Clonal integration supports the expansion from terrestrial to aquatic environments of the amphibious stoloniferous herb Alternanthera philoxeroides

Effects of clonal integration on land plants have been extensively studied, but little is known about the role in amphibious plants that expand from terrestrial to aquatic conditions. We simulated expansion from terrestrial to aquatic habitats in the amphibious stoloniferous alien invasive alligator weed ( Alternanthera philoxeroides ) by growing basal ramets of clonal fragments in soils connected (allowing integration) or disconnected (preventing integration) to the apical ramets of the same fragments submerged in water to a depth of 0, 5, 10 or 15 cm. Clonal integration significantly increased growth and clonal reproduction of the apical ramets, but decreased both of these characteristics in basal ramets. Consequently, integration did not affect the performance of whole clonal fragments. We propose that alligator weed possesses a double-edged mechanism during population expansion: apical ramets in aquatic habitats can increase growth through connected basal parts in terrestrial habitats; however, once stolon connections with apical ramets are lost by external disturbance, the basal ramets in terrestrial habitats increase stolon and ramet production for rapid spreading. This may contribute greatly to the invasiveness of alligator weed and also make it very adaptable to habitats with heavy disturbance and/or highly heterogeneous resource supply.     

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

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

Clonal integration increases performance of ramets of the fern Diplopterygium glaucum in an evergreen forest in southeastern China

Physiological integration is a major ecological advantage of clonal growth in angiosperms. Clonal growth is also common in pteridophytes, but almost no study has tested whether clonal integration increases performance in ramets of pteridophytes in natural populations. To test this hypothesis and also whether the positive effect of integration is greater on smaller ramets, we severed the connecting rhizomes of individual ramets of the common, understory fern Diplopterygium glaucum in an evergreen, broadleaf forest in southeastern China. In another experiment, we severed rhizomes around the edges of small plots each containing several ramets. After 19.5 weeks, survival was 100% in intact individual ramets but only 27% in severed ones. Among surviving ramets, final dry mass and lamina mass were also less in severed than in intact ramets, though stalk, rhizome, and root mass and maximum quantum yield of PSII (F_v/F_m) were not reduced. Individual ramets with fewer stalk nodes had lower dry mass but were not more affected by severing than ramets with more stalk nodes. Severance around the edge of plots did not significantly affect the combined final mass of the ramets within a plot. We conclude that clonal integration can have significant positive effects on both survival and growth of individual ramets of ferns in natural populations.     

Clonal integration facilitates spread of Paspalum paspaloides from terrestrial to cadmium‐contaminated aquatic habitats

• Cadmium (Cd) is a hazardous environmental pollutant with high toxicity to plants, which has been detected in many wetlands. Clonal integration (resource translocation) between connected ramets of clonal plants can increase their tolerance to stress. We hypothesised that clonal integration facilitates spread of amphibious clonal plants from terrestrial to Cd‐contaminated aquatic habitats.
• The spread of an amphibious grass Paspalum paspaloides was simulated by growing basal older ramets in uncontaminated soil connected (allowing integration) or not connected (preventing integration) to apical younger ramets of the same fragments in Cd‐contaminated water.
• Cd contamination of apical ramets of P. paspaloides markedly decreased growth and photosynthetic capacity of the apical ramets without connection to the basal ramets, but did not decrease these properties with connection. Cd contamination did not affect growth of the basal ramets without connection to the apical ramets, but Cd contamination of 4 and 12 mg·l^?1 significantly increased growth with connection. Consequently, clonal integration increased growth of the apical ramets, basal ramets and whole clones when the apical ramets were grown in Cd‐contaminated water of 4 and 12 mg·l^?1. Cd was detected in the basal ramets with connection to the apical ramets, suggesting Cd could be translocated due to clonal integration. Clonal integration, most likely through translocation of photosynthates, can support P. paspaloides to spread from terrestrial to Cd‐contaminated aquatic habitats.
• Amphibious clonal plants with a high ability for clonal integration are particularly useful for re‐vegetation of degraded aquatic habitats caused by Cd contamination.

     

Partial mechanical stimulation facilitates the growth of the rhizomatous plant Leymus secalinus: modulation by clonal integration

Background and Aims

Mechanical stimulation (MS) often induces plants to undergo thigmomorphogenesis and to synthesize an array of signalling substances. In clonal plants, connected ramets often share resources and hormones. However, little is known about whether and how clonal integration influences the ability of clonal plants to withstand MS. We hypothesized that the effects of MS may be modulated by clonal integration.

Methods

We conducted an experiment in which ramet pairs of Leymus secalinus were subjected to three treatments: (1) connected ramet pairs under a homogeneous condition [i.e. the proximal (relatively old) and distal (relatively young) ramets were not mechanically stressed]; (2) connected ramet pairs under a heterogeneous condition (i.e. the proximal ramet was mechanically stressed but the distal ramet was not); and (3) disconnected ramet pairs under the same condition as in treatment 2. At the end of the experiment, we harvested all plants and determined their biomass and allocation.

Key Results

Clonal integration had no significant influence on measured traits of distal L. secalinus ramets without MS. However, under MS, plants with distal ramets that were connected to a mother ramet produced more total plant biomass, below-ground biomass, ramets and total rhizome length than those that were not connected. Partial MS exerted local effects on stimulated ramets and remote effects on connected unstimulated ramets. Partial MS increased total biomass, root/shoot ratio, number of ramets and total rhizome length of stimulated proximal ramets, and increased total biomass, root weight ratio, number of ramets and total rhizome length of connected unstimulated ramets due to clonal integration.

Conclusions

These findings suggest that thigmomorphogenesis may protect plants from the stresses caused by high winds or trampling and that thigmomorphogenesis can be strongly modulated by the degree of clonal integration.     

Trampling, defoliation and physiological integration affect growth, morphological and mechanical properties of a root-suckering clonal tree

Background and Aims

Grazing is a complex process involving the simultaneous occurrence of both trampling and defoliation. Clonal plants are a common feature of heavily grazed ecosystems where large herbivores inflict the simultaneous pressures of trampling and defoliation on the vegetation. We test the hypothesis that physiological integration (resource sharing between interconnected ramets) may help plants to deal with the interactive effects of trampling and defoliation.

Methods

In a field study, small and large ramets of the root-suckering clonal tree Populus simonii were subjected to two levels of trampling and defoliation, while connected or disconnected to other ramets. Plant responses were quantified via survival, growth, morphological and stem mechanical traits.

Key Results

Disconnection and trampling increased mortality, especially in small ramets. Trampling increased stem length, basal diameter, fibrous root mass, stem stiffness and resistance to deflection in connected ramets, but decreased them in disconnected ones. Trampling decreased vertical height more in disconnected than in connected ramets, and reduced stem mass in disconnected ramets but not in connected ramets. Defoliation reduced basal diameter, leaf mass, stem mass and leaf area ratio, but did not interact with trampling or disconnection.

Conclusions

Although clonal integration did not influence defoliation response, it did alleviate the effects of trampling. We suggest that by facilitating resource transport between ramets, clonal integration compensates for trampling-induced damage to fine roots.     

Effects of clonal integration on photosynthesis of the invasive clonal plant <Emphasis Type="Italic">Alternanthera philoxeroides</Emphasis>

A greenhouse experiment examined whether clonal integration improves photosynthesis of ramets of alligator weed [Alternanthera philoxeroides (Mart.) Griseb.], a widespread invasive clonal plant in China, in heterogeneous (He) nutrient habitats. The connected pairs of ramets experienced different nutrient levels [high homogeneous (Ho) nutrient, low Ho nutrient, and two He nutrient treatments]. Clonal integration significantly improved the net photosynthetic rate, stomatal conductance, transpiration rate, and minimal and maximal chlorophyll fluorescence of ramets of alligator weed in low nutrient condition. These characteristics may contribute to the success of the ramets of alligator weed in invading contrasting habitats. The clonal integration of the invasive clonal plants may contribute significantly to their invasiveness.     

克隆整合对异质性光照环境下紫竹根际土壤氮素有效性的影响

以根状茎克隆植物紫竹为对象,研究克隆整合对遭受异质性光照胁迫分株根际土壤有机碳(SOC)、总氮(TN)、溶解性有机碳(DOC)、溶解性有机氮(DON)、氨氮(NH_4~+-N)、硝态氮(NO_3~--N)以及微生物群落组成的影响。所取紫竹克隆片段由一个母本分株和一个子代分株组成,母本分株置于全光照下,而子代分株置于80%遮阴环境中,同时母本分株与子代分株间的根茎保持连接或割断处理。结果表明:与切断处理相比,紫竹遮荫子代分株根际土壤的SOC、TN、DOC、NH_4~+-N在保持根状茎连接时显著更高,这表明异质性光照环境下克隆整合可能改善紫竹连接遮荫子代分株根际土壤的氮素有效性。克隆整合提高了连接遮阴状态下紫竹子代分株根际土壤中的放线菌、真菌和革阴细菌的PLFAs浓度。通过对遮阴子代分株根际土壤微生物群落PLFAs主成分分析得出克隆整合导致遮阴子代分株根际土壤微生物群落结构发生显著变化。该研究结果暗示了紫竹可能通过克隆整合作用降低土壤中某些对氮利用有效性影响较低的细菌数量,而增加对土壤氮利用起重要作用的放线菌和真菌的数量,进而改善紫竹对土壤中氮利用的有效性,这有利于增强克隆植物对时空异质性生境的适应能力。     

Trade-offs among growth, clonal, and sexual reproduction in an invasive plant Spartina alterniflora responding to inundation and clonal integration

The purpose of this article was to study the trade-offs among vegetative growth, clonal, and sexual reproduction in an aquatic invasive weed Spartina alterniflora that experienced different inundation depths and clonal integration. Here, the rhizome connections between mother and daughter ramets were either severed or left intact. Subsequently, these clones were flooded with water levels of 0, 9, and 18 cm above the soil surface. Severing rhizomes decreased growth and clonal reproduction of daughter ramets, and increased those of mother ramets grown in shallow and deep water. The daughter ramets disconnected from mother ramets did not flower, while sexual reproduction of mother ramets was not affected by severing. Clonal integration only benefited the total rhizome length, rhizome biomass, and number of rhizomes of the whole clones in non-inundation conditions. Furthermore, growth and clonal reproduction of mother, daughter ramets, and the whole clone decreased with inundation depth, whereas sexual reproduction of mother ramets and the whole clones increased. We concluded that the trade-offs among growth, clonal, and sexual reproduction of S. alterniflora would be affected by inundation depth, but not by clonal integration.     

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.     

克隆整合对无芒雀麦在异质性盐分环境中存活和生长的影响

无芒雀麦是浑善达克沙地植物群落中占优势的多年生根茎禾草.研究了克隆整合特性对无芒雀麦在异质性盐分环境中存活和生长的影响.结果表明,克隆整合显著提高了无芒雀麦分株在高盐环境中的存活能力,耗-益分析表明无芒雀麦在高盐斑块中分株的生物量、分株数、根茎节数和根茎总长显著受益于克隆整合,而与之相连的非盐分斑块中的分株却没有产生显著的损耗.因而,克隆整合特性是无芒雀麦对异质性环境形成的重要适应对策,它使无芒雀麦能够扩展到不适合植物生长的高盐分斑块中,从而增加了无芒雀麦在浑善达克沙地中的存活和生长,提高了其在半干旱沙化地区的适合度.     

附生蕨类植物的克隆性研究进展

石松类及蕨类植物在高等植物中处于比较特殊的进化与系统发育地位, 同时具有孢子植物(孢子)与种子植物(维管束)的双重特征。附生蕨类植物是蕨类植物中占据独特生境的一个大类群, 其生活史策略及进化历史与其附生生长的森林生态系统紧密相关。大部分附生蕨类植物的克隆生长习性及克隆生活史性状在其生态适应中具有重要作用, 但这方面未引起广泛关注。本文主要综述了中国山地森林中附生蕨类植物的根状茎克隆生长、克隆性与生态适应性、不同克隆生长方式与进化等方面, 并展望了蕨类植物克隆性在森林生态系统过程与功能中的作用, 以及今后如何将蕨类植物生态学研究与气候变化、植被恢复、土地利用变化等全球变化的主流方向进行结合。     

Effects of nitrogen addition on clonal integration between mother and daughter ramets of Moso bamboo: a 13C-CO2 pulse labeling study

用¹³C-CO₂脉冲标记法研究氮添加对毛竹母子分株克隆整合的影响 相连分株间的资源共享(即克隆整合)是克隆植物的显著特征。克隆整合使毛竹(Phyllostachys pubescens)对多种环境条件具有较强的适应性。但毛竹通过克隆整合获得性能提升的机制尚不明确。本 研究区分并分析了毛竹光合碳的顶向整合和基向整合,以探究毛竹克隆片段如何在土壤氮异质性条件下 提高整体收益。本研究以由两株不同年龄毛竹分株组成 的克隆片段为研究对象,分株间通过根状茎连接。 每个分株设置氮添加或无氮添加两种处理,并通过单株母株或子株的¹³C-CO₂脉冲标记对光合碳的顶向整合和基向整合进行区分。研究结果表明,子株氮添加显著促进了光合碳由母株向子株的顶向整合,无论母株是否有氮添加,均向氮添加子株转运更多的光合碳。转运自无氮添加母株的光合碳主要分配至氮添加子株的 叶片,而转运自氮添加母株的光合碳则主要分配至根系。与无氮添加处理的子株相比,氮 添 加子株转运更多的光合碳至母株,且该过程不受母株氮添加的影响。转运自氮添加子株的光合碳主要 分配至无氮添加母株的根系和氮添加母株的叶片。这些结果说明,母株优先将更多的资源投入到具有高 养分可利用性的子株,随后子株作为更高效的资源采集点,根据母株的养分状况针对富集资源进行获取。通过这种方式,克隆植物可以降低资源获取成本并提高资源获取效率,最大限度地提升其整体表现。     

A tropical epiphytic orchid uses a low‐light interception strategy in a spatially heterogeneous light environment

Light is considered a non‐limiting factor for vascular epiphytes. Nevertheless, an epiphyte's access to light may be limited by phorophyte shading and the spatio‐temporal environmental patchiness characteristic of epiphytic habitats. We assessed the extent to which potential light interception in Rodriguezia granadensis, an epiphytic orchid, is determined by individual factors (plant size traits and leaf traits), or environmental heterogeneity (light patchiness) within the crown of the phorophyte, or both. We studied 104 adult plants growing on Psidium guajava trees in two habitats with contrasting canopy cover: a dry tropical forest edge, and isolated trees in a pasture. We recorded the number of leaves and the leaf area, the leaf position angles, and the potential exposure of the leaf surface to direct irradiance (silhouette area of the leaf blade), and the potential irradiance incident on each plant. We found the epiphytes experience a highly heterogeneous light environment in the crowns of P. guajava. Nonetheless, R. granadensis plants displayed a common light interception strategy typical of low‐light environments, resembling terrestrial, forest understory plants. Potential exposure of the total leaf surface to direct irradiance correlated positively with plant size and within‐plant variation in leaf orientation. In many‐leaved individuals, within‐plant variation in leaf angles produced complementary leaf positions that enhanced potential light interception. This light interception strategy suggests that, in contrast to current wisdom, enhancing light capture is important for vascular epiphytes in canopies with high spatio‐temporal heterogeneity in light environments.     

极端干旱生境下胡杨克隆水分整合特征及其生态意义

克隆水分整合是克隆植物有效利用异质性生境资源的重要对策,也是在恶劣生境下得以定植、存活的重要生存策略。本研究以塔里木河下游的胡杨（Populus euphratica Oliv.）为对象,结合野外调查、稳定同位素技术、生理生态监测实验,调查胡杨的克隆水分整合特征并分析其生态意义。结果显示,胡杨母株与其克隆幼株间存在显著水分生理整合,水分整合以顶向传输为主,水分整合过程与整合水量受到母株生理节律及母株到幼株间隔子长度的影响;克隆幼株因水分整合而能够获取与母株相似的深层土壤水分,比同一区域实生幼株拥有更好的水分获取能力,并使胡杨克隆幼株比实生幼株保持相对更高的水力导度、叶片水分含量及叶水势;在极端干旱生境下的这种水分获取及利用策略使胡杨克隆幼株比实生幼株具有更高的光化学效率与光合性能,有助于克隆幼株在不利生境下定植、存活并保持更高的生存优势。     

Water integration patterns in two rhizomatous dune perennials of different clonal fragment size

In clonal plants, a clonal fragment is a basic unit. There may exist clonal integration that is often regarded adaptive. By definition, a larger clonal fragment comprises more interconnected ramets and/or occupies a larger area of the habitat. When the habitat is so heterogeneous in terms of essential resources that the resources the whole clonal fragment needs can be captured only in a limited number of microhabitats, a large clonal fragment may require extensive and/or intensive clonal integration. Therefore, we hypothesize that in an environment where the resources are distributed in a highly heterogeneous way, a species forming large clonal fragments possesses more extensive and/or intensive clonal integration gaining essential resources than that forming small ones.This hypothesis was tested in a field experiment with two rhizomatous species growing in inland-dune habitats with high heterogeneity of water resource, one (Psammochloa villosa) forming large clonal fragments and the other (Hedysarum laeve) forming small ones. In P. villosa, tracer-(water-soluble acid fuchsin)-labeled water could be transported along the rhizome at a velocity of about 0.93 m per hour and 4 ramets per hour, which was much faster than that in H. laeve. Similarly, the acid fuchsin-labeled water was transported to a longer length in the rhizomes (3.96 m vs. 1.12 m) and to more ramets (14 vs. 3) in P. villosa than in H. laeve. In P. villosa, the acid fuchsin-labeled water reached the ends of the rhizomes and all ramets along the rhizomes were dyed purple. In H. laeve, however, the acid fuchsin-labeled water could not reach the rhizome tips and along the rhizomes only some ramets were labeled. The results suggest that P. villosa is an extensive and intensive integrator, whereas H. laeve is a restrictive and less intensive integrator, supporting the hypothesis. The possible mechanisms and the ecological implications of the findings were discussed.     

Effects of stolon severing on the expansion of Alternanthera philoxeroides from terrestrial to contaminated aquatic habitats

Few studies have examined the effects of clonal integration (translocation of resources between interconnected ramets) during the expansion of amphibious clonal plants from terrestrial to aquatic habitats. We conducted a greenhouse experiment to simulate the expansion of plants from terrestrial to contaminated aquatic habitats in the amphibious stoloniferous herb Alternanthera philoxeroides (alligator weed). The proximal ramets (i.e. relatively old) of clonal fragments grown in uncontaminated soils were connected to (allowing clonal integration) or disconnected from (preventing clonal integration) distal ramets (i.e. relatively young) grown either in uncontaminated water (control, no CuSO₄) or in four copper‐contaminated water treatments containing 31.25, 62.5, 125 and 250 mg/L CuSO₄, respectively. When a stolon connection was severed, all distal ramets grown in the contaminated water died. When the stolon connection was intact, however, the survival rate of the distal ramets was 85–100% when they were grown at the three lower levels of contamination and 43.75% at the highest level. Moreover, the survival rate and growth of the distal ramets grown in the three lower levels of contamination treatments did not differ from those in the control (uncontaminated water). These results suggest that clonal integration could greatly improve the survival and growth of alligator weed subjected to moderate levels of copper stress. Although clonal integration could also increase the survival rate of the connected distal ramets subjected to the highest level of copper stress (250 mg/L CuSO₄) compared with that of disconnected distal ramets, the survival rate and growth measures were still significantly lower than those in the control. This suggests that clonal integration plays a limited role in the survival and growth of alligator weed when it is subjected to severe stress by high levels of copper contamination.     

Diversity of dry forest epiphytes along a gradient of human disturbance in the tropical Andes

Question: Disturbance effects on dry forest epiphytes are poorly known. How are epiphytic assemblages affected by different degrees of human disturbance, and what are the driving forces? Location: An inter‐Andean dry forest landscape at 2300 m elevation in northern Ecuador. Methods: We sampled epiphytic bryophytes and vascular plants on 100 trees of Acacia macracantha in five habitats: closed‐canopy mixed and pure acacia forest (old secondary), forest edge, young semi‐closed secondary woodland, and isolated trees in grassland. Results: Total species richness in forest edge habitats and on isolated trees was significantly lower than in closed forest types. Species density of vascular epiphytes (species per tree) did not differ significantly between habitat types. Species density of bryophytes, in contrast, was significantly lower in edge habitat and on isolated trees than in closed forest. Forest edge showed greater impoverishment than semi‐closed woodland and similar floristic affinity to isolated trees and to closed forest types. Assemblages were significantly nested; habitat types with major disturbance held only subsets of the closed forest assemblages, indicating a gradual reduction in niche availability. Distance to forest had no effect on species density of epiphytes on isolated trees, but species density was closely correlated with crown closure, a measure of canopy integrity. Main conclusions: Microclimatic changes but not dispersal constraints were key determinants of epiphyte assemblages following disturbance. Epiphytic cryptogams are sensitive indicators of microclimate and human disturbance in montane dry forests. The substantial impoverishment of edge habitat underlines the need for fragmentation studies on epiphytes elsewhere in the Tropics.     

Clonal Integration Enhances the Performance of a Clonal Plant Species under Soil Alkalinity Stress

Clonal plants have been shown to successfully survive in stressful environments, including salinity stress, drought and depleted nutrients through clonal integration between original and subsequent ramets. However, relatively little is known about whether clonal integration can enhance the performance of clonal plants under alkalinity stress. We investigated the effect of clonal integration on the performance of a typical rhizomatous clonal plant, Leymus chinensis, using a factorial experimental design with four levels of alkalinity and two levels of rhizome connection treatments, connected (allowing integration) and severed (preventing integration). Clonal integration was estimated by comparing physiological and biomass features between the rhizome-connected and rhizome-severed treatments. We found that rhizome-connected treatment increased the biomass, height and leaf water potential of subsequent ramets at highly alkalinity treatments but did not affect them at low alkalinity treatments. However, rhizome-connected treatment decreased the root biomass of subsequent ramets and did not influence the photosynthetic rates of subsequent ramets. The biomass of original ramets was reduced by rhizome-connected treatment at the highest alkalinity level. These results suggest that clonal integration can increase the performance of clonal plants under alkalinity stress. Rhizome-connected plants showed dramatically increased survival of buds with negative effects on root weight, indicating that clonal integration influenced the resource allocation pattern of clonal plants. A cost-benefit analysis based on biomass measures showed that original and subsequent ramets significantly benefited from clonal integration in highly alkalinity stress, indicating that clonal integration is an important adaptive strategy by which clonal plants could survive in local alkalinity soil.     

紫耳箭竹克隆形态可塑性对典型冠层结构及光环境的响应

在重庆金佛山国家自然保护内,选择了3种典型群落类型(落叶阔叶林、常绿落叶阔叶混交林和常绿阔叶林),使用Hemiview数字植物冠层分析系统量化群落冠层结构和光环境特征,并对林下紫耳箭竹(Fargesia decurvata)的形态可塑性特征进行调查,分析冠层结构和光环境特征改变下紫耳箭竹形态可塑性的差异,并探讨它们之间的相互关系。结果表明:(1)随着落叶阔叶林"常绿落叶阔叶混交林"常绿阔叶林演替的进行,群落的冠层开度降低,叶面积指数增加,平均叶倾角变小,趋于水平化,冠层对光的截获能力提高,林下光照的强度降低(P0.05)。(2)随着光照强度的降低,紫耳箭竹分株矮小化,叶片变窄,生物量积累降低,但通过增大比茎长、叶面积率和比叶面积提高对光的利用效率,并增大分枝角度和比隔长有效适应弱光环境。(3)在光照条件差的常绿阔叶林下,紫耳箭竹降低对地下茎的投资,将较多的生物量用于秆的增高增长和叶片的生长;而在光照条件好的落叶阔叶林环境下,紫耳箭竹降低对枝、叶生物量的分配,则加大对地下茎的投资,可认为是克隆植物对水分资源所表现的一种觅食行为。研究表明,紫耳箭竹种群随着冠层结构的改变发生了明显的可塑性变化,这些可塑性变化是种群对冠层结构和光环境差异的适应性反应的结果,有利于增强种群对异质生境中光资源的获取和利用;群落内部可以通过调控冠层结构的改变协调和控制小径竹种群的发展。