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     

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)     

Modelling the morphological response to nutrient availability in the clonal plant Trientalis europaea L.

The morphological responses to changes in environmental quality shown by many clonal plants have been interpreted as an expression of foraging behaviour, as they allow the ramets to become concentrated in more favourable microhabitats. The morphological response to increased nutrient availability in the pseudoannual plant Trientalis europaea was studied in a field experiment. The response was largely size-dependent and consistent with enhanced clonal growth. Fertilized ramets produced more tubers and a larger main tuber. In contrast, stolon length was not affected by the treatment. A spatially explicit simulation model calibrated with data from the field experiment examined the population dynamics of T. europaea ramets in a spatially hetereogeneous, temporally constant, environment. The model showed that T. europaea was effective at concentrating its ramets in favourable patches, but this process was strongly influenced by patch size. The analysis of this response at the clone level showed that ramet aggregation was mainly due to the enhanced performance of clones located initially in the favourable patches, or clones that located a favourable patch by chance. In these clones, the simultaneous increase of ramet size and survival accelerated the production of ramets. The temporal and spatial scale at which the aggregation of ramets in favourable patches was manifested suggests that the effectiveness of the morphological response in T. europaea is favoured by a high spatio-temporal predictability in the environment. Overall, the model emphasized the important role of population dynamics in understanding the nature of the foraging response.     

Length of the Spacer Rather than its Plasticity Relates to Species Distribution in Various Natural Habitats

Much evidence from laboratory experiments and theoretical studies show that plasticity of clonal growth traits like lateral spread provides advantage in heterogeneous conditions. However, few tests of whether species with plastic clonal growth have an advantage over non-plastic species in natural conditions exist. I analyzed whether spacer length (i.e., length of a stolon or a rhizome branch between two ramets) and the variation of it (as a surrogate to its plasticity) relate to species presence and abundance in open meadow, wooded meadow and forest habitats with varying environmental conditions at Laelatu wooded meadow, Estonia. In fertilized, compared to unfertilized conditions, a weak and non-conclusive advantage of both long spacers and high variability of spacer length was detected. Abandonment in open meadows lead to a prevalence of species with shorter spacers, while on abandoned wooded meadows the species with longer spacers dominated. There was no difference in variability of spacer length between managed and abandoned meadows. In more heterogeneous forest habitats high variability of spacer length was more common, but the effect of lengthy spacers was more pronounced. The results suggest that while the variability of spacer length indeed corresponds with larger abundance in some conditions, the actual length of spacers has a more pronounced relationship with abundance of plants in natural vegetation.     

Small-scale heterogeneity in soil quality influences photosynthetic efficiency and habitat selection in a clonal plant

BACKGROUND AND AIMS: In clonal plants, internode connections allow translocation of photosynthates, water, nutrients and other substances among ramets. Clonal plants form large systems that are likely to experience small-scale spatial heterogeneity. Physiological and morphological responses of Fragaria vesca to small-scale heterogeneity in soil quality were investigated, together with how such heterogeneity influences the placement of ramets. As a result of their own activities plants may modify the suitability of their habitats over time. However, most experiments on habitat selection by clonal plants have not generally considered time as an important variable. In the present study, how the foraging behaviour of clonal plants may change over time was also investigated. METHODS: In a complex of environments with different heterogeneity, plant performance was determined in terms of biomass, ramet production and photosynthetic activity. To identify habitat selection, the number of ramets produced and patch where they rooted were monitored. KEY RESULTS: Parent ramets in heterogeneous environments showed significantly higher maximum and effective quantum yields of photosystem II than parents in homogeneous environments. Parents in heterogeneous environments also showed significantly higher investment in photosynthetic biomass and stolon/total biomass, produced longer stolons, and had higher mean leaf size than parents in homogeneous environments. Total biomass and number of offspring ramets were similar in both environments. However, plants in homogeneous environments showed random allocation of offspring ramets to surrounding patches, whereas plants in heterogeneous environments showed preferential allocation of offspring to higher-quality patches. CONCLUSIONS: The results suggest that F. vesca employs physiological and morphological strategies to enable efficient resource foraging in heterogeneous environments and demonstrate the benefits of physiological integration in terms of photosynthetic efficiency. The findings indicate that short-term responses cannot be directly extrapolated to the longer term principally because preferential colonization of high-quality patches means that these patches eventually show reduced quality. This highlights the importance of considering the time factor in experiments examining responses of clonal plants to heterogeneity.     

Responses to mineral nutrient availability and heterogeneity in physiologically integrated sedges from contrasting habitats

Clonal plants from poor habitats benefit less from morphologically plastic responses to heterogeneity than plants from more productive sites. In addition, physiological integration has been suggested to either increase or decrease the foraging efficiency of clonal plants. We tested the capacity for biomass production and morphological response in two closely related, rhizomatous species from habitats that differ in resource availability, Carex arenaria (from poor sand dunes) and C. disticha (from nutrient-richer, moister habitats). We expected lower total biomass production and reduced morphological plasticity in C. arenaria, and that both species would produce more ramets in high nutrient patches, either in response to signals transported through physiological integration, or by locally determined responses to nutrient availability. To investigate mineral nutrient heterogeneity, plants were grown in boxes divided into two compartments with homogeneous or heterogeneous supply of high (H) or low (L) nutrient levels, resulting in four treatments, H-H, H-L, L-H and L-L. Both C. arenaria and C. disticha produced similar biomass in high nutrient treatments. C. disticha responded to high nutrients by increased biomass production and branching of the young parts and by altering root:shoot ratio and rhizome lengths, while C. arenaria showed localised responses to high nutrients in terms of local biomass and branch production in high nutrient patches. The results demonstrated that although it has a conservative morphology, C. arenaria responded to nutrient heterogeneity through morphological plasticity. An analysis of costs and benefits of integration on biomass production showed that young ramets of both species benefited significantly from physiological integration, but no corresponding costs were found. This suggests that plants from resource-poor but dynamic habitats like sand dunes respond morphologically to high nutrient patches. The two species responded to nutrient heterogeneity in different traits, and this is discussed in terms of local and distant signalling of plant status.     

Effects of clonal integration on plant plasticity in Fragaria chiloensis

The ability of clonal plants to transport substances between ramets located in different microsites also allows them to modify the plastic responses of individual ramets to local environmental conditions. By equalising concentrations of substances between ramets, physiological integration might decrease responses to local conditions. However, integration has also been observed to increase plasticity and induce novel plastic responses in ramets. To ask how integration modifies plant plasticity in the clonal herb, Fragaria chiloensis, ramets were given either low light and high nitrogen or high light and low nitrogen, simulating a pattern of resource patchiness in their native habitat. Ramets in contrasting light/nitrogen treatments were either connected or single. Effects of light/nitrogen and connection were measured at three levels of morphological organisation, the organ, the ramet, and the clonal fragment. Connection between ramets reduced or had no effect on plastic responses in leaf size at the level of the plant organ. This suggested that integration dampened certain plastic responses. Connection induced a new plastic response at the level of the clonal fragment, an increase in allocation to vegetative reproduction in patches of low light and high nitrogen. It is concluded that clonal integration can have different effects on plant plasticity at different levels of plant organisation. It appears that, at least in this species, integration can increase plasticity at the level of the clonal fragment and concentrate vegetative reproduction in particular microsite types.     

Clonal plant allocation to daughter ramets is a simple function of parent size across species and nutrient levels

The evolution of clonal growth is a widespread phenomenon among plant species, characterized by the production of genetically identical clonal fragments (ramets) via rhizomes or stolons that form an interconnected clonal organism (genet). Clonal plant species are known to differ in their investment into ramet production, and exhibit considerable variation in ramet morphology both within and among species. While patterns of resource allocation are thought to be linked to a number of plant characteristics, many analyses are limited by uncertainty in how clonal plants determine the morphology and resources allocated to new ramets. In this study, we attempted to discern what aspects of parent ramets best predicted resource allocation to new daughter ramets, and the relationship between resource allocation and daughter ramet rhizome morphology. We grew two sedge species, Schoenoplectus tabernaemontani and Eleocharis elliptica, in a greenhouse under two levels of fertilizer addition. By harvesting daughter ramets that had initiated stem production, yet remained aphotosynthetic, we were able to isolate parental investment into non-independent daughter ramets at a point where daughter ramet spacer length became fixed. Our results indicate that parent ramets allocated a non-linear proportion of parent rhizome biomass to the production of daughter ramets. Moreover, this relationship was unaffected by environmental nutrient availability. Daughter ramet biomass, in turn, was strongly correlated with daughter ramet spacer length. These observations shed light on key processes governing clonal growth in plants, and their potential application in unifying allocational and morphological perspectives to explore the fitness implications of variability in clonal growth.     

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。     

Costs of plasticity in foraging characteristics of the clonal plant Ranunculus reptans

In clonal plants, evolution of plastic foraging by increased lengths of leaves and internodes under unfavourable conditions may be constrained by costs and limits of plasticity. We studied costs and limits of plasticity in foraging characteristics in 102 genotypes of the stoloniferous herb Ranunculus reptans. We grew three replicates of each genotype with and three without competition by the naturally co-occuring grass Agrostis stolonifera. We used regression and correlation analyses to investigate potential costs of plasticity in lengths of leaves and stolon internodes, developmental instability costs of these traits, and a developmental range limit of these traits. We used randomization procedures to control for spurious correlations between parameters calculated from the same data. Under competition the number of rosettes, rooted rosettes, and flowers was 58%, 40%, and 61% lower, respectively, than in the absence of competition. Under competition lengths of leaves and stolon internodes were 14% and 6% smaller, respectively, than in the absence of competition. We detected significant costs of plasticity in stolon internode length in the presence of competition when fitness was measured in terms of the number of rosettes and the number of flowers (selection gradients against plasticity were 0.250 and 0.214, respectively). Within-environment variation (SD) in both foraging traits was not positively correlated with the corresponding plasticity, which indicates that there were no developmental instability costs. More plastic genotypes did not have less extreme trait values than less plastic genotypes for both foraging traits, which indicates that there was no developmental range limit. We conclude that in R. reptans costs of plasticity more strongly constrain evolution of foraging in the horizontal plane (i.e., stolon internode length) than in the vertical plane (i.e., leaf length).     

Multivariate selection shapes environment-dependent variation in the clonal morphology of a red seaweed

Within-individual strategies of variation (e.g., phenotypic plasticity) are particularly relevant to modular organisms, in which ramets of the same genetic individual may encounter diverse environments imposing diverse patterns of selection. Hence, measuring selection in heterogeneous environments is essential to understanding whether environment-dependent phenotypic change enhances the fitness of modular individuals. In sublittoral marine habitats, competition for light and space among modular taxa generates extreme patchiness in resource availability. Little is known, however, of the potential for plasticity within individuals to arise from spatially-variable selection in such systems. We tested whether plasticity enhances genet-level fitness in Asparagopsis armata, a clonal seaweed in which correlated traits mediate morphological responses to variation in light. Using the capacity for rapid, clonal growth to measure fitness, we identified aspects of ramet morphology targeted by selection in two contrasting light environments and compared patterns of selection across environments. We found that directional selection on single traits, coupled with linear and nonlinear selection on multi-trait interactions, shape ramet morphology within environments and favor different phenotypes in each. Evidence of environment-dependent, multivariate selection on correlated traits is novel for any marine modular organism and demonstrates that seaweeds, such as A. armata, may potentially adapt to environmental heterogeneity via plasticity in clonal morphology.     

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。     

Morphological responses to local light conditions in clonal herbs from contrasting habitats,and their modification due to physiological integration

Morphological responses to light and effects of physiological integration on local morphological responses are examined for Hydrocotyle vulgaris and Lamiastrum galeobdolon, stoloniferous herbs from open fenlands and forest understoreys, respectively. An assessment was made of whether these clonal herbs of similar morphology but from contrasting habitats show different foraging behaviour for light. In a garden experiment, the plants wer subjected to four levels of light availability, and to a split treatment in which the primary stolons grew along the border of patches of the two intermediate light levels. In this treatment the plant parts on opposite sides of the primary stolons were in contrasting light environments. Petiole extension was more responsive to light conditions in Hydrocotyle than in Lamiastrum, while the opposite was true for leaf area. Both species showed similar responses in stolon internode length and specific leaf area (SLA). Integration did not significantly modify local responses in stolon internode length in either species. Local responses in petiole length, leaf area and SLA of Hydrocotyle ramets were not significantly affected by physiological integration, except for the SLA of ramets in high light which was evened out by integration. In contrast, in Lamiastrum, local responses in petiole length, leaf area and SLA of many ramets in the shaded and/or light patch were significantly evened out by integration. As a result, interconnected ramets in patches of different light supply developed very different morphologies in Hydrocotyle, but not in Lamiastrum. The results indicate that the species differed in ramet morphological responses to light intensity as well as in effects of integration on local morphological responses, and suggest that species from different habitats show different foraging behaviour for light.     

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。     

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.     

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.     

How past and present influence the foraging of clonal plants?

Clonal plants spreading horizontally and forming a network structure of ramets exhibit complex growth patterns to maximize resource uptake from the environment. They respond to spatial heterogeneity by changing their internode length or branching frequency. Ramets definitively root in the soil but stay interconnected for a varying period of time thus allowing an exchange of spatial and temporal information. We quantified the foraging response of clonal plants depending on the local soil quality sampled by the rooting ramet (i.e. the present information) and the resource variability sampled by the older ramets (i.e. the past information). We demonstrated that two related species, Potentilla reptans and P. anserina, responded similarly to the local quality of their environment by decreasing their internode length in response to nutrient-rich soil. Only P. reptans responded to resource variability by decreasing its internode length. In both species, the experience acquired by older ramets influenced the plastic response of new rooted ramets: the internode length between ramets depended not only on the soil quality locally sampled but also on the soil quality previously sampled by older ramets. We quantified the effect of the information perceived at different time and space on the foraging behavior of clonal plants by showing a non-linear response of the ramet rooting in the soil of a given quality. These data suggest that the decision to grow a stolon or to root a ramet at a given distance from the older ramet results from the integration of the past and present information about the richness and the variability of the environment.     

Two types of division of labour in clonal plants: benefits, costs and constraints

Clonal plants spread vegetatively within their habitats by forming rooted ramets on stolons or rhizomes. Each of these ramets is capable of an independent existence after establishment. Nevertheless, ramets remain physically connected by stolon or rhizome internodes for variable periods of time, thereby allowing for resource movement and signal transduction within clones.Interconnected ramets of clonal plants, though potentially independent and totipotent, can specialize functionally in the performance of limited numbers of tasks such as the uptake of resources from above- vs below-ground sources, carbohydrate storage, vegetative spread and sexual reproduction. Such specialization and cooperation is comparable to a division of labour in economic systems or in colonies of social animals. The ecological significance of division of labour in clonal plants may be found in the increased efficiency of entire clones in exploiting their environments.Two different types of division of labour in clonal plants will be discussed in this review. The first type is an environmentally-induced specialization of ramets in the uptake of locally abundant resources (plastic division of labour), which can be found in several stoloniferous species. Evidence exists that this response increases resource uptake in spatially heterogeneous environments. The second type of division of labour, which occurs mainly in rhizomatous species, relates to a developmentally-programmed specialization and cooperation between interconnected ramets. This response pattern is thought to enhance plant performance by restricting the number of tasks for individual ramets and thereby significantly increasing the efficiency of task performance. In some plants, such an inherent division of labour is likely to contribute to nutrient extraction from poor and unpredictably variable sources.In this article not only benefits but also potential costs and constraints on division of labour in clonal plants are shown. The aim is to provide a review of existing knowledge and to develop concepts and hypotheses for future research.     

林冠环境对亚高山针叶林下缺苞箭竹生物量分配和克隆形态的影响

通过对亚高山针叶林的林下、小林窗(130 m²)、中林窗(300 m²)和大林窗(500 m²) 4种林冠环境中缺苞箭竹(Fargesia denudata)分株种群特征进行调查, 研究其生物量分配格局和克隆形态可塑性。研究结果表明: (1)分株生物量、基径、高度以及分株各构件生物量随林冠郁闭度减小均表现为先增加后减小的趋势, 在小林窗中达到最大值; (2)大林窗中, 根生物量分配和数量显著高于其他林冠环境, 随林冠郁闭度增大, 分株不断增大叶生物量分配、比叶面积和分枝百分比等地上部分投资, 以适应低光环境; (3)比茎长和比地下茎长随林冠郁闭度增大表现为先减小后增加的趋势, 在小林窗值最低, 分枝强度在小林窗和中林窗中显著高于林下和大林窗环境。结果显示, 缺苞箭竹在不同林冠环境中具有生物量分配和克隆形态的可塑性, 以利于种群对光资源的有效利用。小林窗环境是缺苞箭竹较适宜的生境, 生物量积累最多, 长势最好。