Search

Interactions among species are a fundamental aspect of biodiversity and drive ecosystem functioning and services. Species interactions include direct (pairwise) interactions among two species and indirect interactions that occur when a third species interacts and changes the pairwise direct interaction. In a three‐species interaction network, these interactions can be transitive (where one species outperforms all others) or intransitive (where each species outperforms another). Here, we investigate how direct and indirect interactions influence ecosystem functions in crop systems and how diversification and evolutionary adaptation can influence those interactions and therefore ecosystem functions.A common garden experiment was conducted with crop communities in monocultures, 2‐ and 3‐species mixtures that had either a common or no coexistence history (i.e. co‐adaptation) for the three previous years. Net, direct and indirect interaction intensities were estimated and compared between the diversity levels and coexistence histories. Furthermore, species interaction networks were inspected for transitive/intransitive interactions.We found evidence for less intense competition in mixtures and for reduced negative direct interaction intensity and enhanced facilitative effects upon co‐adaptation. We could further show that indirect interactions were generally less important for co‐adaptation than direct interactions. Additionally, we showed that co‐adaptation has the potential to shift interactions in the species interaction networks from competitive intransitive into pairwise competitive interactions where interactions occurred mainly between two species.Synthesis. Co‐adapted crop species with reduced negative interactions might have the potential to enhance productivity, especially in more diverse cropping systems. This supports the notion that intercropping is a vital part towards a more sustainable agriculture and one with further yield potential when developing cultivars optimised for growth in mixtures. [ABSTRACT FROM AUTHOR]

Inspired by grassland biodiversity experiments studying the impact of plant diversity on primary productivity, the Crop Diversity Experiment setup in 2018 aimed at testing whether these biodiversity benefits also hold for annual crop systems and whether crop mixtures also achieved transgressive overyielding, i.e. yield in mixture that was higher than the most productive monoculture. The first 3 years of the experiment demonstrated that crop mixtures do not only increase yield compared with an average monoculture but often also compared with the highest yielding monoculture. The crop diversity effects were stronger under more stressful environmental conditions and were often achieved in mixtures with legume crops. However, we observed transgressive overyielding also under favorable conditions and in mixtures without legumes. With our investigation of the underlying mechanisms of the yield benefits we found both direct complementarities between crop species and indirect effects via other organisms. The former included chemical, spatial and temporal complementarity in N uptake, complementary root distribution leading to complementary water uptake, as well as spatial and temporal complementarity in light use. Among the indirect mechanisms we identified complementary suppression of weeds and more abundant plant growth-promoting microbes in crop mixtures, apart from complementarity in pest and disease suppression not yet studied in the Crop Diversity Experiment but demonstrated elsewhere. In consequence, the Crop Diversity Experiment supports not only the assumption that the ecological processes identified in biodiversity experiments also hold in crop systems, but that diversification of arable crop systems provides a valuable tool to sustainably produce food. [ABSTRACT FROM AUTHOR]

More diverse plant communities are generally more productive than monocultures. This benefit of species diversity is supposed to stem from resource partitioning of species in mixtures where different species use the resources spatially, temporally, or chemically in distinct ways. With respect to water, the simultaneous cultivation of crops with distinct water uptake patterns might reduce niche overlaps and thus result in higher productivity. However, little is known about whether and how spatial water uptake patterns of crop species differ among different planting arrangements and whether these changes result in increased niche partitioning and explain overyielding in mixtures.Stable isotopes of water and a Bayesian model were used to investigate the spatial water uptake patterns of six different crop species and how these patterns change depending on the planting arrangement (monocultures vs mixtures). Niche overlaps and niche widths in spatial water uptake were compared among the different crop diversity levels and linked to productivity. Furthermore, spatial water uptake was related to competition intensity and overyielding in mixtures.We found evidence for increased niche partitioning in spatial water uptake, and therefore complementary spatial root distributions of crop species, and higher expected productivity in mixtures compared to expected productivity in monocultures both due to inherent species‐level differences in water uptake and plasticity in the water uptake pattern of species. We also found a significant relationship of competition and overyielding with observed patterns in spatial water uptake. These results suggest that competition was most intense in shallow soil layers and enhanced overyielding was related to a gradual increase of water uptake in deeper soil layers. Thus, overyielding might be related to a more complete spatial exploitation of available water sources.Synthesis. Differences in spatial water uptake and niche partitioning of intercropped species, driven most likely by a complementary spatial root distribution, might explain why mixtures outperform monocultures. These findings underpin the potential of intercropping systems for a more sustainable agriculture with a more efficient use of soil resources and hence reduced input demands. [ABSTRACT FROM AUTHOR]

Intercropping, by capitalizing on positive biodiversity–productivity relationships, represents a promising option to increase agricultural sustainability. However, the complexity and context-dependency of plant–plant interactions can make it challenging for farmers to find suitable crop combinations. Furthermore, intercropping is usually implemented with standard inter-row spacing and plant densities based on monoculture practices, which might not be the ideal configuration to maximize yield. Here we present a spatially-explicit yield analysis method based on plant ecological interaction models that allowed to optimize crop species combinations and spatial configurations for maximal yield in intercropped systems. We tested this method with three crop species, namely oat, lupine, and camelina. In a first step, field experiments in which crop density and adjacent crop type were varied provided us with indications on which species would compete more with each other. The results showed us that oat and camelina strongly competed with each other. In addition, the distance experiments allowed us to understand how the changes in yield associated with the presence of neighbors vary with distance. This allowed us to find the sets of parameters (identity of neighbors, sowing density, distances between individuals) that optimise intercrop yield (measured as Land Equivalent Ratio [LER]) for the three considered species. Specifically, we show that alternating rows of species led to higher LERs than a homogeneous species mixing, and that 3-species combinations are not necessarily more performant than the best 2-species combinations. In addition, we show that increasing the density of oat is generally beneficial for LER, while increasing the density of lupine is not. By modelling crop yield from simple and reproducible density and distance experiments, our results allow to optimize crop mixtures in terms of species combinations and spatial configurations, for maximal crop yield. [ABSTRACT FROM AUTHOR]

Recent meta-analyses demonstrate that intercropping can increase the land use efficiency of crop production by 20–30 % on average, indicating a strong potential contribution to sustainable intensification. However, there is substantial variability around this average: individual studies range from half to double the land productivity of monocrops. Legume-cereal intercrops and intercrops with high temporal niche separation tend to be more productive than the average, but these two combination types are not always suitable. There is a need to explore other possibilities to achieve productive intercrops. We explored whether two simple functional traits involved in radiation use, plant vegetative height and specific leaf area (SLA), could be used to predict intercrop productivity. Height and SLA together are associated with key plant life-history and resource economy strategies determining competitiveness and tolerance of competition, especially with regard to light, and could therefore be expected to underpin overyielding in intercrops. In the first year of our study, we grew crops as monocrops at one site in Kenya and measured their height and SLA. In the second year, we grew crops in monocrop, intercrop, and single plant treatments at two sites in Kenya and one site in Nigeria. Together, these treatments allowed us to identify whether each intercrop combination overyielded or underyielded, and whether any overyielding was driven by facilitation and/or differences in inter- vs intraspecific competition. We then related the strength of these interactions to the two traits. We found that intercrop grain yields varied in relation to the height and SLA of each species in the intercrop, but together these traits explained less than a third of variation in intercrop land equivalence ratios (LER). More variation could be explained by allowing for the effect of site, suggesting that the two traits interact with site conditions to determine yield. Biomass LERs responded differently to grain LERs, suggesting that plasticity in resource allocation in response to intercropping conditions may further influence yields. Our study found some evidence that combining species with traits indicating contrasting responses to competition (an avoidant species with a tolerant species) could increase resource use complementarity and thus intercrop overyielding. However, it was clear that other factors (such as additional traits, or the trait by site interaction) are needed to refine our understanding of intercrop productivity. A trait-based framework has potential to predict intercrop productivity, but simple measures of height and SLA alone are insufficient. • Intercrop yields vary from much lower than sole crops to much higher across different studies. • Plant traits may mediate competition and facilitation between intercrops, and predict yields. • Two simple traits, vegetative height and specific leaf area (SLA), explained some variance in yields. • Yields were higher when tall species with a low SLA were paired with short species with a high SLA. • Height and SLA alone cannot reliably predict intercrop yields: other traits should be explored. [ABSTRACT FROM AUTHOR]