the minimum value of underground parts reached 20% in nutrient-poor patches. Results demonstrate that clonal V. spiralis can maintain itself preferentially in favourable nutrient-rich sediments, whereas nutrient-poor conditions could be escaped by plastic biomass allocation.     

Soil nutrient patchiness affects nutrient use efficiency,though not photosynthesis and growth of parental Glechoma longituba ramets: both patch contrast and direction matter

Aims Most plants are clonal in nature. Clonal ramets can share water, nutrients and photosynthate, especially when they experience patchy resources. Patch contrast (i.e. a difference in resources among patches) and patch direction (i.e. source–sink relations) are among the basic attributes of spatial patchiness. Here, I hypothesize that young established ramets in nutrient-rich patches support old ramets in nutrient-poor patches when ramets are subjected to different patch contrasts and patch directions.Methods In a greenhouse experiment, old and young ramets of Glechoma longituba were grown in four combinations consisting of patch contrast and patch direction. Minus patch direction refers to a patch combination in which parent ramets grow in nutrient-rich patches while connected daughter ramets grow in nutrient-poor ones and plus patch direction is the opposite direction. I measured photosynthesis and fluorescence traits, harvested all ramets, took morphological measures, weighed their dry mass and determined their nutrient uptake and use.Important findings For parental ramets of G. longituba, patch contrast and patch direction and their interactions had no significant effects on net photosynthetic rate, maximal fluorescence yield, photochemical quenching (quenching refers to any process which decreases the fluorescence intensity of a given substance), non-photochemical quenching, nutrient uptake, biomass and stolon weight ratio. Patch direction alone significantly affected root weight ratio. Large patch contrast enhanced N use efficiency (NUE) and P use efficiency (PUE); plus patch direction decreased NUE, but increased PUE; the patch contrast by patch direction interaction affected PUE and K use efficiency (KUE). There were significant interactions between patch direction and patch contrast on PUE and KUE. It is concluded that soil nutrient patchiness may influence nutrient use strategies, but not nutrient uptake, photosynthesis and growth of parent ramets of G. longituba connected to daughter ramets, and that patch contrast and patch direction jointly affect PUE and KUE.     

Foraging responses of clonal plants to multi-patch environmental heterogeneity: spatial preference and temporal reversibility

Background and aims

Plant root placement is highly plastic in order to acquire patchily distributed nutrients and to ensure their survival, growth and reproduction. Considering the spatial extension of clonal organs, we selected two clonal plants (Leymus chinensis (Trin.) Tzvel. and Hierochloe glabra Trin.) to determine the spatio-temporal effects of environmental heterogeneity on belowground organs and newly-born ramets.

Methods

Small-scale and multi-patch heterogeneous environments were manipulated by creating four patches filled with different types of soil in a same pot. The four patches were composed of sandy soil, sandy loam, loam soil and humus soil, respectively. Ramet number, bud number, mean spacer length, rhizome length, and biomass allocation within each patch were measured to identify plant foraging responses.

Results

The preferential patch of L. chinensis was humus soil patch which was the highest in nutrient availability, whereas H. glabra preferred to place ramets in sandy loam and loam soil patches. When growing in homogeneous environments, both species randomly rooted their offspring ramets in the four compartments. In heterogeneous environments, foraging responses were detected in ramet placement, aboveground biomass and total rhizome length. However, there were no differences in bud number or belowground biomass among four types of patches in heterogeneous environments, which might suggest that there would be no inter-patch differences in seedling establishment in the next year.

Conclusions

Plants show selective allocation of offspring ramets to preferential patches in the presence of multi-patch environmental heterogeneity. Responses of H. glabra to multi-patch heterogeneity were faster than those of L. chinensis, demonstrating that the foraging patterns are species-specific. Clonal plants can rapidly respond to environmental heterogeneity, whereas foraging responses are potentially reversible over a longer temporal scale.     

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.     

Buffalograss decreases ramet propagation in infertile patches to enhance interconnected ramet proliferation in fertile patches

Buffalograss (Buchloë dactyloides) is known for its low-nutrient tolerance. However, in natural habitats, nutrients are usually patchily distributed. For clonal plants like buffalograss, physiological integration is an important strategy to cope with adverse environmental conditions. In order to examine how integration helps buffalograss to survive in patchy conditions, a greenhouse experiment was conducted for 91 days. Interconnected ramet pairs of stoloniferous buffalograss were planted in two partitioned same-sized containers, and subjected to identical or contrasting nutrient supply. In contrast to normally perceived resource-sharing concepts, results showed that buffalograss genets reduced production of new ramets in nutrient-poor patches promoting at the same time propagation of interconnected ramets in nutrient-rich patches. Ramets in nutrient-rich patches gained significant benefit from heterogeneous treatments, whereas nutrient-poor ramets performed even worse than in uniform low-nutrient treatment. Younger ramets developed more biomass than elder ramets with the same amounts of nutrient supply under homogeneous treatment, while elder ramets were more tolerant when nutrients were scarce. Heterogeneity had a particular strong effect on stolons and new ramet production in nutrient-rich patches. Rooted ramets in nutrient-poor patches suffered from a by-pass of nutrients to interconnected ramets on nutrient-rich substrate that probably resulted from different transpiration rates. We conclude that this resource-sharing strategy is advantageous for buffalograss to concentrate more ramets in fertile patches, and facilitate the survivorship of more buffalograss ramets in adverse environments with uneven nutrient supply.     

Root growth and plant biomass in <Emphasis Type="Italic">Lolium perenne</Emphasis> exploring a nutrient-rich patch in soil

We investigated soil exploration by roots and plant growth in a heterogeneous environment to determine whether roots can selectively explore a nutrient-rich patch, and how nutrient heterogeneity affects biomass allocation and total biomass before a patch is reached. Lolium perenne L. plants were grown in a factorial experiment with combinations of fertilization (heterogeneous and homogeneous) and day of harvest (14, 28, 42, or 56 days after transplanting). The plant in the heterogeneous treatment was smaller in its mean total biomass, and allocated more biomass to roots. The distributions of root length and root biomass in the heterogeneous treatment did not favor the nutrient-rich patch, and did not correspond to the patchy distribution of inorganic nitrogen. Specific root length (length/biomass) was higher and root elongation was more extensive both laterally and vertically in the heterogeneous treatment. These characteristics may enable plants to acquire nutrients efficiently and increase the probability of encountering nutrient-rich patches in a heterogeneous soil. However, heterogeneity of soil nutrients would hold back plant growth before a patch was reached. Therefore, although no significant selective root placement in the nutrient-rich patch was observed, plant growth before reaching nutrient-rich patches differed between heterogeneous and homogeneous environments.     

Do Physiological Integration and Soil Heterogeneity Influence the Clonal Growth and Foraging of <Emphasis Type="Italic">Schoenoplectus pungens</Emphasis>?

Physiological integration and foraging behavior have both been proposed as advantages for clonal growth in heterogeneous environments. We tested three predictions concerning their short- and long-term effects on the growth of the clonal perennial sedge Schoenoplectus pungens (Pers.) Volk. ex Schinz and R. Keller: (1) growth would be greatest for clones with connected rhizomes and on heterogeneous soil, (2) clones would preferentially place biomass in the nutrient-rich patches of a spatially heterogeneous environment, and (3) physiological integration would decrease a clone’s ability to forage. We tested our predictions by growing S. pungens clones for 2 years in an experimental garden with two severing treatments (connected and severed rhizomes) crossed with two soil treatments (homogeneous and heterogeneous nutrient distribution). Severing treatments were only carried out in the first year. As predicted, severing significantly decreased total biomass and per capita growth rate in year one and individual ramet biomass both in year one and the year after severing stopped. This reduction in growth was most likely caused by severing damage, because the total biomass and growth rate in severed treatments did not vary with soil heterogeneity. Contrary to our prediction, total biomass and number of ramets were highest on homogeneous soil at the end of year two, regardless of severing treatment, possibly because ramets in heterogeneous treatments were initially planted in a nutrient-poor patch. Finally, as predicted, S. pungens concentrated ramets in the nutrient-rich patches of the heterogeneous soil treatment. This foraging behavior seemed enhanced by physiological integration in the first year, but any possible enhancement disappeared the year after severing stopped. It seems that over time, individual ramets become independent, and parent ramets respond independently to the conditions of their local microsite when producing offspring, a life-history pattern that may be the rule for clonal species with the spreading “guerrilla” growth form.     

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.