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.

Summary 1.Interactions among neighbouring plants are often mediated by foraging choices of pollinators. For example, the presence of a conspicuous species may increase the number of pollinators attracted to its vicinity, indirectly increasing visitation rates also to neighbouring plants. Because pollinator choices are frequently density-dependent, the presence of a conspicuous species at high densities may also increase competition for pollination services. Additionally, models predict that plant density will interact with spatial distribution in manipulating the pollinator behaviour, yet experimental evidence for this effect is missing. 2. We performed a field experiment in which we introduced a highly conspicuous species in different densities and spatial configurations in a full-factorial manner into a species-rich meadow and studied its effect on neighbouring plants. 3. We showed that the highly conspicuous species strongly contributed to the attractiveness of its local patch and thus benefited its neighbours. However, because of the strong density effect, the conspicuous species changed its role and became a competitor for pollinators when its density increased. 4. We supported our theoretical assumptions and showed that when the introduced conspicuous species was regularly distributed among other plants in the patch, it increased visitation rate, and in some cases also seed set, to conspicuous neighbours relative to when it was aggregated, at least at low densities. 5. Synthesis. We suggest that complex interactions between density and spatial distribution of plant species at the patch scale are highly relevant for the interpretation of pollinator behaviour and therefore should be treated as factors of floral attractiveness in future studies.

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Summary1. An unresolved question in plant ecology is how the balance between positive and negative interactions changes with environmental conditions. Recently, the debate has been expanded by Brooker & Kikvidze (2008), reintroducing the importance, rather than the intensity, of interactions as the appropriate concept for empirical studies. Importance is the difference in performance with interactions relative to all affecting factors. 2. Positive interactions among plants (facilitation) are common in nature. However, the advocated importance indices fail to address facilitation and can thus not be applied to studying shifts in the balance between positive and negative interactions. The deficiencies of the current importance indices are both conceptual as well as mathematical. In particular, their maximum performance estimator is based on the assumption that it can only be attained in the absence of neighbours. 3. We suggest two major improvements. First, we use the target's overall maximum performance as a surrogate for the plant optimum, i.e. we allow the possibility of higher performance with neighbours (i.e. facilitation). Second, we propose an alternative approach to treat the factors affecting measured performance. The new index is limited in its range [)1, 1], is symmetrical for competition and facilitation, and it preserves the intuitive nature of the original index by Brooker & Kikvidze (2008). 4. Synthesis. The concept of interaction importance is useful for studying interactions along environmental gradients. In contrast to previous indices, our index accommodates facilitative interactions and thus offers a significant conceptual and methodological advancement. We advocate its use in future empirical studies.

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