Coadaptation of coexisting plants enhances productivity in an agricultural system.

Growing crops in more diverse crop systems (i.e., intercropping) is one way to produce food more sustainably. Even though intercropping, compared to average monocultures, is generally more productive, the full yield potential of intercropping might not yet have been achieved as modern crop cultivars are bred to be grown in monoculture. Breeding plants for more familiarity in mixtures, i.e., plants that are adapted to more diverse communities (i.e., adaptation) or even to coexist with each other (i.e., coadaptation) might have the potential to sustainably enhance productivity. In this study, the productivity benefits of familiarity through evolutionary adaptation and coevolutionary coadaptation were disentangled in a crop system through an extensive common garden experiment. Furthermore, evolutionary and coevolutionary effects on species-level and community-level productivity were linked to corresponding changes in functional traits. We found evidence for higher productivity and trait convergence with increasing familiarity with the plant communities. Furthermore, our results provide evidence for the coevolution of plants in mixtures leading to higher productivity of coadapted species. However, with the functional traits measured in our study, we could not fully explain the productivity benefits found upon coevolution. Our study investigated coevolution among randomly interacting plants and was able to demonstrate that coadaptation through coevolution of coexisting species in mixtures occurs and promotes ecosystem functioning (i.e., higher productivity). This result is particularly relevant for the diversification of agricultural and forest ecosystems, demonstrating the added value of artificially selecting plants for the communities they are familiar with.

Network motifs involving both competition and facilitation predict biodiversity in alpine plant communities.

Biological diversity depends on multiple, cooccurring ecological interactions. However, most studies focus on one interaction type at a time, leaving community ecologists unsure of how positive and negative associations among species combine to influence biodiversity patterns. Using surveys of plant populations in alpine communities worldwide, we explore patterns of positive and negative associations among triads of species (modules) and their relationship to local biodiversity. Three modules, each incorporating both positive and negative associations, were overrepresented, thus acting as "network motifs." Furthermore, the overrepresentation of these network motifs is positively linked to species diversity globally. A theoretical model illustrates that these network motifs, based on competition between facilitated species or facilitation between inferior competitors, increase local persistence. Our findings suggest that the interplay of competition and facilitation is crucial for maintaining biodiversity.

Nitrogen fixation by common beans in crop mixtures is influenced by growth rate of associated species.

BACKGROUND: Legumes can fix atmospheric nitrogen (N) and facilitate N availability to their companion plants in crop mixtures. However, biological nitrogen fixation (BNF) of legumes in intercrops varies largely with the identity of the legume species. The aim of our study was to understand whether BNF and concentration of plant nutrients by common bean is influenced by the identity of the companion plant species in crop mixtures. In this greenhouse pot study, common beans were cultivated with another legume (chickpea) and a cereal (Sorghum). We compared BNF, crop biomass and nutrient assimilation of all plant species grown in monocultures with plants grown in crop mixtures. RESULTS: We found beans to exhibit low levels of BNF, and to potentially compete with other species for available soil N in crop mixtures. The BNF of chickpeas however, was enhanced when grown in mixtures. Furthermore, biomass, phosphorous and potassium values of chickpea and Sorghum plants were higher in monocultures, compared to in mixtures with beans; suggesting competitive effects of beans on these plants. Concentration of calcium, magnesium and zinc in beans was higher when grown with chickpeas than with Sorghum. CONCLUSIONS: It is generally assumed that legumes benefit their companion plant species. Our study highlights the contrary and shows that the specific benefits of cereal-legume mixtures are dependent on the growth rate of the species concerned. We further highlight that the potential of legume-legume mixtures is currently undervalued and may play a strong role in increasing N use efficiency of intercrop-based systems.

Using plant traits to understand the contribution of biodiversity effects to annual crop community productivity.

Increasing biodiversity generally enhances productivity through selection and complementarity effects not only in natural, but also in agricultural, systems. However, the quest to explain why diverse cropping systems are more productive than monocultures remains a central goal in agricultural science. In a mesocosm experiment, we constructed monocultures, two- and four-species mixtures from eight crop species with or without fertilizer and both in temperate Switzerland and dry, Mediterranean Spain. We measured physical factors and plant traits and related these in structural equation models to selection and complementarity effects to explain seed yield differences between monocultures and mixtures. Increased crop diversity increased seed yield in Switzerland. This positive biodiversity effect was driven to almost the same extent by selection and complementarity effects, which increased with plant height and specific leaf area (SLA), respectively. Also, ecological processes driving seed yield increases from monocultures to mixtures differed from those responsible for seed yield increases through the diversification of mixtures from two to four species. Whereas selection effects were mainly driven by one species, complementarity effects were linked to larger leaf area per unit leaf weight. Seed yield increases due to mixture diversification were driven only by complementarity effects and were not mediated through the measured traits, suggesting that ecological processes beyond those measured in this study were responsible for positive diversity effects on yield beyond two-species mixtures. By understanding the drivers of positive biodiversity-productivity relationships, we can improve our ability to predict species combinations that enhance ecosystem functioning and can promote sustainable agricultural production.

Crop-weed relationships are context-dependent and cannot fully explain the positive effects of intercropping on yield.

Implementing sustainable weed control strategies is a major challenge in agriculture. Intercropping offers a potential solution to control weed pressure by reducing the resources available for weeds; however, available research on the relationship between crop diversity and weed pressure and its consequences for crop yield is not yet fully conclusive. In this study, we performed an extensive intercropping experiment using eight crop species and 40 different species mixtures to examine how crop diversity affects weed communities and how the subsequent changes in weeds influence crop yield. Mesocosm experiments were carried out under field conditions in Switzerland and in Spain, which differ drastically in terms of climate, soil and weed community, and included monocultures, two- and four-species mixtures, and a control treatment without crops. Weed communities were assessed in terms of biomass, species number and evenness, and community composition. Results indicate that intercropping reduces weed biomass and diversity in Spain but not in Switzerland. In Switzerland, despite the lack of a crop diversity effect on weeds, crop yield increased with crop species number. Moreover, in Switzerland, where soil resources were abundant, increasing crop yield correlated with reduced weed biomass. In Spain, where water and nutrients were limited, crop yield was not related to weed biomass or diversity. The presented research applies plant community ecology in the context of agricultural crop production systems. We demonstrate that, in our study, increased crop yield in mixtures was not due to increased weed suppression in diverse crop communities, and so must be the result of other ecological processes. We further show that crop-weed relationships vary across environmental conditions; more specifically, our study shows that weeds are less detrimental to crop yield in harsher environments compared to benign abiotic conditions, where alternative strategies are needed to control weed pressure and ensure the yield benefits provided by intercropping.

Plant domestication disrupts biodiversity effects across major crop types.

Plant diversity fosters productivity in natural ecosystems. Biodiversity effects might increase agricultural yields at no cost in additional inputs. However, the effects of diversity on crop assemblages are inconsistent, probably because crops and wild plants differ in a range of traits relevant to plant-plant interactions. We tested whether domestication has changed the potential of crop mixtures to over-yield by comparing the performance and traits of major crop species and those of their wild progenitors under varying levels of diversity. We found stronger biodiversity effects in mixtures of wild progenitors, due to larger selection effects. Variation in selection effects was partly explained by within-mixture differences in leaf size. Our results indicate that domestication might disrupt the ability of crops to benefit from diverse neighbourhoods via reduced trait variance. These results highlight potential limitations of current crop mixtures to over-yield and the potential of breeding to re-establish variance and increase mixture performance.

Increasing water availability and facilitation weaken biodiversity-biomass relationships in shrublands.

Positive biodiversity-ecosystem-functioning (BEF) relationships are commonly found in experimental and observational studies, but how they vary in different environmental contexts and under the influence of coexisting life forms is still controversial. Investigating these variations is important for making predictions regarding the dynamics of plant communities and carbon pools under global change. We conducted this study across 433 shrubland sites in northern China. We fitted structural equation models (SEMs) to analyze the variation in the species-richness-biomass relationships of shrubs and herbs along a wetness gradient and general liner models (GLMs) to analyze how shrub or herb biomass affected the species-richness-biomass relationship of the other life form. We found that the positive species-richness-biomass relationships for both shrubs and herbs became weaker or even negative with higher water availability, likely indicating stronger interspecific competition within life forms under more benign conditions. After accounting for variation in environmental contexts using residual regression, we found that the benign effect of greater facilitation by a larger shrub biomass reduced the positive species-richness-biomass relationships of herbs, causing them to become nonsignificant. Different levels of herb biomass, however, did not change the species-richness-biomass relationship of shrubs, possibly because greater herb biomass did not alter the stress level for shrubs. We conclude that biodiversity in the studied plant communities is particularly important for plant biomass production under arid conditions and that it might be possible to use shrubs as nurse plants to facilitate understory herb establishment in ecological restoration.

Plant interactions shape pollination networks via nonadditive effects.

Plants grow in communities where they interact with other plants and with other living organisms such as pollinators. On the one hand, studies of plant-plant interactions rarely consider how plants interact with other trophic levels such as pollinators. On the other, studies of plant-animal interactions rarely deal with interactions within trophic levels such as plant-plant competition and facilitation. Thus, to what degree plant interactions affect biodiversity and ecological networks across trophic levels is poorly understood. We manipulated plant communities driven by foundation species facilitation and sampled plant-pollinator networks at fine spatial scale in a field experiment in Sierra Nevada, Spain. We found that plant-plant facilitation shaped pollinator diversity and structured pollination networks. Nonadditive effects of plant interactions on pollinator diversity and interaction diversity were synergistic in one foundation species networks while they were additive in another foundation species. Nonadditive effects of plant interactions were due to rewiring of pollination interactions. In addition, plant facilitation had negative effects on the structure of pollination networks likely due to increase in plant competition for pollination. Our results empirically demonstrate how different network types are coupled, revealing pervasive consequences of interaction chains in diverse communities.

A global meta-analysis of the relative extent of intraspecific trait variation in plant communities.

Recent studies have shown that accounting for intraspecific trait variation (ITV) may better address major questions in community ecology. However, a general picture of the relative extent of ITV compared to interspecific trait variation in plant communities is still missing. Here, we conducted a meta-analysis of the relative extent of ITV within and among plant communities worldwide, using a data set encompassing 629 communities (plots) and 36 functional traits. Overall, ITV accounted for 25% of the total trait variation within communities and 32% of the total trait variation among communities on average. The relative extent of ITV tended to be greater for whole-plant (e.g. plant height) vs. organ-level traits and for leaf chemical (e.g. leaf N and P concentration) vs. leaf morphological (e.g. leaf area and thickness) traits. The relative amount of ITV decreased with increasing species richness and spatial extent, but did not vary with plant growth form or climate. These results highlight global patterns in the relative importance of ITV in plant communities, providing practical guidelines for when researchers should include ITV in trait-based community and ecosystem studies.

Intraspecific genetic diversity and composition modify species-level diversity-productivity relationships.

Biodiversity regulates ecosystem functions such as productivity, and experimental studies of species mixtures have revealed selection and complementarity effects driving these responses. However, the impacts of intraspecific genotypic diversity in these studies are unknown, despite it forming a substantial part of the biodiversity. In a glasshouse experiment we constructed plant communities with different levels of barley (Hordeum vulgare) genotype and weed species diversity and assessed their relative biodiversity effects through additive partitioning into selection and complementarity effects. Barley genotype diversity had weak positive effects on aboveground biomass through complementarity effects, whereas weed species diversity increased biomass predominantly through selection effects. When combined, increasing genotype diversity of barley tended to dilute the selection effect of weeds. We interpret these different effects of barley genotype and weed species diversity as the consequence of small vs large trait variation associated with intraspecific barley diversity and interspecific weed diversity, respectively. The different effects of intra- vs interspecific diversity highlight the underestimated and overlooked role of genetic diversity for ecosystem functioning.

Improving intercropping: a synthesis of research in agronomy, plant physiology and ecology.

Intercropping 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 species ­ for 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.

The effects of foundation species on community assembly: a global study on alpine cushion plant communities.

Foundation species can change plant community structure by modulating important ecological processes such as community assembly, yet this topic is poorly understood. In alpine systems, cushion plants commonly act as foundation species by ameliorating local conditions. Here, we analyze diversity patterns of species' assembly within cushions and in adjacent surrounding open substrates (83 sites across five continents) calculating floristic dissimilarity between replicate plots, and using linear models to analyze relationships between microhabitats and species diversity. Floristic dissimilarity did not change across biogeographic regions, but was consistently lower in the cushions than in the open microhabitat. Cushion plants appear to enable recruitment of many relatively stress-intolerant species that otherwise would not establish in these communities, yet the niche space constructed by cushion plants supports a more homogeneous composition of species than the niche space beyond the cushion's influence. As a result, cushion plants support higher α-diversity and a larger species pool, but harbor assemblies with lower ß-diversity than open microhabitats. We conclude that habitats with and without dominant foundation species can strongly differ in the processes that drive species recruitment, and thus the relationship between local and regional species diversity.

Facilitative plant interactions and climate simultaneously drive alpine plant diversity.

Interactions among species determine local-scale diversity, but local interactions are thought to have minor effects at larger scales. However, quantitative comparisons of the importance of biotic interactions relative to other drivers are rarely made at larger scales. Using a data set spanning 78 sites and five continents, we assessed the relative importance of biotic interactions and climate in determining plant diversity in alpine ecosystems dominated by nurse-plant cushion species. Climate variables related with water balance showed the highest correlation with richness at the global scale. Strikingly, although the effect of cushion species on diversity was lower than that of climate, its contribution was still substantial. In particular, cushion species enhanced species richness more in systems with inherently impoverished local diversity. Nurse species appear to act as a 'safety net' sustaining diversity under harsh conditions, demonstrating that climate and species interactions should be integrated when predicting future biodiversity effects of climate change.

The context dependence of beneficiary feedback effects on benefactors in plant facilitation.

Facilitative effects of some species on others are a major driver of biodiversity. These positive effects of a benefactor on its beneficiary can result in negative feedback effects of the beneficiary on the benefactor and reduced fitness of the benefactor. However, in contrast to the wealth of studies on facilitative effects in different environments, we know little about whether the feedback effects show predictable patterns of context dependence. We reanalyzed a global data set on alpine cushion plants, previously used to assess their positive effects on biodiversity and the nature of the beneficiary feedback effects, to specifically assess the context dependence of how small- and large-scale drivers alter the feedback effects of cushion-associated (beneficiary) species on their cushion benefactors using structural equation modelling. The effect of beneficiaries on cushions became negative when beneficiary diversity increased and facilitation was more intense. Local-scale biotic and climatic conditions mediated these community-scale processes, having indirect effects on the feedback effect. High-productivity sites demonstrated weaker negative feedback effects of beneficiaries on the benefactor. Our results indicate a limited impact of the beneficiary feedback effects on benefactor cushions, but strong context dependence. This context dependence may help to explain the ecological and evolutionary persistence of this widespread facilitative system.

A global analysis of bidirectional interactions in alpine plant communities shows facilitators experiencing strong reciprocal fitness costs.

Facilitative interactions are defined as positive effects of one species on another, but bidirectional feedbacks may be positive, neutral, or negative. Understanding the bidirectional nature of these interactions is a fundamental prerequisite for the assessment of the potential evolutionary consequences of facilitation. In a global study combining observational and experimental approaches, we quantified the impact of the cover and richness of species associated with alpine cushion plants on reproductive traits of the benefactor cushions. We found a decline in cushion seed production with increasing cover of cushion-associated species, indicating that being a benefactor came at an overall cost. The effect of cushion-associated species was negative for flower density and seed set of cushions, but not for fruit set and seed quality. Richness of cushion-associated species had positive effects on seed density and modulated the effects of their abundance on flower density and fruit set, indicating that the costs and benefits of harboring associated species depend on the composition of the plant assemblage. Our study demonstrates 'parasitic' interactions among plants over a wide range of species and environments in alpine systems, and we consider their implications for the possible selective effects of interactions between benefactor and beneficiary species.

Degeneration of foundation cushion species induced by ecological constraints can cause massive changes in alpine plant communities.

Foundational cushion plants can re-organize community structures and sustain a prominent proportion of alpine biodiversity, but they are sensitive to climate change. The loss of cushion species can have broad consequences for associated biota. The potential plant community changes with the population dynamics of cushion plants remain, however, unclear. Using eight plant communities along a climatic and community successional gradient, we assessed cushion population dynamics, the underlying ecological constraints and hence associated plant community changes in alpine communities dominated by the foundational cushion plant Arenaria polytrichoides. The population dynamics of Arenaria are attributed to ecological constraints at a series of life history stages. Reproductive functions are constrained by increasing associated beneficiary plants; subsequent seedling establishment is constrained by temperature, water and light availability, extreme climate events, and interspecific competition; strong competitive exclusion may accelerate mortality and degeneration of cushion populations. Along with cushion dynamics, species composition, abundance and community structure gradually change. Once cushion plants completely degenerate, previously cushion-dominated communities shift to relatively stable communities that are overwhelmingly dominated by sedges. Climate warming may accelerate the degeneration process of A. polytrichoides. Degeneration of this foundational cushion plant will possibly induce massive changes in alpine plant communities and hence ecosystem functions in alpine ecosystems. The assessment of the population dynamics of foundation species is critical for an effective conservation of alpine biodiversity.

Foundation species influence trait-based community assembly.

Here, we incorporate facilitation into trait-based community assembly theory by testing two mutually compatible facilitative mechanisms: changes in the environmental filter, causing either an increase in the range of trait values (i.e. a range expansion effect) and/or a shift in trait distributions (i.e. a range shift effect); and changes in trait spacing, suggesting an effect on niche differentiation. We analyzed the distribution of three functional traits - leaf dry matter content, specific leaf area and lateral spread - of plant communities dominated by a cushion-forming foundation species at four sites differing in elevation and aspect. We found support for environmental filtering and niche differentiation mechanisms by cushions, with filtering effects (in particular range shifts) increasing with environmental severity at higher elevation. The effect size of cushions on trait distribution was similar to that of environmental gradients caused by elevation and aspect. The consideration of intraspecific trait variability improved the detection of cushion effects on trait distributions. Our results highlight the importance of facilitation in the modification of taxonomic and functional diversity of ecological communities, and indicate that facilitation can occur through combined effects on environmental filtering and niche differentiation, with strong environmental context dependence of each mechanism.

Effect of Drought on Bean Yield Is Mediated by Intraspecific Variation in Crop Mixtures.

Increasing plant diversity in agricultural systems provides promising solutions for sustainably increasing crop yield. It remains unclear; however, how plant-plant interactions in diverse systems are mediated by plant genetic variation. We conducted a greenhouse experiment in which we grew three varieties of common beans with three companion plant species (chickpeas, sorghum, and sunflower) in different combinations (crop mixtures, bean cultivar mixtures, and monocultures), with and without drought stress. We hypothesized that under drought stress, the effect of companion plant species on bean yield would be mediated by the drought tolerance potential of the species. We further hypothesized that this effect would vary across different bean cultivars. Overall, we show that the effect of companion plant species on bean yield was not influenced by drought stress; instead, it was dependent on the identity of the bean variety. This could partially be explained by variation in growth rate between bean varieties, where the fastest growing variety recorded the highest yield increase in plant mixtures. The effect of companion plant species on chickpea biomass, however, was potentially influenced by chickpea drought tolerance potential; chickpea biomass was recorded to be higher in plant mixtures than in its monoculture under drought conditions. Our study highlights that to develop plant mixtures, it is not only important to consider the functional traits of the interacting plant species, but also those of the different plant varieties. We further suggest that stress tolerance can be a useful trait for initial selection of plant varieties when developing crop mixtures.

Rapid transgenerational adaptation in response to intercropping reduces competition.

By capitalising on positive biodiversity-productivity relationships, intercropping provides opportunities to improve agricultural sustainability. Intercropping is generally implemented using commercial seeds that were bred for maximal productivity in monocultures, thereby ignoring the ability of plants to adapt over generations to the surrounding neighbourhood, notably through increased complementarity, that is reduced competition or increased facilitation. This is why using monoculture-adapted seeds for intercropping might limit the benefits of crop diversity on yield. However, the adaptation potential of crops and the corresponding changes in complementarity have not been explored in annual crop systems. Here we show that plant-plant interactions among annual crops shifted towards reduced competition and/or increased facilitation when the plants were growing in the same community type as their parents did in the previous two generations. Total yield did not respond to this common coexistence history, but in fertilized conditions, we observed increased overyielding in mixtures with a common coexistence history. Surprisingly, we observed character convergence between species sharing the same coexistence history for two generations, in monocultures but also in mixtures: the six crop species tested converged towards taller phenotypes with lower leaf dry matter content. This study provides the first empirical evidence for the potential of parental diversity affecting plant-plant interactions, species complementarity and therefore potentially ecosystem functioning of the following generations in annual cropping systems. Although further studies are required to assess the context-dependence of these results, our findings may still have important implications for diversified agriculture as they illustrate the potential of targeted cultivars to increase complementarity of species in intercropping, which could be achieved through specific breeding for mixtures.

Plants have two ways of interacting with each other: they can compete with each other if they use the same resources; or they can 'help' each other in what is known as facilitation, for example, when a larger plant protects a smaller plant in harsh environments. These interactions can vary over several generations in response to changes in the environment or the surrounding plant community. For instance, in plant communities formed by many different species, like in most natural systems, competition usually decreases over time as the plants 'learn' to grow together. In agriculture, intercropping ­ defined as growing at least two species of crop at the same time on the same field ­ takes advantage from a reduction in competition. The idea is that planting two species that grow differently together will lead to less competition than having a single crop because the two species will use slightly different resources, or use them at different times. However, intercropping has traditionally overlooked changes in the interactions between plants as a result of the crop species evolving after being grown together for generations. Indeed, farmers that practice intercropping generally use standard seeds that have been bred to produce high yields when planted on their own, in what is known as monoculture. If plants can adapt and become less competitive when they are grown together over several generations, then using these standard seeds might limit the success of intercropping. Stefan, Engbersen and Schöb wanted to know whether crop species adapt to the levels of plant diversity surrounding them over generations, and if so, how they do it. To find this out, they investigated how competition and facilitation changed when six crop species (wheat, oat, lentil, coriander, flax and camelina) that grow annually were grown together in different combinations over several generations. Stefan, Engbersen and Schöb started off with seeds normally used for growing these crops on their own, and planted them either on their own, or in different combinations of two or four species. They then repeated the experiment over the course of three years, each year using seeds from the previous year, recording both crop yields and changes in how the plants interacted with each other. The experiments showed that interactions among these annual crops shifted towards reduced competition and/or increased facilitation when the plants were growing alongside the same crops as their parents did in the previous two generations. Improving and promoting the development of intercropping is essential for agricultural sustainability, as it could offer alternatives to intensive monocultures (crops grown on their own that require increased resources). Stefan, Engbersen and Schöb's findings are relevant for programmes aimed at developing seeds for intercropping, as they highlight the importance of including diversity when developing these seeds. However, before these results can be used in the field, longer experiments (of more than three generations) in different environments should be carried out to confirm the findings. Another question that remains open is what the mechanisms underlying adaptations to intercropping are: more in-depth research will be needed to determine whether the changes observed have a genetic basis.

Active and adaptive plasticity in a changing climate.

Better understanding of the mechanistic basis of plant plasticity will enhance efforts to breed crops resilient to predicted climate change. However, complexity in plasticity's conceptualisation and measurement may hinder fruitful crossover of concepts between disciplines that would enable such advances. We argue active adaptive plasticity is particularly important in shaping the fitness of wild plants, representing the first line of a plant's defence to environmental change. Here, we define how this concept may be applied to crop breeding, suggest appropriate approaches to measure it in crops, and propose a refocussing on active adaptive plasticity to enhance crop resilience. We also discuss how the same concept may have wider utility, such as in ex situ plant conservation and reintroductions.