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.     

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.     

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     

The effects of clonal integration on morphological plasticity and placement of daughter ramets in black locust (Robinia pseudoacacia)

We studied the field response of Robinia pseudoacacia L. to light, total soil nitrogen, available soil phosphorus and soil pH. Results indicated that there was very strong clonal integration between mother and daughter ramets. Mother ramets can provide nitrogen and phosphorus to daughter ramets sufficient for their continued growth through strong clonal integration, but cannot provide enough photosynthate. With clonal integration, soil nitrogen and phosphorus availability had no effect on biomass allocation to roots, number of ramets and length of connection roots. Biomass allocation to roots increased markedly and responded to nitrogen and phosphorus availability, when the connections were severed. Light had a significant effect on the percent of biomass allocation to leaves and number of ramets, but no effect on the length of connection roots. Daughter ramets allocated more resources to leaves, and clones placed more daughter ramets in high light patches than in low light patches. Soil pH had a significant effect on ramet number and connection root length. Clones concentrated in alkaline patches and escaped from acid patches through selective placement of daughter ramets and changing the length of connection roots. We suggest that the clonal integration may be very strong and provide sufficient soil resources to daughter ramets, then affect the daughter ramets’ morphology and placement, if the size of a specific ramet is significantly larger than the other ramets in an arbor clone.     

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匍匐茎草本植物克隆内资源共享和分株功能特化(英文)

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 (PotentillareptansL .var.sericophyllaFranch .)、鹅绒委陵菜 (P .anserinaL .)和金戴戴 (Halerpestesruthenica (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     

Physiological integration impacts nutrient use and stoichiometry in three clonal plants under heterogeneous habitats

Physiological integration facilitates clonal plants to deal with heterogeneous resources. However, little is known about how nutrient patchiness affects its use and stoichiometry in clonal plants. We conducted an experiment with Cynodon dactylon, Glechoma longituba, and Potentilla reptans to address the effects of physiological integration on nutrient use efficiency and N:P ratios. For C. dactylon, the effects of nutrient patchiness on N use efficiency (NUE), P use efficiency (PUE), and N:P ratio were stronger in daughter ramets than in parent ramets; for G. longituba, nutrient patchiness affected PUE and N:P ratio of parent and daughter ramets, but not NUE; for P. reptans, nutrient patchiness decreased NUE, PUE, and N:P ratio, regardless of parent or daughter ramets. PUE was associated with N:P ratios in three clonal plants and this association of NUE with N:P ratios varied with species. Our findings suggest that physiological integration alters nutrient use efficiency and N:P ratios of clonal plants under patchy nutrients and that these effects are linked to clonal species identity.     

Consequences of ramets helping ramets: No damage and increased nutrient use efficiency in nurse ramets of Glechoma longituba

Source–sink relationships determine the transfer of resources among interconnected ramets of clonal plants and these relationships depend on resource regimes experienced by ramets. We hypothesized that if ramets growing in nutrient-richer patches support ramets in nutrient-poorer patches, then this support would weaken the performance of supporters themselves. This hypothesis was tested in an experiment, in which two interconnected ramet-groups of Glechoma longituba were subjected to four levels of nutrient-patch contrast. For nurse ramets, midday leaf water potential, net photosynthetic rate, fluorescence yield, specific root length, biomass, and root weight ratio remained unchanged along the gradient of patch contrast, indicating that no damage occurs to these ramets. Nitrogen use efficiency and phosphorus use efficiency of nurse ramets significantly increased with increasing patch contrast, suggesting that these increases may allow nurse ramets from habitats higher in patch contrast to have a greater capacity for export of nutrients than those ramets from habitats lower in patch contrast. In addition, we discuss the likely implications of these findings.     

根状茎型植物扁秆荆三棱对土壤养分异质性尺度和对比度的生长响应

土壤养分的空间异质性在自然界普遍存在, 而克隆植物被认为能很好地适应和利用土壤养分异质性。尽管尺度和对比度是异质性的两个重要属性, 但有关土壤养分异质性的尺度和对比度及其交互作用对克隆植物生长和分株分布格局影响的研究仍比较缺乏。在一个温室实验中, 根状茎型草本克隆植物扁秆荆三棱(Bolboschoenus planiculmis) (异名扁秆藨草(Scirpus planiculumis))被种植在由高养分斑块和低养分斑块组成的异质性环境中。实验为两种尺度处理(大斑块和小斑块)和两种对比度处理(高对比度和低对比度)交叉组成的4种处理组合。在每个处理中, 高养分和低养分斑块的总面积相同; 在所有4种处理中, 土壤养分的总量也完全相同。无论在整个克隆(植株)水平, 还是在斑块水平, 尺度、对比度及其交互作用对扁秆荆三棱的生物量、分株数、根状茎长和块茎数的影响均不显著。然而, 在斑块水平, 扁秆荆三棱在高养分斑块中的生物量、分株数、根状茎长和块茎数均显著高于低养分斑块, 而在高养分斑块中相邻分株间的距离(间隔物长)小于低养分斑块, 并且这种效应均不依赖于斑块尺度的大小和对比度的高低。因此, 在土壤养分异质性环境中, 扁秆荆三棱可以通过缩短间隔物长, 并可能通过提高根状茎的分枝强度, 把较多的分株和潜在分株放置在养分条件好的斑块中。这种响应格局体现出克隆植物的觅食行为, 有利于整个克隆对异质性资源的吸收和利用。然而, 该实验中的尺度和对比度对扁秆荆三棱分株的放置格局均没有显著效应。作者推测, 在一个更大的斑块尺度和(或)对比度范围内, 扁秆荆三棱对土壤养分异质性的响应可能不同。因此, 下一步的研究应涉及更广泛的尺度和对比度。     

Clonal integration ameliorates the carbon accumulation capacity of a stoloniferous herb,Glechoma longituba,growing in heterogenous light conditions by facilitating nitrogen assimilation in the rhizosphere

Background and Aims Enhanced availability of photosynthates increases nitrogen (N) mineralization and nitrification in the rhizosphere via rhizodeposition from plant roots. Under heterogeneous light conditions, photosynthates supplied by exposed ramets may promote N assimilation in the rhizosphere of shaded, connected ramets. This study was conducted to test this hypothesis.Methods Clonal fragments of the stoloniferous herb Glechoma longituba with two successive ramets were selected. Mother ramets were subjected to full sunlight and offspring ramets were subjected to 80 % shading, and the stolon between the two successive ramets was either severed or left intact. Measurements were taken of photosynthetic and growth parameters. The turnover of available soil N was determined together with the compostion of the rhizosphere microbial community.Key Results The microbial community composition in the rhizosphere of shaded offspring ramets was significantly altered by clonal integration. Positive effects of clonal integration were observed on NAGase activity, net soil N mineralization rate and net soil N nitrification rate. Increased leaf N and chlorophyll content as well as leaf N allocation to the photosynthetic machinery improved the photosynthetic capability of shaded offspring ramets when the stolon was left intact. Clonal integration improved the growth performance of shaded, connected offspring ramets and whole clonal fragments without any cost to the exposed mother ramets.Conclusions Considerable differences in microbial community composition caused by clonal integration may facilitate N assimilation in the rhizosphere of shaded offspring ramets. Increased N content in the photosynthetic machinery may allow pre-acclimation to high light conditions for shaded offspring ramets, thus promoting opportunistic light capture. In accordance with the theory of the division of labour, it is suggested that clonal integration may ameliorate the carbon assimilation capacity of clonal plants, thus improving their fitness in temporally and spatially heterogeneous habitats.     

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。     

Habitat selection in spatially heterogeneous environments: a test of foraging behaviour in the clonal submerged macrophyte Vallisneria spiralis

1. To test whether clonal macrophytes can select favourable habitats in heterogeneous environments, clonal fragments of the stoloniferous submerged macrophyte Vallisneria spiralis were subjected to conditions in which light intensity and substratum nutrients were patchily distributed. The allocation of biomass accumulation and ramet production of clones to the different patches was examined. 2. The proportion of both biomass and ramet number of clones allocated to rich patches was significantly higher than in poor patches. The greatest values of both clone and leaf biomass were produced in the heterogeneous light treatment, in which clones originally grew from light‐rich to light‐poor patches, while clones produced the most offspring ramets in the treatments with heterogeneous substratum nutrients. Similarly, root biomass had the highest values in nutrient‐rich patches when clones grew from nutrient‐rich to nutrient‐poor patches. 3. The quality of patches in which parent ramets established significantly influenced the foraging pattern. When previously established in rich patches, a higher proportion of biomass was allocated to rich patches, whereas a higher proportion of ramet number was allocated to rich patches when previously established in poor patches. 4. Results demonstrate that the clonal macrophyte V. spiralis can exhibit foraging in submerged heterogeneous environments: when established under resource‐rich conditions V. spiralis remained in favourable patches, whereas if established in adverse conditions it could escape by allocating more ramets to favourable patches.     

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.     

Effects of Clonal Integration on Microbial Community Composition and Processes in the Rhizosphere of the Stoloniferous Herb Glechoma longituba (Nakai) Kuprian

The effects of rhizodeposition on soil C and N availabilities lead to substantial changes of microbial community composition and processes in the rhizosphere of plants. Under heterogeneous light, photosynthates can be translocated or shared between exposed and shaded ramets by clonal integration. Clonal integration may enhance the rhizodeposition of the shaded ramets, which further influences nutrient recycling in their rhizosphere. To test the hypothesis, we conducted a pot experiment by the stoloniferous herb Glechoma longituba subjected to heterogeneous light. Microbial biomass and community composition in the rhizosphere of shaded offspring ramets, assessed by phospholipid fatty acids (PLFAs) analysis, were markedly altered by clonal integration. Clonal integration positively affected C, N availabilities, invertase and urease activities, N mineralization (N_min) and nitrification rates (N_nitri) in the rhizosphere of shaded offspring ramets. However, an opposite pattern was also observed in phenoloxidase (POXase) and peroxidase (PODase) activities. Our results demonstrated that clonal integration facilitated N assimilation and uptake in the rhizosphere of shaded offspring ramets. The experiment provides insights into the mechanism of nutrient recycling mediated by clonal integration.     

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.     

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.     

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.     

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.     

Environmental constraints on the growth,photosynthesis and reproductive development of Dryas octopetala at a high Arctic polar semi-desert,Svalbard

Opportunities exist in high Arctic polar semidesert communities for colonisation of unvegetated ground by long-lived clonal plants such as Dryas octopetala. This can be achieved by lateral spread of vegetative ramets, or by sexual reproduction and seedling recruitment. The objectives of this study were (1) to determine whether these two means of proliferation show differential sensitivity to contrasting components of the abiotic environment (temperature, soil nutrient and water availability) and (2) to evaluate the potential for D. octopetala to respond to climate change by an increase in cover and biomass in polar semi-desert communities. Factorial environmental manipulations of growing season temperature, soil nutrient and water status were conducted over 3 years at a polar semi-desert community in Svalbard, Norway (78°56.12N, 11°50.4E) and both clonal and sexual reproductive performance, together with instantaneous net photosynthesis (P_n), were recorded during the third season (1993). D. octopetala capitalised rapidly on an amelioration in the availability of inorganic nutrients (N, P and K) by an expansion in leaf area and biomass supported by increased P_n per unit leaf weight, and by apparent luxury uptake of nutrients (particularly P). Several facets of sexual reproductive development and seed viability were markedly improved by elevated temperatures or soil nutrient availability. Thus although D. octopetala is a long-lived clonal plant, with many traits characteristic of stress resistance syndrome, it showed considerable phenotypic plasticity in response to environmental manipulations. The results support the hypothesis that clonal growth confers survival potential during unfavourable years, together with the ability to capitalise on nutrient flushes and recycle nutrients internally. Continued investment in sexual reproduction ensures that seed setting is successful during favourable years, even if these occur infrequently. Cimate warming in the high Arctic could thus result in changes in the cover, biomass and the relative significance of clonal versus sexual proliferation of D. octopetala (and thus the genetic diversity of the population) but the long-term responses will probably be constrained by lack of available nutrients.