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1. The stress-gradient hypothesis (SGH) predicts that the frequency of facilitative and competitive interactions will vary inversely across abiotic stress gradients, with facilitation being more common in conditions of high abiotic stress relative to more benign abiotic conditions. With notable exceptions, most tests of the SGH have studied the interaction between a single pair or a few pairs of species, and thus have evaluated shifts in the magnitude and direction of pair-wise interactions along stress gradients, rather than shifts in the general frequency of interactions. 2. The SGH has been supported by numerous studies in many ecosystems, has provided a crucial foundation for studying the interplay between facilitation and competition in plant communities, and has a high heuristic value. However, recent empirical research indicates that factors like the variation among species and the nature of the stress gradient studied add complexity not considered in the SGH, creating an opportunity to extend the SGH's general conceptual framework. 3. We suggest that one approach for extending the SGH framework is to differentiate between the original idea of how 'common' interactions might be along stress gradients and the ubiquitous empirical approach of studying shifts in the strength of pair-wise interactions. Furthermore, by explicitly considering the life history of the interacting species (relative tolerance to stress vs. competitive ability) and the characteristics of the stress factor (resource vs. non-resource) we may be able to greatly refine specific predictions relevant to the SGH. 4. We propose that the general pattern predicted by the SGH would hold more frequently for some combinations of life histories and stress factor, particularly when the benefactor and beneficiary species are mostly competitive and stress-tolerant, respectively. However, we also predict that other combinations are likely to yield different results. For example, the effect of neighbours can be negative at both ends of the stress gradient when both interacting species have similar 'competitive' or 'stress-tolerant' life histories and the abiotic stress gradient is driven by a resource (e.g. water). 5. Synthesis. The extension of the SGH presented here provides specific and testable hypotheses to foster research and helps to reconcile potential discrepancies among previous studies. It represents an important step in incorporating the complexity and species-specificity of potential outcomes into models and theories addressing how plant-plant interactions change along stress gradients.

1. Traditionally, techniques of plant manipulation during restoration have focused on the reduction of competition by 'problematic' existing vegetation. However, the increasing recognition of facilitation as a main process regulating the composition of communities has brought a change in the practice of restoration towards a better awareness of the benefits inherent to conserving neighbouring vegetation. 2. Here, I provide the results of a meta-analysis of published studies that have manipulated interactions among plants with the objective of restoring degraded terrestrial systems. I created four different data sets corresponding to the variables most commonly used to measure plant performance (i.e. emergence, survival, growth and density), and asked whether the benefits of facilitation as a restoration tool vary depending on the study duration, the life-form of the neighbour and target species, and the ecosystem type. 3. Neighbour effects varied strongly among performance estimators. Positive effects were frequently found for emergence and survival, whereas neutral or negative interactions predominated for growth and density. 4. No clear support existed for a relationship between study duration and neighbour effect. 5. The life-form of the interacting species, particularly of neighbours, largely influenced the interaction outcome. Herbs had strong negative effects, especially on other herb species, whereas shrubs had large facilitative effects, especially on trees. 6. Semiarid and tropical systems showed in general more positive neighbour effects than wetlands and particularly mesic temperate systems, where negative interactions predominated. However, these results were largely influenced by the over-representation of herb species in wetlands and temperate habitats, survival facilitation being found in all systems when only woody species were considered. 7. Synthesis. Pre-existing vegetation can have large impacts on species establishment in degraded habitats. Inhibition predominates in herbaceous communities typical of early-successional stages, whereas facilitation prevailes in communities dominated by shrubs and trees. Even productive systems (e.g. mesic temperate habitats) appear suitable for the application of facilitation as a restoration tool of woody communities. Whereas restoring herbaceous communities seems largely reliable on removal techniques, augmenting populations of nurse shrubs and trees should be considered a promising strategy for restoring woody late-successional communities.

1. Plants mediate multiple interactions between below‐ground (BG) and above‐ground (AG) heterotrophic communities that have no direct physical contact. These interactions can be positive or negative from the perspective of each player, can go from the BG to the AG community or vice versa, and comprise representatives of different phyla. Here we highlight emerging general patterns and discuss future research directions. 2. Ecologists initially postulated that root herbivores induce general stress responses, which increase the levels of primary (nutritional) compounds in the undamaged plant compartment and thereby facilitate future attack by AG herbivores. However, damage can also reduce the levels of primary compounds or increase contents of secondary (defensive) metabolites. Both effects may cause resistance phenotypes that play an important role in mediating BG-AG interactions. Systemically induced resistance does not only affect other herbivores but also pathogens in the AG and BG compartment and may inhibit beneficial organisms such as natural enemies of herbivores, microbial root symbionts and pollinators. Conversely, symbiotic mutualists such as mycorrhiza and rhizobia may affect AG and BG defence levels. Finally, BG-AG interactions may be costly if they impede optimal defence strategies in the undamaged compartment. 3. Synthesis. In order to better understand the adaptive value of BG-AG induced responses for the players involved and to identify the driving evolutionary forces, we need a better integration of studies at the community level with experiments on model systems that allow unravelling the genetic and physiological mechanisms of BG-AG interactions. Experiments preferably should be carried out at realistic densities and using the natural temporal sequence at which the various associations are established, because we can expect plants to be adapted only to events that are common over evolutionary time spans. Detailed mechanistic knowledge will help to reproduce relevant interactions in experiments that study multiple species in the field. This step will ultimately allow us to evaluate the importance of plant‐mediated interactions between BG and AG communities for the fitness of the species involved and for the structuring of natural communities.

Environmental conditions and plant size may both alter the outcome of inter-specific plant– plant interactions, with seedlings generally facilitated more strongly than larger individuals in stressful habitats. However, the combined impact of plant size and environmental severity on interactions is poorly understood. Here, we tested explicitly for the first time the hypothesis that ontogenetic shifts in interactions are delayed under increasingly severe conditions by examining the interaction between a grass, Agrostis magellanica, and a cushion plant, Azorella selago, along two severity gradients. The impact of A. selago on A. magellanica abundance, but not reproductive effort, was related to A. magellanica size, with a trend for delayed shifts towards more negative interactions under greater environmental severity. Intermediate-sized individuals were most strongly facilitated, leading to differences in the size-class distribution of A. magellanica on the soil and on A. selago. The A. magellanica size-class distribution was more strongly affected by A. selago than by environmental severity, demonstrating that the plant–plant interaction impacts A. magellanica population structure more strongly than habitat conditions. As ontogenetic shifts in plant–plant interactions cannot be assumed to be constant across severity gradients and may impact species population structure, studies examining the outcome of interactions need to consider the potential for size- or age-related variation in competition and facilitation.

1. The idea that the role of facultative interactions increases as environmental conditions become more stressful has become a ruling paradigm in ecology. Here, we review three reasons why positive interactions may actually be more prominent than generally thought under moderately stressful rather than under extreme conditions. 2. First, there is evidence that in some communities the net effect of amelioration of shortage of a limiting resource, such as water under the canopy of nurse plants, may be beneficial under moderate conditions whereas it can be overruled by increased competition for the same resource in very harsh environments. 3. Secondly, we show that even in situations where the relative role of facilitation increases monotonically with stress, the absolute effect should as a rule be largest at intermediately stressful conditions. This is because under the harshest conditions, facultative amelioration of conditions is insufficient to allow growth altogether. Therefore, while facilitation will expand the range of conditions where an organism may occur, the largest absolute effects on biomass will always occur under less stressful conditions. 4. A third reason why facilitation may be more important under moderate conditions than previously thought is that in any ecosystem, the suite of organisms is adapted to local conditions. This implies that even under conditions that appear benign, facilitation may play an unexpectedly large role as organisms are simply more sensitive than those found under harsher overall conditions. 5. Synthesis. We argue that while facilitation will extend the range of conditions where an organism can occur, it should also boost performance of the species well into the more moderate range of conditions. Broadening our search image for facultative effects towards milder environments will reveal wider than expected prevalence of positive interactions and their effects on stability and diversity in nature.

1. Environmental gradients may influence a plant's physiological status and morphology, which in turn may affect plant-plant interactions. However, little is known about the relationship between environmental variation, physiological and morphological variability of plants and variation in the balance between competition and facilitation. 2. Mountain ranges in dry environments have opposing altitudinal environmental gradients of temperature and aridity, which limit plant growth at high and low elevations. This makes them particularly suitable for exploring the relationships between environmental conditions, plant phenotype and plant-plant interactions. We hypothesized that different environmental Stressors will differently affect the physiological status of a nurse plant. This, then, manifests itself as variation in nurse plant morphological traits, which in turn mediates plant-plant interactions by altering microhabitat conditions for the nurse and associated species. 3. In an observational study, we measured a series of functional traits of Arenaria tetraquetra cushions as indicators of its physiological status (e.g. specific leaf area, relative water content) and morphology (e.g. cushion compactness, size). Measurements were taken along the entire elevation range where A. tetraquetra occurs. Furthermore, we analysed how these functional traits related to soil properties beneath cushions and the number of associated species and individuals compared with open areas. 4. Cushions at high elevation showed good physiological status; they were compact and large, had higher soil water and organic matter content compared with open areas and showed the strongest facilitation effect of the whole elevation gradient — that is, the highest increase in species richness and abundance of beneficiaries compared with open areas. Physiological data at low elevation indicated stressful abiotic conditions for A. tetraquetra, which formed loose and small cushions. These cushions showed less improved soil conditions and had reduced facilitative effects compared with those at high elevation. 5. Synthesis. Functional traits of the nurse species varied distinctively along the two opposing stress gradients, in parallel to the magnitude of differences in microenvironmental conditions between cushions and the surrounding open area, and also to the facilitation effect of cushions. Our data, therefore, provides a strong demonstration of the generally overlooked importance of a nurse plant's vigour and morphology for its facilitative effects.

Some plant species are able to distinguish between neighbours of different genetic identity and attempt to pre‐empt resources through root proliferation in the presence of unrelated competitors, but avoid competition with kin. However, studies on neighbour recognition have met with some scepticism because the mechanisms by which plants identify their neighbours have remained unclear. In order to test whether root exudates could mediate neighbour recognition in plants, we performed a glasshouse experiment in which plants of Deschampsia caespitosa were subjected to root exudates collected from potential neighbours of different genetic identities, including siblings and individuals belonging to the same or a different population or species. Our results show that root exudates can carry specific information about the genetic relatedness, population origin and species identity of neighbours, and trigger different responses at the whole root system level and at the level of individual roots in direct contact with locally applied exudates. Increased root density was mainly achieved through changes in morphology rather than biomass allocation, suggesting that plants are able to limit the energetic cost of selfish behaviour. This study reveals a new level of complexity in the ability of plants to interpret and react to their surroundings.

1. Declines in availability of plant resources to pollinators are a major cause of pollinator loss. The management of plant communities to enhance floral resources is often proposed as a way to sustain pollinator populations. Nectar, the main energetic resource for pollinators, plays a central role in behaviour and composition of pollinator communities. Abiotic and biotic factors are known to influence nectar traits at both the species and community levels, but the impact of plant community composition itself has never been investigated. 2. Below-ground interactions in plant communities can induce changes in plant development through (i) plant-derived litter amendment of the soil and (ii) competition for soil resources between plants. We tested how plant below-ground interactions affect above-ground nectar traits involved in plant attractiveness to pollinators. 3. A short-term pot experiment was carried out with three temperate grassland species Mimulus guttatus, Lamium amplexicaule, and Medicago sativa, showing distinct litter stoichiometry and competitive abilities for soil resources. Litter amendment (none, mono and tri-specific litter) and plant interaction treatments (monocultures, two-and three-species mixtures) were crossed in a factorial design. 4. Litter amendment to the soil led to an increase in total nectar sugar content in L. amplexicaule plants but not in the two other species. We also found that the presence of M. guttatus, a competitive species, reduced the total nectar sugar content in L. amplexicaule through a concomitant decrease in nectar volume per flower and in floral display size, but not in other species. Species-specific responses of nectar traits to variation in soil nitrogen availability were thus observed, suggesting consequences for plant species and community attractiveness to pollinators. However, we did not find evidence that the legume M. sativa affected nectar traits of any neighbouring plants. 5. Synthesis. Our results demonstrate that litter inputs and competition between plants for soil resources can alter nectar traits linked to plant attractiveness to pollinators. This supports the idea that below-ground plant-plant interactions for soil resources can influence above-ground plant-plant interactions for pollination services. This offers promising perspectives in studying the role of below-ground-above-ground interactions on higher trophic levels.

1. The stress-gradient hypothesis (SGH) predicts an increasing importance of facilitative mechanisms relative to competition along gradients of increasing environmental stress. Although developed across a variety of ecosystems, the SGH's relevance to the dynamic tree—grass systems of global savannas remains unclear. Here, we present a meta-analysis of empirical studies to explore emergent patterns of tree—grass relationships in global savannas in the context of the SGH. 2. We quantified the net effect of trees on understorey grass production relative to production away from tree canopies along a rainfall gradient in tropical and temperate savannas and compared these findings to the predictions of the SGH. We also analysed soil and plant nutrient concentrations in subcanopy and open-grassland areas to investigate the potential role of nutrients in determining grass production in the presence and absence of trees. 3. Our meta-analysis revealed a shift from net competitive to net facilitative effects of trees on subcanopy grass production with decreasing annual precipitation, consistent with the SGH. We also found a significant difference between sites from Africa and North America, suggesting differences in tree—grass interactions in the savannas of tropical and temperate regions. 4. Nutrient analyses indicate no change in nutrient ratios along the rainfall gradient, but consistent nutrient enrichment under tree canopies. 5. Synthesis. Our results help to resolve questions about the SGH in semi-arid systems, demonstrating that in mixed tree—grass systems, trees facilitate grass growth in drier regions and suppress grass growth in wetter regions. Relationships differ, however, between African and North American sites representing tropical and temperate bioclimates, respectively. The results of this meta-analysis advance our understanding of tree—grass interactions in savannas and contribute a valuable data set to the developing theory behind the SGH.