白三叶无性系植物种群整合作用格局的研究

利用酸性品红和＾１４Ｃ－标记光合同化物,分析了在维管系统上的资源运输。在木质部中染料的运输是向顶的,在韧皮部中标记的光合同化物的运输方式与木质部中酸性品红的移动是相反的,具有强烈的基性,定量分析表明,尽管有一些向顶的运输,但向基运输是占绝对优势的,但随着无性系生长时间的不同,运输将会发生改变。一个无性系的变化取决于复杂的整合作用,使它们成为许多半自发的整合生理单位（ＩＰＵｓ）,不同匍匐茎的分枝,就     

Sectoriality and physiological organisation in herbaceous plants: an overview

The physiological organisation of plants is considered in relation to the carbon economy of plant parts. Although assimilate is partitioned according to the relative strength of sinks, in many species there is also a very close relationship between partitioning and shoot phyllotaxy, giving rise to sectorial patterns of allocation whereby only certain sinks are supported by any source leaf. Essentially these sinks are in the same orthostichy as the source leaf. This constraint of the vascular architecture on assimilate distribution to developing sinks such as leaves, flowers and fruits is not always absolute, as following the loss of their principal source leaves these sinks can in many cases be supplied with assimilate by other leaves via new inter-orthostichy pathways. The supply of assimilate to major sinks such as developing fruits becomes more and more localised with time so that a fruit in an axillary position becomes largely supported by its subtending leaf; the reproductive node—a metamer-can thus be regarded as a relatively autonomous unit of the plant (an IPU). Similary, once established after a developmental phase of assimilate import, tiller ramets and branches in unitary plants tend to become physiologically autonomous modules. However, the functional autonomy of tillers is reversed following defoliation or shading as they are then sustained by the import of assimilate, subject to its availability, from unaffected tillers. Consequently the plant becomes physiologically integrated by the flow of assimilate from one part to another. The mainly autonomous ramets of many stoloniferous and rhizomatous species display a similar pattern of physiological integration in response to source manipulation, but in some species the ramets appear to maintain their independent functioning as a normal feature of the carbon allocation within the clone. In other clonal species, as the clone develops and becomes more structurally complex, vascular constraints start to restrict the movement of resources, and the clone becomes composed of a number of semi-autonomous IPUs. In unitary plants branches appear to remain very physiologically isolated in terms of their carbon economy once they become established, irrespective of a range of source-sink manipulations.These different patterns of physiological integration and organisation are discussed in relation to different strategies of assimilate utilisation and conservation.     

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.     

Nutrient addition does not increase the benefits of clonal integration in an invasive plant spreading from open patches into plant communities

• One benefit of clonal integration is that resource translocation between connected ramets enhances the growth of the ramets grown under stressful conditions, but whether such resource translocation reduces the performance of the ramets grown under favourable conditions has not produced consistent results. In this study, we tested the hypothesis that resource translocation to recipient ramets may reduce the performance of donor ramets when resources are limiting but not when resources are abundant.
• We grew Mikania micrantha stolon fragments (each consisting of two ramets, either connected or not connected) under spatially heterogeneous competition conditions such that the developmentally younger, distal ramets were grown in competition with a plant community and the developmentally older, proximal ramets were grown without competition. For half of the stolon fragments, slow‐release fertiliser pellets were applied to both the distal and proximal ramets.
• Under both the low and increased soil nutrient conditions, the biomass, leaf number and stolon length of the distal ramets were higher, and those of the proximal ramets were lower when the stolon internode was intact than when it was severed. For the whole clone, the biomass, leaf number and stolon length did not differ between the two connection treatments. Connection did not change the biomass of the plant communities competing with distal ramets of M. micrantha.
• Although clonal integration may promote the invasion of M. micrantha into plant communities, resource translocation to recipient ramets of M. micrantha will induce a cost to the donor ramets, even when resources are relatively abundant.

     

The effects of physiological integration on biomass partitioning in plant modules: an experimental study with the stoloniferous herb Glechoma hederacea

Morphological and physiological plasticity are crucial attributes enabling plants to acquire resources from heterogeneous habitats. Although physiological integration can modify biomass partitioning in modules, especially when connected modules experience different conditions, its ecological importance has been largely overlooked. This experiment examined its effects on above- and belowground biomass partitioning by modules in the stoloniferous herb Glechoma hederacea. We studied how biomass allocation to roots by younger ramets was affected by connection to older ramets, and by nutrient conditions. A lower proportion of biomass was allocated to roots by younger ramets growing under low nutrient (LN) conditions when connected to older ramets in high nutrient (HN) conditions than when they were isolated, demonstrating localised modification of biomass partitioning due to physiological integration. The proportion of biomass allocated to roots by younger ramets was also lower when connected to older ramets in HN conditions than when connected to older ramets in LN conditions. Thus, the effect of integration on biomass partitioning depended on the nutrient conditions experienced by connected ramets. Such changes in biomass partitioning would result in more extensive stolon growth, and greater lateral displacement of new ramets. Understanding the ecological implications of phenotypic plasticity in plants will require further examination of the effects of physiological integration when connected modules experience contrasting growing conditions. This study demonstrates that such integration affects the biomass allocation strategy of connected ramets, enhancing resource acquisition in heterogeneous habitats. The widespread success of clonality in many communities is likely to be strongly promoted by this characteristic.     

The effects of developmental stage and source leaf position on integration and sectorial patterns of carbohydrate movement in an annual plant, Perilla frutescens (Lamiaceae)

A well-integrated plant shows extensive carbohydrate translocation through the plant body. Even in highly integrated plants, however, translocation patterns will be sectorial if vascular tissue restricts carbon movement to sectors along stems. Both integration and sectorial translocation patterns are sensitive to plant architecture and thus may change as a plant develops. These patterns should vary also with the position of the source leaf because leaves at each node are unique in age and vascular relationship to the rest of the plant. I measured the effects of developmental stage and location of the source leaf on integration and sectoriality in an annual plant, Perilla frutescens, by labeling plants with C at one of three leaves and four developmental stages. Stage and source leaf affected both integration and sectoriality. Most notably, integration declined and sectoriality increased during seed fill, when resource demand at each node was high. Furthermore, translocation was least extensive from the leaf supporting the largest number of seeds on its axillary branch. These results suggest that plants are not homogeneous collections of subunits; rather, the role of each leaf in a plant's carbon budget is a function of its age and location on the plant.     

Environmental heterogeneity and clonal growth: a study of the capacity for reciprocal translocation in Glechoma hederacea L.

Clonal fragments of Glechoma hederacea L. (Lamiaceae) were subjected to environments in which light and nutrients were supplied with a strictly negative association in space, i.e. when one of these resources was in ample supply the other was scarce. Treatments were chosen to simulate environments in which clones grew either within homogeneous conditions or across patch types (heterogeneous conditions). The hypothesis was tested that reciprocal translocation (i.e. exchange of both nutrients and assimilates) between connected groups of ramets would increase biomass production of clones growing under heterogeneous conditions compared to that of clones growing in homogeneous conditions. A cost-benefit analysis was carried out to test this hypothesis. Results suggested that reciprocal translocation did not occur at the structural scale considered in this experiment; no evidence was found for a significant effect on whole clone biomass of assimilate and/or nutrient translocation between clone parts experiencing contrasting levels of resource supply. It is suggested that predominantly acropetal movement of resources and the pattern of integrated physiological unit formation in G. hederacea are the main properties responsible for the lack of mutual physiological support between connected clonal fragments growing in differing habitat conditions. These properties are expected to promote clonal expansion and the exploitation of new territory, rather than sustaining clone parts in sub-optimal patches of habitat for prolonged periods of time.     

Physiological integration helps a clonal macrophyte spread into competitive environments and coexist with other species

Physiological integration may help clonal macrophytes invade or escape from existing communities. No studies have tested the above hypothesis in aquatic plants. In an outdoor pond experiment, we subjected clonal fragments of the submerged macrophyte Vallisneria spiralis L. to heterogeneous environments in which V. spiralis spread from bare habitats towards vegetated habitats occupied by Myriophyllum spicatum L. or V. spiralis spread from vegetated habitats towards bare habitats. V. spiralis stolons between ramets in bare habitats and in vegetated habitats were either intact or severed. We investigated the habitat selection of V. spiralis by examining the allocation of biomass and ramets to heterogeneous habitats during its vegetative spread phase. Results showed that the stolon connection had different effects on the habitat selection of V. spiralis with regard to invasion and escape. When V. spiralis spread from bare to vegetated habitats, in comparison to severing the stolon, the stolon connection eventually facilitated a 49% increase in biomass and a 27% increase in number of ramets allocated to vegetated habitats. However, when V. spiralis spread from vegetated to bare habitats, biomass and ramets allocated to bare habitats were not significantly changed by the stolon connection (only a 5% increase in biomass and a 6% increase in number of ramets). These results indicate that clonal integration facilitated V. spiralis not to escape from but invade into vegetated habitats. The study provides evidence that physiological integration is important for survival and tolerance of ramets in competitively stressful environments and can help clonal macrophytes coexist with other species.     

Hydraulic conductivity and PAT determine hierarchical resource partitioning and ramet development along Fragaria stolons

Co-ordination of metabolic and physiological activity between plant parts is key to the control of growth and development. Here the movement of resources and their allocation between mother plants and daughter ramets along Fragaria stolons was quantified with respect to hierarchy. Gradients of internodal ramet leaf water potential (ψ) and stolon and ramet hydraulic conductivities (L) were measured together with apparent stolon IAA movement via the polar auxin transport pathway (PAT). These processes are linked with measurements of stolon vascular development. The pattern of tissue differentiation and lignification in sequential stele sections of stolons demonstrated the rapid acquisition of the capacity for water transport, with transpiration potentially varying systematically with stolon lignification and the acropetal decline in stolon xylem ψ. Stolon and ramet L declined acropetally, with L across older ramets being significantly lower than that of the connecting stolons. The capacity for polar IAA transport increased with stolon age; this was due to increased transport intensity in older tissues. The partitioning of dry matter was strongly hierarchical with younger ramets smaller than older ramets, while foliar concentrations of N, P, and K were greater for the younger ramets. The results show that stolon anatomy develops rapidly at the apical end, facilitating hierarchical ramet development, which is evident as a basipetal increase in L. The rapid development of transport tissue functionality enables young unrooted ramets to acquire water, in order to supply an expanding leaf area, as well as mineral ions disproportionally with respect to older ramets. This facilitates colonization and self-rooting of apical ramets. The unidirectional increase in basipetal PAT along stolons facilitates hierarchical ramet development.     

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.     

Physiological integration enhanced the tolerance of Cynodon dactylon to flooding

Many flooding‐tolerant species are clonal plants; however, the effects of physiological integration on plant responses to flooding have received limited attention. We hypothesise that flooding can trigger changes in metabolism of carbohydrates and ROS (reactive oxygen species) in clonal plants, and that physiological integration can ameliorate the adverse effects of stress, subsequently restoring the growth of flooded ramets. In the present study, we conducted a factorial experiment combining flooding to apical ramets and stolon severing (preventing physiological integration) between apical and basal ramets of Cynodon dactylon, which is a stoloniferous perennial grass with considerable flooding tolerance. Flooding‐induced responses including decreased root biomass, accumulation of soluble sugar and starch, as well as increased activity of superoxide dismutase (SOD) and ascorbate peroxidase (APX) in apical ramets. Physiological integration relieved growth inhibition, carbohydrate accumulation and induction of antioxidant enzyme activity in stressed ramets, as expected, without any observable cost in unstressed ramets. We speculate that relief of flooding stress in clonal plants may rely on oxidising power and electron acceptors transferred between ramets through physiological integration.     

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.     

Physiological integration modifies δ15N in the clonal plant Fragaria vesca,suggesting preferential transport of nitrogen to water-stressed offspring

Background and Aims

One of the most striking attributes of clonal plants is their capacity for physiological integration, which enables movement of essential resources between connected ramets. This study investigated the capacity of physiological integration to buffer differences in resource availability experienced by ramets of the clonal wild strawberry plant, Fragaria vesca. Specifically, a study was made of the responses of connected and severed offspring ramets growing in environments with different water availability conditions (well watered or water stressed) and nitrogen forms (nitrate or ammonium).

Methods

The experimental design consisted of three factors, ‘integration’ (connected, severed) ‘water status’ (well watered, water stressed) and ‘nitrogen form’ (nitrate, ammonium), applied in a pot experiment. The effects of physiological integration were studied by analysing photochemical efficiency, leaf spectral reflectance, photosynthesis and carbon and nitrogen isotope discrimination, the last of which has been neglected in previous studies.

Key Results

Physiological integration buffered the stress caused by water deprivation. As a consequence, survival was improved in water-stressed offspring ramets that remained connected to their parent plants. The nitrogen isotope composition (δ¹⁵N) values in the connected water-stressed ramets were similar to those in ramets in the ammonium treatment; however, δ¹⁵N values in connected well-watered ramets were similar to those in the nitrate treatment. The results also demonstrated the benefit of integration for offspring ramets in terms of photochemical activity and photosynthesis.

Conclusions

This is the first study in which carbon and nitrogen isotopic discrimination has been used to detect physiological integration in clonal plants. The results for nitrogen isotope composition represent the first evidence of preferential transport of a specific form of nitrogen to compensate for stressful conditions experienced by a member clone. Water consumption was lower in plants supplied with ammonium than in plants supplied with nitrate, and therefore preferential transport of ammonium from parents to water-stressed offspring could potentially optimize the water use of the whole clone.     

Physiological integration among ramets of Lathyrus sylvestris L.

Summary Lathyrus sylvestris is a pioneer legume often found in disturbed habitats. Mainly reproduced through vegetative propagation, this clonal species presents a system of ramets that remain connected for several years. The existence of carbon transfer among ramets within a clone has been studied using ¹⁴C in situ. Assimilate translocation from primary to secondary ramets was observed in all clones when the primary ramet was exposed to ¹⁴CO₂. The amount of transfer ranged from trace up to 90% of the total ¹⁴C incorporated. However, in only half of the clones there was consistent enrichment of the secondary ramet (5 to 89%) suggesting that interramets transfer of carbon may be facultative. Furthermore, when significant export occurred from the primary ramet, it was always principally towards only one ramet even when the clone included more than one. The transfer of ¹⁴C from secondary to primary ramets was shown to be significant only when photosynthesis of the latter was decreased by shading. In this case import of carbon was never more than 60% of the incorporated ¹⁴C.No correlation was found between age or size of the ramets and the intensity of transfer. The shading effect let suppose that transfers are mainly driven by carbon limitation due to changing environmental conditions and not to the state of ramet maturity. The adaptative advantage of such facultative physiological integration between ramets of a clone is discussed.     

Relative costs and benefits of clonal integration of Zoysia japonica under various N:P ratios

Clonal plants adaptively respond to abiotic stress, but to date little is known about under what circumstanses clonal integration is beneficial, or costly. To study the costs and benefits of clonal integration on clonal growth, we cultivated Zoysia japonica under three ratios of N:P (N:P ≈ 7, 14:1 and 21:1), and four types of stolon severing treatments (connected stolon CS, light severing LS, moderate severing MS, serious severing SS). The results showed that Z. japonica performed best at a low ratio of N:P (N:P ≈ 7:1). When the stolons were connected (CS), the growth of primary A‐ramets, multiple‐nodes and stolons benefited from clonal integration; however, the growth of primary B‐ramets on the primary stolons, A and B ramets was significantly reduced (p < 0.05). In the moderate stolon severing treatment MS_7:1 (N:P ≈ 7:1), clonal integration appeared more cost effective than in all other treatments. On the whole, with increasing stolon severing intensity, the cost of clonal integration increased and the clonal growth of Z. japonica declined significantly. The pattern of biomass allocation may be useful for Z. japonica to adapt to the various environments, and clonal integration plays a significant role under adverse environmental conditions.     

Physiological integration in the clonal perennial herb Trifolium repens L.

Summary Translocation of ¹⁴C-labelled carbohydrates between the parent stolon and branches, and among branches, of Trifolium repens plants was investigated in two glasshouse experiments to determine patterns of physiological organisation in this clonal species. Differential defoliation treatments were applied to the parent stolon and/or branches to test the sensitivity of translocation to the short-term carbon needs of defoliated sinks. Strong reciprocal exchange of carbohydrate between the parent stolon and branches was observed, with 18 41% of the ¹⁴C exported from leaves on the parent stolon moving to branches, while branches simulta-neously exported 25% (for old source branches) to 54% (for young source branches) of the ¹⁴C they assimilated to the parent plant, including translocation to other branches. Branch-to-branch translocation occurred both acropetally and basipetally. Parent-to-branch, branch-to-parent and branch-to-branch carbon fluxes all increased in response to defoliation of the sink, at the expense of carbon supply to stolon tissue or roots of the source module. Reduced export to stolon tissue of the parent axis played a major role in facilitating C reallocation from leaves on the parent stolon to defoliated branches. The observed patterns of C allocation and translocation could be adequately explained by accepted source-sink theory, and are consistent with a high degree of intra-plant physiological integration in resource supply and utilisation. This information provides mechanistic explanations for aspects of the growth dynamics and ecological interactions of T. repens in the patchy environment of a grazed pasture.     

Physiological integration increases the survival and growth of the clonal invader <Emphasis Type="Italic">Carpobrotus edulis</Emphasis>

Clonal growth seems to be a common trait for many of the most aggressive invasive plant species. However, little research has been conducted to determine the role of clonality in the successful invasion of new areas by exotic species. Carpobrotus edulis (L.) N.E. Br. is a mat-forming succulent plant, native to South Africa that is invasive in coastal dunes of Australia, New Zealand, USA and Southern Europe. Although Carpobrotus edulis is a clonal plant, there is no information on the role of clonality for the invasion by this species, therefore the objective of this study was to test whether or not physiological integration improves the performance of C. edulis invading coastal sand dunes. To do that, a 6-month field experiment was designed in which the stolon connections between the apical ramets and the C. edulis mats were severed to prevent physiological integration. This treatment was applied to ramets growing under high and low competition with the native species. Apical ramets with intact stolon connections were used as control. Integration improved the survivorship and growth of apical ramets, both in high and low competition. Connected ramets showed a more pronounced increase of clonal growth (estimated as stolon length) during the experimental period and a higher total biomass and number of ramets at the completion of the experiment. In terms of survivorship, the benefit of integration was greater under high competition. Physiological integration can therefore be considered an important factor in the invasiveness of C. edulis, both in open space and in direct competition with the native plants.     

Clonal integration in Fragaria vesca growing in metal-polluted soils: parents face penalties for establishing their offspring in unsuitable environments

Clonal plants often establish descendent ramets in sites with contrasting presence of favourable and unfavourable factors. Connections between ramets allow translocation of essential resources from established ramets to developing ramets and, as consequence, integration confers net benefits to ramets growing under unfavourable conditions. Therefore, integrated ramets may survive in habitat patches that would be lethal to independent ramets or non-clonal plants. This experiment aimed to investigate the physiological and morphological responses of the clonal plant Fragaria vesca growing in heterogeneous substrate with patches of contrasting quality (i.e. uncontaminated or heavy-metal-contaminated). We observed that parents reduced their photosynthetic efficiencies and growth as consequence of maintaining their offspring. This cost did not affect survival of the parents. Physiological integration brings about benefits to offspring ramets growing both at uncontaminated and heavy-metal-contaminated soils. The benefits of integration were detected in both physiological and morphological traits, enhancing the survivorship of offspring ramets in the Cu-polluted soils. We conclude that integration improves the performance of developing ramets of F. vesca growing in heavy-metal-contaminated habitats, allowing clone systems to overcome the establishment risks and maintain their presence in these less favourable sites.     

Clonal integration in Ranunculus reptans: by‐product or adaptation?

We studied fitness consequences of clonal integration in 27 genotypes of the stoloniferous herb Ranunculus reptans in a spatially heterogeneous light environment. We grew 216 pairs of connected ramets (eight per genotype) with mother ramets in light and daughter ramets in shade. In half of the pairs we severed the stolon connection between the two ramets at the beginning of the experiment. During the experiment, 52.7% of the ramet pairs with originally intact connection physically disintegrated. We detected significant variation among genotypes in this regard. Survival of planted ramets was 13.3% higher for originally connected pairs. Moreover, there was significant variation among genotypes in survival, in the difference in survival between plant parts developing from mother and daughter ramets, and in the effect of integration on this difference. In surviving plants connection between ramets decreased size differences between mother and daughter parts. Variation among genotypes was significant in growth and reproduction and marginally significant in the effect of physiological integration on growth and reproduction. Connected daughter ramets had longer leaves and internodes than daughters in severed pairs indicating that integration stimulated plant foraging in both the vertical and the horizontal plane. Observed effects of integration on fitness components in combination with genetic variation in maintenance and effects of connection indicate that clonal integration in R. reptans has the capability to evolve, and therefore suggest that clonal integration is adaptive. If genetic variation in integration is common, future studies on clonal integration should always use defined genetic material and many clones to allow extrapolation of results to population and wider levels.     

UV-B辐射方向对白三叶克隆整合的影响

增强UV-B辐射会对植物生长和生理生化过程产生有害效应。克隆植物中,相连的克隆分株对经常共享资源和激素,然而鲜有关于异质性UV-B辐射下UV-B辐射方向对克隆整合的影响及克隆植物形态结构变化的报道。模拟同质(克隆分株片段均处于自然背景辐射)和异质(克隆分株一端处于自然背景辐射,另一端处于补加的UV-B辐射)UV-B辐射,以克隆植物白三叶为材料,进行连接和隔断处理,研究UV-B辐射方向对克隆整合强度变化、叶片形态结构特化及生理可塑性的影响。结果表明:异质性UV-B辐射下,15N同位素标记端保留的15N百分比高于同质UV-B辐射处理,转移到无标记相连端的15N含量则降低,紫外辐射处理和同位素标记是否处于同一分株端对结果无显著性影响,说明克隆植物白三叶生理整合存在但整合强度降低,辐射方向与克隆整合强度无关;隔断处理组气孔长度增加,栅栏组织增厚,但连接处理组却无此变化,表明生理整合在白三叶叶片形态结构特化中发挥作用。UV-B辐射下,最小荧光、电子传递速率及光化学淬灭系数降低但非光化学淬灭系数升高,而生理整合却使结果相反;叶绿素和紫外吸收物可在异质性UV-B辐射相连的两端运输分享。以上均表明异质UV-B辐射环境中,UV-B辐射胁迫端克隆分株通过生理整合从非胁迫端获益,并以此提高胁迫环境中克隆植物对资源的利用效率。