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.

     

养分异质条件下结缕草克隆分株生长及光合作用的生理整合

通过对连接和断开的分株进行不同养分处理,研究养分异质条件下结缕草克隆分株生长及光合作用的变化.结果表明:在养分异质条件下,处于中、高养分水平的母株可以提高与其相连子株的叶长、叶宽、根质量、叶质量、光合速率、气孔导度、蒸腾速率和水分利用效率,高养分条件下分别提高16.0%、8.3%、24.4%、58.1%、30.3%、54.0%、9.2%和21.9%,降低根冠比和胞间CO2浓度,在高养分下分别降低21.6%和31.5%;处于中、高养分水平的子株对与其相连母株的生长及光合特性没有显著影响.在养分异质条件下,结缕草母株对子株存在生理整合,养分梯度越大,整合强度越强.结缕草克隆子株可以从母株获益,但母株不能从子株获益,子株是生理整合单向获益者.     

Differences in physiological integration between invasive and noninvasive introduced clonal species of Carpobrotus

AimsClonal growth is associated with invasiveness in introduced plant species, but few studies have compared invasive and noninvasive introduced clonal species to investigate which clonal traits may underlie invasiveness. To test the hypothesis that greater capacity to increase clonal growthviaphysiological integration of connected ramets increases invasiveness in clonal plants, we compared the effects of severing connections on accumulation of mass in the two species of the creeping, succulent, perennial, herbaceous genusCarpobrotusthat have been introduced on sand dunes along the Pacific Coast of northern California, the highly invasive speciesCarpobrotus edulisand the co-occurring, noninvasive speciesCarpobrotus chilensis.     

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.     

Trait values and not invasive status determine competitive outcomes between native and invasive species under varying soil nutrient availability

Invasion by exotic plants is often associated with nutrient enrichment of soils, particularly on soils of naturally low fertility. As a consequence, it is likely that the outcome of competitive interactions between native and invasive plants may be mediated by soil nutrient availability. We independently investigated competitive effect and response as well as the occurrence of asymmetric competition among native and invasive plants on soils of varying nutrient availability, using a glasshouse experiment. Seedlings of eight co‐occurring pairs of invasive and native species from low fertility Hawkesbury Sandstone‐derived soil were grown under low and high nutrient availability. We tested the hypotheses that native species would be competitively superior at low nutrient availability and have trait values associated with a resource conservation strategy while invasive species would be competitively superior at high nutrient availability and have trait values associated with a resource acquisition strategy. We found that nutrient availability did not mediate competitive interactions between invasive and native species. Instead, two invasive and one native species were always competitively superior irrespective of nutrient availability. Competitively superior species displayed a mixture of both resource conservation and acquisition strategies at low and high nutrient availability. In support of previous studies, we found that the a priori classification of invasive and native species does not predict competitive superiority at varying nutrient levels. Rather, species specific differences in trait values provide a competitive advantage in response to nutrient availability.     

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.

     

养分异质条件下结缕草克隆分株生长、碳水化合物及可溶性蛋白的生理整合

通过对连接和断开的结缕草分株进行不同的养分处理,研究了养分异质条件下结缕草克隆分株生长、碳水化合物及可溶性蛋白含量的变化.结果表明: 在养分异质条件下,处于中、高养分水平的母株可以显著提高与其相连子株的地上、地下及总生物量,高养分下分别提高32.5%、22.1%和24.8%,降低根冠比、可溶性糖及非结构性碳水化合物(NSC)含量,高养分下分别降低7.7%、15.2%和13.1%,但是对淀粉、纤维素、可溶性蛋白含量无显著影响.处于中、高养分水平的子株对与其相连的母株生长、碳水化合物及可溶性蛋白含量没有显著影响.养分异质条件下结缕草母株对子株生物量、根冠比、可溶性糖及NSC含量有明显的生理整合,其整合强度与养分梯度呈正比,而对淀粉、纤维素及可溶性蛋白含量没有显现显著的生理整合效应.子株对母株各项指标都没有显著的生理整合,结缕草母株和子株间是一种单向的生理整合.     

海口城市空间差异对外来入侵植物分布格局的影响

【目的】城市是外来入侵物种引入和二次传播的主要地区,城市环境中的入侵植物会对生物多样性和社会经济造成影响。阐明城市空间差异对入侵植物分布格局的影响,可为海口市生物入侵科学管理提供参考。【方法】以海口市主城区为研究区域,通过对461块随机样方的调查,记录外来入侵植物种类、盖度、平均株高和样方环境信息。从入侵植物的样方发生率、物种组成、丰富度和多度等方面,比较其在城市化梯度和土地利用类型上的发生差异和分布特征。【结果】(1)调查共记录外来入侵植物36种,隶属13科31属,其中菊科为优势科;以一年生和原产地为热带美洲的草本居多。(2)城市核心区和城市边缘区的入侵植物丰富度显著高于城市郊区,但城市郊区的多度显著高于城市核心区和城市边缘区。(3)在居住区、交通用地、空闲地、商业/单位/工矿用地中,入侵植物的发生概率与不透水面百分比呈负相关,而在林地、农业用地和城市公共绿地中,入侵植物更容易发生在不透水面百分比较高的区域。【结论】城市化梯度和土地利用类型均显著影响入侵植物的分布格局,不透水面的比例作为一个重要的解释因子,可以为入侵植物的发生提供预判参考。     

Clonal integration affects growth, photosynthetic efficiency and biomass allocation, but not the competitive ability, of the alien invasive Alternanthera philoxeroides under severe stress

Background and Aims: Many notorious alien invasive plants are clonal, but littleis known about some roles and aspects of clonal integration.Here, the hypothesis is tested that clonal integration affectsgrowth, photosynthetic efficiency, biomass allocation and competitiveability of the exotic invasive weed Alternanthera philoxeroides(Amaranthaceae). Methods: The apical parts of Alternanthera were grown either with orwithout the lawn grass Schedonorus phoenix (tall fescue) andtheir stolon connections to the basal parts grown without competitorswere either severed or left intact. Key Results: Competition greatly reduced the maximum quantum yield of photosystemII (F_v/F_m) and growth (biomass, number of ramets and leaves,total stolon length and total leaf area) of the apical Alternanthera,but not the biomass of S. phoenix. Stolon connections significantlyincreased F_v/F_m and growth of Alternanthera. However, such effectson growth were smaller with than without competition and stolonconnections did not alter the relative neighbour effect of Alternanthera.Stolon connections increased Alternanthera's biomass allocationto roots without competition, but decreased it with competition. Conclusions: Clonal integration contributed little to Alternanthera's competitiveability, but was very important for Alternanthera to exploreopen space. The results suggest that the invasiveness of Alternantheramay be closely related to clonal integration.     

匍匐茎草本植物形态可塑性、整合作用与觅食行为研究进展

综述了匍匐茎型克隆植物在形态可塑性、整合作用及觅食行为方面的研究进展。资源斑块性分布是生境异质性的特征之一,适应于异质性生境,匍匐茎植物对环境资源表现了一系列可塑性反应。本文着重从匍匐茎植物对光、水、肥的可塑性反应及其整合作用以及觅食行为等方面的研究进行总结分析,以期对匍匐茎型克隆植物进行更广泛深入的研究。     

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.     

西双版纳外来入侵植物及其共存种叶片氮、磷化学计量特征

诊断外来植物和本地种的资源利用方式是入侵生态学研究的热点问题。叶片氮(N)、磷(P)含量和化学计量特征可以反映受侵地区植物的N、P吸收能力和限制状况, 为把握外来植物的入侵能力、为本地植物共存或消失的机制提供基础科学依据。该研究以我国西南地区典型外来入侵植物(飞机草(Chromolaena odorata)、紫茎泽兰(又叫破坏草, Ageratina adenophora))及其共存的本地植物为对象, 探讨不同入侵条件下入侵种及其共存的本地植物的N、P利用策略。在云南西双版纳孔明山研究区, 调查了无入侵条件下和飞机草与紫茎泽兰的不同入侵程度(按入侵种生物量比例划分)下的物种数量和生物量, 分析了主要植物的叶片N、P含量和N:P值。结果显示: 尽管群落地上生物量随飞机草和紫茎泽兰入侵程度增加而增加, 但本地植物种数随飞机草和紫茎泽兰入侵程度增加而显著减少。飞机草与紫茎泽兰的叶片N、P平均含量无显著差异, 但均显著高于无入侵条件下的本地植物以及与其共存的本地植物。两种入侵植物的N、P含量均随它们所占样方总生物量的比例增大而升高, 本地植物N含量也有相似的变化趋势。当对比入侵和无入侵两类样方的同种植物N、P和N:P变化时发现多数本地种叶片P含量呈降低趋势, N含量和N:P呈升高趋势。根据叶片N、P绝对含量和N:P值及其随入侵的变化规律, 推测入侵可能提高了植物的N可利用性, 但本地植物仍然受N限制; 入侵植物N:P值总体小于10与其具有相对于N吸收的较高的P吸收能力有关。该研究揭示了西南典型外来入侵植物具有较强的N、P吸收富集能力。     

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.     

异质性光照条件下匍匐茎草本野草莓的克隆整合

通过盆栽实验,研究了匍匐茎草本野草莓在异质性光照条件下的克隆整合。结果显示克隆整合显著增强了野草莓胁迫分株段的生长,损—益分析表明未受胁迫分株没有显著损耗,整个克隆片段的生长得到显著提高。在局部遮荫处理,克隆整合对克隆形态可塑性的修饰作用没有观察到。最后,讨论了克隆植物对环境的生态适应意义。     

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。     

Clonal plants in a spatially heterogeneous environment: effects of integration on Serengeti grassland response to defoliation and urine-hits from grazing mammals

It has been hypothesized that clonal integration between ramets inenvironments with spatially variable rates of herbivory and nutrientavailabilities leads to increased growth and fitness in the genet. An increasein genet growth could potentially influence ecosystem processes such as primaryproductivity and nutrient cycling. I tested the idea that clonal integrationwould lead to greater aboveground productivity, compensatory response todefoliation, and N re-distribution in a factorial experiment in fieldplots in the Serengeti Ecosystem, Tanzania. Each plot had either all stolonssevered or left intact, had repeated defoliation (to simulate grazing bythe African buffalo) or none, and was either located next to a plot thatreceived urea (to simulate a urine-hit) or next to a plot thatremained untreated. Plots that received stolon severing treatments had32% less peak biomass than did connected control plots, and this suggeststhat plants grew better when ramets remained connected. However, compensatoryresponse by plots to repeated defoliation was inconsistent with the hypothesizedbenefits of ramet integration at the ecosystem level: productivityresponse to defoliation was similar between plots with connected vs. severedramets (i.e. no stolon severing × defoliation interaction wasfound). When averaged across other treatments, defoliated plots hadincreased productivity compared to nondefoliated plots during the growingseason. Thus, ramet connection and defoliation did increase productivity whenthey were considered alone, but productivity response to defoliation wasunaffected by ramet connections. Urea additions, which led to a 78%increase in productivity in adjacent urine-hits, had noconsistent effect on productivity but did increase leaf percent N in adjoiningstudy plots. Thus, in the Serengeti, urine hits probably have very localizedeffects on productivity during the initial growing season.     

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

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

崩岗生态修复不同人工林林下入侵植物和本土植物对群落稳定性的影响

探寻基于自然的崩岗生态修复群落构建方案,以广东三岭山国家森林公园崩岗区早期恢复阶段的4种(湿地松Pinus elliottii、尾叶桉Eucalyptus urophylla、大叶相思Acacia auriculaeformis和樟树Cinnamomum camphora)林分为对象,进行样方调查,以林下入侵植物和本土植物作为切入点,以物种多样性指数、生态位宽度、生态位重叠度、生态响应指数和种间联结指数探究不同林分的群落稳定性,优化林下植物配置。结果表明：(1)4种林分中共有林下植物104种,隶51科92属,其中菊科(Asteraceae)种类最多,有33种;其次是禾本科(Poaceae),有24种。林下本土植物数目为湿地松>尾叶桉>樟树>大叶相思,林下入侵植物数目为湿地松=樟树>大叶相思>尾叶桉。与其他三种林分相比,尾叶桉林的林下入侵物种数目、多样性指数和生态位宽度均最少,同时入侵植物与本土植物的生态位重叠度也最小。(2)林下植物群落未来发展趋势最佳的是湿地松林,其次是尾叶桉林、樟树林,最差为大叶相思林,且除大叶相思林外,其它3种林分正向发展的平均速率大...