An analysis of the costs and benefits of physiological integration between ramets in the clonal perennial herb Glechoma hederacea

Summary The costs and benefits, measured in terms of dry weight, of physiological integration between clonal ramets, were analysed in two experiments conducted on the clonal herb Glechoma hederacea. Firstly, integration between consecutively-produced ramets was examined in an experiment in which stolons grew from one set of growing conditions (either unshaded or shaded and either nutrient-rich or nutrient-poor) into conditions in which light or nutrient level was altered. Comparisons were made between the dry weight of the parts of the clones produced before and after growing conditions were changed, and the dry weights of the corresponding part of control clones subjected to constant growing conditions. In a second experiment, integration between two distinct parts of G. hederacea clones was investigated. In this experiment clones were grown from two connected parent ramets and the parts of the clone produced by each parent ramet were subjected independently to either nutrient-rich or nutrient-poor conditions. Ramets in resource-rich conditions provided considerable physiological support to those in resource-poor conditions. This was measured as a dry weight gain compared with the weight of the corresponding part of the control clones growing in resource-poor conditions. However, when stolons grew from resource-poor conditions into resource-rich conditions, there was no similar evidence of the resourcepoor ramtes receiving support from resource-rich ramets. Physiological integration did not result in dry weight gains when this would have necessitated basipetal translocation of resources.Because of the predominantly acropedal direction of movement of translocates in G. hederacea, the structure of the clone was important in determining the effectiveness of integration between ramets. Where physiological integration was effective, the cost to the supporting ramets in terms of dry weight was insignificant. Physiological integration allows clones to maintain a presence in less favourable sites with insignificant cost to ramets in favourable sites, thereby reducing the probability of invasion by other plants, and providing the potential for rapid clonal growth if conditions improve. Integrated support of ramets in unfavourable conditions also enables the clone to grow through unfavourable sites, thus increasing the probability of encountering more favourable conditions by wider foraging.     

Performance of a clonal species in patchy environments: effects of environmental context on yield at local and whole-plant scales

Yield of the clonal plant Glechoma hederacea was compared at different spatial scales, in heterogeneous and homogeneous environments providing the same amount of nutrients. For the heterogeneous treatments, environments were created with different patch sizes and different degrees of contrast in nutrient concentration between patches of different quality. Total clone yield differed by almost 2.5-fold across treatments, being highest in environments with large patches and high contrast, lowest in environments with small patches and high contrast, and intermediate under homogeneous conditions. Compared with plants in homogeneous conditions, there were significant increases or decreases in yield at all scales of measurement in many of the heterogeneous treatments. These effects on yield reflected a combination of local responses to growing conditions and modification of these responses due to physiological integration with other parts of the plant growing in contrasting conditions, supporting the proposal of de Kroon et al. (2005 New Phytol 166:73–82). The results show that plant yield at all scales is strongly dependent on environmental context, and that maximum yield can only be realized under a limited range of heterogeneous conditions.     

Clonal integration is beneficial for resource sharing in a creeping amphibian herb (<Emphasis Type="Italic">Alteranthera philoxeroides</Emphasis>)

Clonal integration facilitates the growth and reproduction of clonal plants by providing the ability to share resources among ramets in heterogeneous environments. The benefits of clonal integration for plant growth may depend on a contrast in resource availability and may encounter costs, especially when a young part of the clone is growing across a border between richer and poorer conditions than the old part. We studied a clonal amphibian plant growing across a border between an aquatic and a terrestrial ecosystem, which typically differ in the availability of resources. We asked whether the young part of the clone is supporting the old part with phosphorus and whether this support has costs. We performed an experiment with Alternanthera philoxeroides where plants grow from water to a terrestrial habitat. The terrestrial habitat had either a low or high phosphorus supply, and the connection between the old and young parts of the clone was either left intact or split. We determined that the young part of the clone growing in a terrestrial habitat supported the old part with phosphorus when growing on a substrate rich in phosphorus. We have found no cost of this resource translocation; on the contrary, whole clones increased not only their accumulation of phosphorus, but also of nitrogen. Our study shows how an amphibian plant may profit from heterogeneous habitats by resource sharing in a clonal network.     

异质水分环境下野牛草相连分株间光合同化物的生理整合及其调控

异质环境下,克隆植物通过生理整合机制使资源在分株间实现共享,提高了其对异质性环境的适应能力,具有重要的生态进化意义,研究生理整合机制及其调控机理可为进一步发掘克隆植物应用潜力提供理论依据。以野牛草3个相连分株为材料,对其中一个分株用30%聚乙二醇6000(PEG-6000)模拟水分胁迫,通过Hoagland营养液培养试验,研究了异质水分环境下光合同化物在野牛草相连分株间的生理整合及分株叶片与根系内源激素ABA与IAA含量的变化规律。结果表明,14C-光合同化物在克隆片断内存在双向运输,但以向顶运输为主,异质水分环境下,受胁迫分株光合同化物的输出率明显降低,而与其相邻分株合成的光合同化物向受胁迫分株方向运输率明显增加;异质水分环境下,各分株ABA含量均明显增加,但以受胁迫的分株叶片及根系ABA的含量增加幅度最大,各分株IAA含量较对照均显著下降(P0.05),且以受胁迫分株IAA含量下降幅度最大;各分株叶片与根系ABA/IAA均显著提高(P0.05),相邻分株ABA/IAA增加幅度低于受胁迫分株。异质水分环境影响野牛草克隆分株间光合同化物的生理整合,且ABA与IAA在分株间光合同化物运输与分配过程中具有重要的调节作用。     

Integration in the clonal plant <Emphasis Type="Italic">Eriophorum angustifolium</Emphasis>: an experiment with a three-member-clonal system in a patchy environment

A clonal plant in heterogeneous environments is usually expected to profit from resource exchange via a clonal network where ramets placed in contrasting environments can specialise so to acquire the most abundant resources. An experiment was designed using the three member clonal system of Eriophorum angustifolium, which consisted of one parent ramet growing in a resource poor environment and two offspring: one was limited in growth by nutrients while the other was light limited; the contrast in availability of limited resources between the offspring ramets was high, medium or none, with the system either connected or severed. The total resource availability was the same in all treatments. We proposed four possible scenarios for the system: offspring ramets will share resources via the deficient parent ramet, and the whole clone will profit from the contrasting environment (scenario 1); offspring ramets will support exclusively the parent ramet, and the whole clone will profit from a homogeneous environment (scenario 2); offspring ramets will stop the export of the limiting resource to the parent ramet, with split and connected treatments not differing (scenario 3); and offspring ramets will exhaust the carbon stored in the biomass of the parental ramet; offspring ramet will profit from connection (scenario 4). In the experiment, the limiting resources were sent to the strongest sink (scenario 2). The parent ramet growing in a deficient environment received the highest support in the treatment where both offspring ramets were growing in the same conditions (no-contrast treatment). Production of new shoots, but not biomass of whole clone, was supported in a homogenous environment. The experiment revealed that multiple stresses might prohibit free exchange of limiting resources via the clonal network and supports the idea that experimental studies on more complex clones are essential for understanding the costs and benefits of clonal growth.     

Spatial division of labour of Schoenoplectus americanus

If connected ramets are growing in heterogeneous environments, Division of Labour (DoL) among ramets potentially will result in more efficient sharing of resources and an overall benefit to the plants. As a result of DoL, connected ramets growing in a heterogeneous environment might achieve more biomass than ramets growing in a homogeneous environment. DoL has been demonstrated to occur in a few clonal plant species, although most studies simply focussed on biomass allocation, not on actual resource capturing such as water and nutrient consumption. The model system for our study is one in which two connected ramet groups of Schoenoplectus americanus were placed into contrasting environments. In one treatment, the connected ramets grew in heterogeneous environments and in the other treatment, the connected ramets grew in the same (i.e. homogeneous) environment. We manipulated two variables (light and salinity) in the experiment. We hypothesized that ramets growing in a shaded and fresh water condition in a heterogeneous environment would use more water than ramets growing in a similar condition but in a homogeneous environment. We further hypothesized that ramets growing in a light and saline condition in a heterogeneous environment would assimilate less water than ramets growing in a similar condition but in a homogeneous environment. These hypotheses are based on the assumption that ramets in a heterogeneous environment would translocate water from ramets growing in a shaded and fresh water condition to ramets growing in a light and saline water condition. We also hypothesized that ramets growing in heterogeneous environments achieve larger biomass than ramets in homogeneous environments. Ramets grown in light and saline conditions in heterogeneous environments allocated more biomass to aboveground parts, had taller shoots, larger Specific Green (leaf) Area and consumed less water than ramets grown in similar conditions but in a homogeneous environment. Results confirm the hypothesis that connected ramets in heterogeneous environments are specialised to capture locally abundant resources and share these with connected ramets growing in other habitats. The result of DoL is that the entire connected ramet system benefits and achieves higher biomass.     

Clonal integration affects allocation in the perennial herb <Emphasis Type="Italic">Alternanthera philoxeroides</Emphasis> in N-limited homogeneous environments

Most work on clonal growth in plants has focused on the advantages of clonality in heterogeneous habitats. We hypothesized (1) that physiological integration of connected ramets within a clone can also increase plant performance in homogeneous environments, (2) that this effect depends on whether ramets differ in ability to take up resources, and (3) that only ramets with relatively low uptake ability benefit. We tested these hypotheses using the perennial amphibious herb Alternanthera philoxeroides. We grew clonal fragments and varied numbers of rooted versus unrooted ramets, connection between the apical and basal parts of fragments, and availability of nitrogen. Patterns of final size and mass of fragments did not support these hypotheses. By some measures, severance did reduce the growth of more apical ramets and increase the growth of less apical ones, consistent with net apical transfer of resources. Rooting of individual ramets strongly influenced their growth: second and third most apical ramets each grew most when they were the most apical rooted ramet, and this pattern was more pronounced under higher nitrogen levels. This adds to the evidence that signalling between ramets is an important aspect of clonal integration. Overall, the results indicate that physiological integration between ramets within clones in homogeneous environments can alter the allocation of resources between connected ramets even when it does not affect the total growth of clonal fragments.     

Combined effects of resource heterogeneity and simulated herbivory on plasticity of clonal integration in a rhizomatous perennial herb

Previous lines of investigation assuming potential advantage of clonal integration generally have neglected its plasticity in complex heterogeneous environments. Clonal plants adaptively respond to abiotic heterogeneity (patchy resource distribution) and herbivory‐induced heterogeneity (within‐clone heterogeneity in ramet performance), but to date little is known about how resource heterogeneity and simulated herbivory jointly affect the overall performance of clones. Partial damage within a clone caused by herbivory might create herbivory‐induced heterogeneity in a resource‐homogeneous environment, and might also decrease or increase the extent of heterogeneity under resource‐heterogeneous conditions. We conducted a greenhouse experiment in which target‐ramets of Leymus chinensis segments within homogeneous or heterogeneous nutrient treatments were subject to clipping (0% or 75% shoot removal). In homogeneous environments with high (9:9) nutrient availability, ramet biomass of L. chinensis with intact or severed rhizomes is 0.70 or 0.69 g. Conversely, target‐ramet biomass with intact rhizomes is obviously lower than that of the severed target‐ramets in the homogeneous environments with medium (5:5) and low (1:1) nutrient availability. High resource availability and the presence of herbivory can alleviate negative effects of rhizome connection under homogeneous conditions, by providing copious resource or creating herbivory‐induced heterogeneity respectively. Herbivory tolerance of clonal fragments with connected rhizomes was higher than that of fragments with severed rhizomes under heterogeneous conditions. These findings confirmed the unconditional advantage of clonal integration on reproduction under the combined influence of resource heterogeneity and simulated herbivory. Moreover, our results made clear the synergistically interactive effects of resource heterogeneity and simulated herbivory on costs and benefits of clonal integration. This will undoubtedly advance our understanding on the plasticity of clonal integration under complex environmental conditions.     

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.     

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 resource heterogeneity on nitrogen translocation within clonal fragments of Sasa palmata: an isotopic (15N) assessment

BACKGROUND AND AIMS: Clonal fragments of the rhizomatous dwarf bamboo Sasa palmata, which widely predominates in temperate regions of Japan, were grown under heterogeneous resource conditions such as gap understories or nutrient-patchy grassland. Clonal fragments develop multiple ramets with long rhizomes and appear to be physiologically integrated by the translocation of assimilates. The glasshouse experiment reported here was designed to clarify the mechanisms of physiological integration of nitrogen more precisely. METHODS: To assess how resource conditions influence the amount of nitrogen translocation, and which organ acts as the strongest sink, two experiments were conducted that traced movement of 15N label between interconnected pairs of ramets to compare homogeneous and heterogeneous light and soil nitrogen conditions. KEY RESULTS: The amount of 15N translocated to leaves was between 9% and 11% greater in high-N and high-light ramets in the heterogeneous compared with homogeneous treatments. Under heterogeneous soil nitrogen conditions, translocation increased from individual ramets in resource-rich patches to ramets in resource-poor patches, while the reverse was true under heterogeneous light environments, reflecting differences in the positions of leaves that act as the strongest sinks. Neither the mass increments nor the total mass of clonal fragments was significantly affected by heterogeneity of either light or nutrients, possibly because the experimental period was too short for differences to manifest themselves. CONCLUSIONS: This study clearly demonstrated that nitrogen is readily translocated between ramets, particularly under heterogeneous resource conditions. The translocation patterns were governed by functional 'division of labour' mechanisms that resulted in net nitrogen movement from understory sites to gaps, thereby enhancing the carbon acquisition of the whole fragment. Thus, physiological integration may provide benefits for S. palmata when it is growing under heterogeneous conditions in which there are deficits of certain environmental resources.     

Causes and consequences of sectoriality in the clonal herb Glechoma hederacea

The causes of sectoriality and consequences for clone behaviour are examined using data from the stoloniferous herb Glechoma hederacea. The proximal causes of physiological integration patterns are investigated using anatomical studies, acid fuchsin dye to reveal patterns of xylem continuity between ramets, and ¹⁴C as a label to reveal quantitative photoassimilate translocation patterns in the phloem. Dye movement in the xylem was acropetal and sectorial, and the sectoriality was determined by phyllotaxy. Patterns of ¹⁴C-labelled photoassimilate allocation were qualitatively similar to those of xylem based resources, although there was some basipetal movement of photoassimilate. The patterns of physiological integration and independence between ramets are shown to be governed by rules which depend on vascular continuity and discontinuity between ramets. Physiological support to stolon apices results in acquisition of relative branch autonomy (branches become semi-autonomous integrated physiological units, IPUs).This paper evaluates whether observed physiological integration patterns may be modified by altering normal source-sink relationships or by modifying environmental conditions. An experiment using different defoliation intensities, and different defoliation patterns at the same overall intensity, demonstrated that the precise positions of leaves removed from a clone had unique consequences for its subsequent development. Individual ramets of a given clone may be located in microhabitats of differing quality. An experiment in which competition was either present or absent throughout the space occupied by the clone, or patchy in distribution, showed that G. hederacea did not respond to competition at the whole clone level. Instead, connected stolons (IPUs) responded independently to local competition. Sectoriality may promote the restriction of lethal, localised environmental factors within the affected IPU. A study investigating the uptake and translocation of zinc by clones revealed that quantified patterns of zinc distribution resembled patterns of ¹⁴C movement in the phloem, and that there was no significant transport of zinc from one stolon to another.Although sectorial patterns of resource movement in G. hederacea can be modified in the short term, in the long-term, physiological integration may not allow this species to integrate the effects of environmental heterogeneity. A mobile clonal species with a high growth rate and relatively short-lived ramets, such as G. hederacea, is likely to benefit from a semi-autonomous response to patch quality at the level of the stolon, since the alternative of widespread intra-clonal support may increase the residence time of the clone in unfavourable pathches.     

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.     

Directional growth of a clonal bromeliad species in response to spatial habitat heterogeneity

Habitat selection by directional growth of plants has previously been investigated but field evidence for this phenomenon is extremely scarce. In this study we demonstrate directional clonal growth in Aechmea nudicaulis, a monocarpic, perennial bromeliad native to spatially heterogeneous sandy coastal plains (restinga) in Brazil. This habitat is characterized by a matrix of bare sand with interspersed vegetation islands. Due to very high soil surface temperatures and other stress factors such as drought, A. nudicauliscan only germinate inside vegetation islands. Nevertheless, this species is very common on bare sand. In this study we tested the hypothesis that clonal fragments occurring at the border and inside vegetation islands show habitat selection by growing preferentially towards the bare sand habitat (i.e. away from the center of vegetation islands).We randomly chose 116 clonal fragments in two distinct micro-environments (inside vegetation islands, and in the border area between bare sand and vegetation islands) in the natural habitat of A.nudicaulisand measured their growth direction in relation to the island center. We measured the growth directions of entire clonal fragments (defined as the line that connects the oldest and the youngest ramets of a clonal fragment) as well as the growth direction of the youngest internode on each fragment (the growth direction of the youngest ramet in relation to its parent ramet). We used Monte Carlo simulations to test for deviations from randomness in the growth direction of clonal fragments and individual internodes. The clonal fragments of A.nudicaulis showed a significant tendency to grow away from the center of vegetation islands. In other words, the main growth direction of clonal fragments growing inside vegetation islands or at the border between bare sand and vegetation islands was preferentially directed towards bare sand environments. Individual internodes at the border of vegetation islands also exhibited this tendency to grow towards the outside of vegetation islands, but internodes growing inside vegetation islands did not show directional growth. These results provide the first field evidence for habitat selection through directional growth of a clonal plant species.Co-ordinationg editor: J. Tuomi     

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.     

Clonal plants as cooperative systems: Benefits in heterogeneous environments

All natural environments are spatially and temporally heterogeneous. Consequently, their ability to provide essential resources for the growth of plants is variable. Modular plant species produce repeated basic structures which, in the case of clonal species, are called ramets. Ramets belonging to the same clone are distributed throughout the environment in space and time, and therefore they may be located in sites which differ in resource-providing quality. The connections between ramets may allow resources to be shared, enabling the clone to behave as a cooperative system. As a result of such physiological integration, ramets can survive in conditions where there is lethal shortage of a resource because they are connected to, and supported by, ramets located in conditions where there is ample supply of the same resource. Physiological integration between connected ramets presents opportunities for heterogeneous environments to be exploited to an extent that is only just becoming apparent. As heterogeneity is ubiquitous in natural environments, it may be expected that plants, as relatively immobile organisms, will have evolved the capacity to cope with it by making appropriate localized morphological and/or physiological plastic responses. Recent studies suggest that such responses not only enable clonal species to cope with environmental heterogeneity, but that under some circumstances they can benefit more from environments which are heterogeneous rather than homogeneous, even when both types of environment contain the same amount of resources. Studies on Glechoma hederacea (Lamiaceae) that illustrate this phenomenon are described.     

High levels of inter-ramet water translocation in two rhizomatous Carex species,as quantified by deuterium labelling

We studied water trnaslocation between interconnected mother and daughter ramets in two rhizomatous Carex species, using a newly developed quantitative method based on deuterium tracing. Under homogeneous conditions, in which both ramets were subjected either to wet or dry soil, little water was exchanged between the ramets. When the ramet pair was exposed to a heterogeneous water supply, water translocation became unidirectional and strongly increased to a level at which 30–60% of the water acquired by the wet ramet was exported towards the dry ramet. The quantity of water translocated was unrelated to the difference in water potential between the ramets, but highly correlated to the difference in leaf area. In both species, the transpiration of the entire plant was similar under heterogeneous and homogeneous wet conditions. This was a direct result of an increase in water uptake by the wet ramet in response to the dry conditions experienced by the interconnected ramet. In C. hirta, the costs and benefits of integration in terms of ramet biomass paralleled the responses of water consumption. This species achieved a similar whole-plant biomass in heterogeneous and homogeneous wet treatments, and water translocation was equally effective in the acropetal and basipetal directions. In C. flacca, responses of biomass and water consumption did not match and, under some conditions, water translocation imposed costs rather than benefits to the plants of this species. It is concluded that enhanced resource acquisition by donor ramets may be of critical importance for the net benefits of physiological integration in clonal plants.     

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.     

The effect of resource integration on fitness related traits in a clonal dune perennial,Hydrocotyle bonariensis

Summary The costs and benefits of resource integration in Hydrocotyle bonariensis were examined by comparing severed and intact clones grown across multiple resource gradients. Basipetal movement of water, nitrogen and photosynthates was demonstrated to occur between two rhizome branch systems interconnecting hundreds of ramets within a clone. Hydrocotyle clones of this size and larger have been shown to span highly patchy and unpredictable resource conditions in coastal dune environments. The extensive movement of water and nitrogen to portions of a clone deficient in these resources, resulted in a significant net benefit to the clone in terms of fitness-related traits: total biomass, ramet proliferation and seed production. The translocation of photosynthates across light gradients allowed for sexual reproduction and clonal expansion in the shade although this incurred a net cost to the clone in terms of ramet and seed production.     

Short‐term fitness benefits of physiological integration in the clonal herb Hydrocotyle peduncularis

Abstract We test whether physiological integration enhances the short‐term fitness of the clonal herb Hydrocotyle peduncularis (Apiaceae, R. Brown ex A. Richards) subjected to spatial variation in water availability. Our measures of fitness and costs and benefits are based on the relative growth rate of fragmented genets. Physiological integration over a gradient in soil moisture resulted in a highly significant net benefit to genet growth of 0.015 g g^?1 day^?1. This net benefit represents a significant enhancement of the average fitness of fragmented genets spanning the moisture gradient relative to the average of those growing in homogeneous moist or dry conditions. Sections of genet fragments growing in dry conditions in spatially heterogeneous treatments had significantly higher growth than the sections they were connected to that were growing in moist conditions. Within fragments, older (parent) sections growing in moist conditions experienced significant costs from connection to younger (offspring) sections growing in dry conditions. In contrast, offspring sections with ample water did not experience any costs when connected to parent sections growing in dry conditions. However, the net benefit of physiological integration was similar for parent and offspring sections, suggesting that parent and offspring sections contributed equally to the net benefit of physiological integration to genet growth and short‐term fitness.