Plants can have positive effects on each other. For example, the accumulation of nutrients, provision of shade, amelioration of disturbance, or protection from herbivores by some species can enhance the performance of neighbouring species. Thus the notion that the distributions and abundances of plant species are independent of other species may be inadequate as a theoretical underpinning for understanding species coexistence and diversity. But there have been no large-scale experiments designed to examine the generality of positive interactions in plant communities and their importance relative to competition. Here we show that the biomass, growth and reproduction of alpine plant species are higher when other plants are nearby. In an experiment conducted in subalpine and alpine plant communities with 115 species in 11 different mountain ranges, we find that competition generally, but not exclusively, dominates interactions at lower elevations where conditions are less physically stressful. In contrast, at high elevations where abiotic stress is high the interactions among plants are predominantly positive. Furthermore, across all high and low sites positive interactions are more important at sites with low temperatures in the early summer, but competition prevails at warmer sites.

Summary 1.Once neglected, the role of facilitative interactions in plant communities has received considerable attention in the last two decades, and is now widely recognized. It is timely to consider the progress made by research in this field. 2. We review the development of plant facilitation research, focusing on the history of the field, the relationship between plant-plant interactions and environmental severity gradients, and attempts to integrate facilitation into mainstream ecological theory. We then consider future directions for facilitation research. 3. With respect to our fundamental understanding of plant facilitation, clarification of the relationship between interactions and environmental gradients is central for further progress, and necessitates the design and implementation of experiments that move beyond the clear limitations of previous studies. 4. There is substantial scope for exploring indirect facilitative effects in plant communities, including their impacts on diversity and evolution, and future studies should connect the degree of non-transitivity in plant competitive networks to community diversity and facilitative promotion of species coexistence, and explore how the role of indirect facilitation varies with environmental severity. 5. Certain ecological modelling approaches (e.g. individual-based modelling), although thus far largely neglected, provide highly useful tools for exploring these fundamental processes. 6. Evolutionary responses might result from facilitative interactions, and consideration of facilitation might lead to re-assessment of the evolution of plant growth forms. 7. Improved understanding of facilitation processes has direct relevance for the development of tools for ecosystem restoration, and for improving our understanding of the response of plant species and communities to environmental change drivers. 8. Attempts to apply our developing ecological knowledge would benefit from explicit recognition of the potential role of facilitative plant-plant interactions in the design and interpretation of studies from the fields of restoration and global change ecology. 9. Synthesis: Plant facilitation research provides new insights into classic ecological theory and pressing environmental issues. Awareness and understanding of facilitation should be part of the basic ecological knowledge of all plant ecologists.

SummaryIntercropping is a farming practice involving two or more crop species, or genotypes, growing together and coexisting for a time. On the fringes of modern intensive agriculture, intercropping is important in many subsistence or low-input/resource-limited agricultural systems. By allowing genuine yield gains without increased inputs, or greater stability of yield with decreased inputs, intercropping could be one route to delivering 'sustainable intensification'. We discuss how recent knowledge from agronomy, plant physiology and ecology can be combined with the aim of improving intercropping systems. Recent advances in agronomy and plant physiology include better understanding of the mechanisms of interactions between crop genotypes and speciesfor example, enhanced resource availability through niche complementarity. Ecological advances include better understanding of the context-dependency of interactions, the mechanisms behind disease and pest avoidance, the links between above-and below-ground systems, and the role of microtopographic variation in coexistence. This improved understanding can guide approaches for improving intercropping systems, including breeding crops for intercropping. Although such advances can help to improve intercropping systems, we suggest that other topics also need addressing. These include better assessment of the wider benefits of intercropping in terms of multiple ecosystem services, collaboration with agricultural engineering, and more effective interdisciplinary research.

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International audiencePlant communities have traditionally been viewed as either a random collection of individuals or as organismal entities. For most ecologists however, neither perspective provides a modern comprehensive view of plant communities, but we have yet to formalize the view that we currently hold. Here, we assert that an explicit re-consideration of formal community theory must incorporate interactions that have recently been prominent in plant ecology, namely facilitation and indirect effects among competitors. These interactions do not suppport the traditional individualistic perspective. We believe that rejecting strict individualistic theory will allow ecologists to better explain variation occurring at different spatial scales, synthesize more general predictive theories of community dynamics, and develop models for community-level responses to global change. Here, we introduce the concept of the integrated community (IC) which proposes that natural plant communities range from highly individualistic to highly interdependent depending on synergism among: (i) stochastic processes, (ii) the abiotic tolerances of species, (iii) positive and negative interactions among plants, and (iv) indirect interactions within and between trophic levels. All of these processes are well accepted by plant ecologists, but no single theory has sought to integrate these different processes into our concept of communities