植物邻体间的正相互作用

植物间的正负相互作用是构建植被群落的重要因素,也是群落生态学研究的中心内容之一。近20a来,植物间正相互作用的研究得到快速发展。综述了正相互作用的定义,不同植物群落中的直接、间接正相互作用及其发生机制,正相互作用研究的实验和模型方法,正负相互作用随胁迫梯度的变化及正相互作用对群落构建的影响。探讨了正相互作用研究前景:(1)进一步理解正负相互作用的平衡及其对群落构建的影响;(2)加深对全球变暖背景下的正相互作用的认识;(3)需把正相互作用研究同进化联系起来;(4)充分发挥正相互作用在生态系统中的推动力作用,把正相互作用应用到生态恢复中,为恢复退化生态系统服务。     

根系间的相互作用——竞争与互利

植物根系间的相互作用分为竞争和互利两种形式 ,它是决定植物群落动态变化和群落结构的重要因素。根系间的竞争包括植株个体自身根系的竞争以及个体与个体根系间 (同种或异种 )的竞争两方面 ,前者的发生在农林系统中是不可避免的并且很难调控 ,后者可以分为种间植物根系的竞争和种内植物根系的竞争。还阐述了根系的竞争能力和与其密切相关的根系生长率、根组织的新陈代谢、植物的生长形式和根系的空间结构等植物特性 ,同时介绍了根系对水和养分的竞争机理、形式、影响以及竞争强度计算方法。接着具体分析包括根系错位在内的各种根系互利现象和相关机理。影响根系间相互作用的限制性因子有土壤营养的异质性、大气 CO2 浓度、地下草食生物、根系生产力和生物量、根系结构、形态和生理调节、土壤养分的扩散性以及植物间距等。随着科技的进步和各门学科的发展 ,未来根系的研究方向主要体现在结合实践优化农林系统中不同物种间的作用关系、预测根系竞争在全球气候变化下的发展规律、更新实验研究方法及手段研究作用机理等 3个方面。     

Positive interactions, discontinuous transitions and species coexistence in plant communities

The population and community level consequences of positive interactions between plants remain poorly explored. In this study we incorporate positive resource-mediated interactions in classic resource competition theory and investigate the main consequences for plant population dynamics and species coexistence. We focus on plant communities for which water infiltration rates exhibit positive dependency on plant biomass and where plant responses can be improved by shading, particularly under water limiting conditions. We show that the effects of these two resource-mediated positive interactions are similar and additive. We predict that positive interactions shift the transition points between different species compositions along environmental gradients and that realized niche widths will expand or shrink. Furthermore, continuous transitions between different community compositions can become discontinuous and bistability or tristability can occur. Moreover, increased infiltration rates may give rise to a new potential coexistence mechanism that we call controlled facilitation.     

Do biotic interactions shape both sides of the humped‐back model of species richness in plant communities?

A humped-back relationship between species richness and community biomass has frequently been observed in plant communities, at both local and regional scales, although often improperly called a productivity-diversity relationship. Explanations for this relationship have emphasized the role of competitive exclusion, probably because at the time when the relationship was first examined, competition was considered to be the significant biotic filter structuring plant communities. However, over the last 15 years there has been a renewed interest in facilitation and this research has shown a clear link between the role of facilitation in structuring communities and both community biomass and the severity of the environment. Although facilitation may enlarge the realized niche of species and increase community richness in stressful environments, there has only been one previous attempt to revisit the humped-back model of species richness and to include facilitative processes. However, to date, no model has explored whether biotic interactions can potentially shape both sides of the humped-back model for species richness commonly detected in plant communities. Here, we propose a revision of Grime's original model that incorporates a new understanding of the role of facilitative interactions in plant communities. In this revised model, facilitation promotes diversity at medium to high environmental severity levels, by expanding the realized niche of stress-intolerant competitive species into harsh physical conditions. However, when environmental conditions become extremely severe the positive effects of the benefactors wane (as supported by recent research on facilitative interactions in extremely severe environments) and diversity is reduced. Conversely, with decreasing stress along the biomass gradient, facilitation decreases because stress-intolerant species become able to exist away from the canopy of the stress-tolerant species (as proposed by facilitation theory). At the same time competition increases for stress-tolerant species, reducing diversity in the most benign conditions (as proposed by models of competition theory). In this way our inclusion of facilitation into the classic model of plant species diversity and community biomass generates a more powerful and richer predictive framework for understanding the role of plant interactions in changing diversity. We then use our revised model to explain both the observed discrepancies between natural patterns of species richness and community biomass and the results of experimental studies of the impact of biodiversity on the productivity of herbaceous communities. It is clear that explicit consideration of concurrent changes in stress-tolerant and competitive species enhances our capacity to explain and interpret patterns in plant community diversity with respect to environmental severity.     

Positive interactions in plant communities and the individualistic-continuum concept

The individualistic nature of communities is held as a fundamental ecological tenet by many ecologists. The empirical rationale for the individualistic hypothesis is largely based on gradient analyses in which plant species are almost always found to be arranged independently of one another in “continua” along environmental gradients. However, continua are correlative patterns and do not identify the processes that determine them, and so they do not necessarily preclude the possibility of interdependent interactions within plant communities. For example, the common occurrence of positive interactions suggests that plant species may not always be distributed independently of each other. If the distributions and abundances of species are enhanced by the presence of other species, their organization is not merely a coincidence of similar adaptation to the abiotic environment. Interpretations of gradient analyses also appear to assume that interactions among species should be similar at all points along environmental axes, and that groups of species should be associated at all points on a gradient if interdependence is to be accepted. However, virtually all types of ecological interactions have been shown to vary with changes in the abiotic environment, and a number of field experiments indicate that positive effects become stronger as abiotic stress increases. Furthermore, interactions among plants have been shown to shift from competition to facilitation along environmental continua. Thus, significant interdependence may occur even when species do not fully overlap in distribution. Higher-order, indirect interactions between animals and plants, and among plants, also suggest that interdependence within communities occurs. Eliminating a species involved in an indirect interaction may not necessarily mean that its beneficiary will be eliminated from a community, but the prospect that the distribution and abundance of any species in a plant community may be positively affected by the effects that other species have on their competitors suggests that communities are organized by much more than “the fluctuating and fortuitous immigration of plants and an equally fluctuating and variable environment” as stated by Henry Gleason. The ubiquity of direct and indirect positive interactions within plant communities provides a strong argument that communities are more interdependent than current theories allow. Received: 17 February 1997 / Accepted: 23 May 1997     

植物间正相互作用对种群动态和群落结构的影响:基于个体模型的研究进展

植物间的相互作用对种群动态和群落结构有着重要的影响。大量的野外实验已经揭示了正相互作用(互利)在群落中的普遍存在及其重要性。为了弥补野外实验方法的不足, 模型方法被越来越多地应用于正相互作用及其生态学效应的研究中。该文基于个体模型研究, 探讨了植物间正相互作用对种群动态和群落结构的影响。介绍了植物间正相互作用的定义和发生机制、植物间相互作用与环境梯度的关系。正相互作用是指发生在相邻的植物个体之间, 至少对其中一个个体有益的相互作用。植物通过直接(生境改善或资源富集)或间接(协同防御等)作用使局部环境有利于邻体而发生正相互作用。胁迫梯度假说认为互利的强度或重要性随着环境胁迫度的增加而增加, 但是越来越多的经验研究认为胁迫梯度假说需要改进。以网格模型和影响域模型为例, 介绍了基于个体的植物间相互作用模型方法。基于个体模型, 对近年来国内外正相互作用对种群时间动态(如生物量-密度关系)、空间分布格局和群落结构(如群落生物量-物种丰富度关系)影响的研究进行了总结。指出未来的研究应集中在对正相互作用概念和机制的理解, 新的模型, 新的种群、群落, 甚至生态系统问题, 以及在全球变化背景下进行相关的研究。     

Positive interactions among plants

Experimental evidence for positive interactions, or facilitation, among plants has increased markedly during the last 10 years. Experiments documenting facilitation have been conducted in many diverse ecological systems, which suggests that positive interactions may be fundamental processes in plant communities. Here, I review the evidence for facilitation, the mechanisms by which facilitation operates, and the effects facilitation has on community structure. Facilitative mechanisms may act simultaneously with resource competition or allelopathy, and the overall effect of one species on another may be the product of multiple, complex interactions. Positive interactions may also determine community spatial patterns, permit coexistence, enhance diversity and productivity, and drive community dynamics. Once viewed as anecdotal and idiosyncratic, facilitation is now contributing to a more complete understanding of community structure and dynamics.     

Global shifts towards positive species interactions with increasing environmental stress

The study of positive species interactions is a rapidly evolving field in ecology. Despite decades of research, controversy has emerged as to whether positive and negative interactions predictably shift with increasing environmental stress as hypothesised by the stress‐gradient hypothesis (SGH). Here, we provide a synthesis of 727 tests of the SGH in plant communities across the globe to examine its generality across a variety of ecological factors. Our results show that plant interactions change with stress through an outright shift to facilitation (survival) or a reduction in competition (growth and reproduction). In a limited number of cases, plant interactions do not respond to stress, but they never shift towards competition with stress. These findings are consistent across stress types, plant growth forms, life histories, origins (invasive vs. native), climates, ecosystems and methodologies, though the magnitude of the shifts towards facilitation with stress is dependent on these factors. We suggest that future studies should employ standardised definitions and protocols to test the SGH, take a multi‐factorial approach that considers variables such as plant traits in addition to stress, and apply the SGH to better understand how species and communities will respond to environmental change.     

Interspecific interactions in phytophagous insects revisited: a quantitative assessment of competition theory

The importance of interspecific competition is a highly controversial and unresolved issue for community ecology in general, and for phytophagous insects in particular. Recent advancements, however, in our understanding of indirect (plant- and enemy-mediated) interactions challenge the historical paradigms of competition. Thus, in the context of this rapidly developing field, we re-evaluate the evidence for interspecific competition in phytophagous insects using a meta-analysis of published studies. Our analysis is specifically designed to test the assumptions underlying traditional competition theory, namely that competitive interactions are symmetrical, necessitate spatial and temporal co-occurrence, and increase in intensity as the density, phylogenetic similarity, and niche overlap of competing species increase. Despite finding frequent evidence for competition, we found very little evidence that plant-feeding insects conform to theoretical predictions for interspecific competition. Interactions were highly asymmetrical, similar in magnitude within vs. between feeding guilds (chewers vs. sap-feeders), and were unaffected by the quantity of resources removed (% defoliation). There was mixed support for the effects of phylogeny, spatial/temporal separation, and the relative strength of intra- vs. interspecific competition. Clearly, a new paradigm that accounts for indirect interactions and facilitation is required to describe how interspecific competition contributes to the organization of phytophagous insect communities, and perhaps to other plant and animal communities as well.     

Phylogenetic relatedness as a tool in restoration ecology: a meta-analysis

Biotic interactions assembling plant communities can be positive (facilitation) or negative (competition) and operate simultaneously. Facilitative interactions and posterior competition are among the mechanisms triggering succession, thus representing a good scenario for ecological restoration. As distantly related species tend to have different phenotypes, and therefore different ecological requirements, they can coexist, maximizing facilitation and minimizing competition. We suggest including phylogenetic relatedness together with phenotypic information as a predictor for the net effects of the balance between facilitation and competition in nurse-based restoration experiments. We quantify, by means of a Bayesian meta-analysis of nurse-based restoration experiments performed worldwide, the importance of phylogenetic relatedness and life-form disparity in the survival, growth and density of facilitated plants. We find that the more similar the life forms of neighbouring plants are the greater the positive effect of phylogenetic distance is on survival and density. This result suggests that other characteristics beyond life form are also contained in the phylogeny, and the larger the phylogenetic distance, the less is the niche overlap, and therefore the less is the competition. As a general rule, we can maximize the success of the nurse-based practices by increasing life-form disparity and phylogenetic distances between the neighbour and the facilitated plant.