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.

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.

Temporal Differentiation of Resource Capture and Biomass Accumulation as a Driver of Yield Increase in Intercropping.

Intercropping, i.e., the simultaneous cultivation of different crops on the same field, has demonstrated yield advantages compared to monoculture cropping. These yield advantages have often been attributed to complementary resource use, but few studies quantified the temporal complementarity of nutrient acquisition and biomass production. Our understanding of how nutrient uptake rates of nitrogen (N) and phosphorous (P) and biomass accumulation change throughout the growing season and between different neighbors is limited. We conducted weekly destructive harvests to measure temporal trajectories of N and P uptake and biomass production in three crop species (oat, lupin, and camelina) growing either as isolated single plants, in monocultures or as intercrops. Additionally, we quantified organic acid exudation in the rhizosphere and biological N2-fixation of lupin throughout the growing season. Logistic models were fitted to characterize nutrient acquisition and biomass accumulation trajectories. Nutrient uptake and biomass accumulation trajectories were curtailed by competitive interactions, resulting in earlier peak rates and lower total accumulated nutrients and biomass compared to cultivation as isolated single plants. Different pathways led to overyielding in the two mixtures. The oat-camelina mixture was characterized by a shift from belowground temporal niche partitioning of resource uptake to aboveground competition for light during the growing season. The oat-lupin mixture showed strong competitive interactions, where lupin eventually overyielded due to reliance on atmospheric N and stronger competitiveness for soil P compared to oat. Synthesis: This study demonstrates temporal shifts to earlier peak rates of plants growing with neighbors compared to those growing alone, with changes in uptake patterns suggesting that observed temporal shifts in our experiment were driven by competitive interactions rather than active plant behavior to reduce competition. The two differing pathways to overyielding in the two mixtures highlight the importance of examining temporal dynamics in intercropping systems to understand the underlying mechanisms of overyielding.

Positive Effects of Crop Diversity on Productivity Driven by Changes in Soil Microbial Composition.

Intensive agriculture has major negative impacts on ecosystem diversity and functioning, including that of soils. The associated reduction of soil biodiversity and essential soil functions, such as nutrient cycling, can restrict plant growth and crop yield. By increasing plant diversity in agricultural systems, intercropping could be a promising way to foster soil microbial diversity and functioning. However, plant-microbe interactions and the extent to which they influence crop yield under field conditions are still poorly understood. In this study, we performed an extensive intercropping experiment using eight crop species and 40 different crop mixtures to investigate how crop diversity affects soil microbial diversity and activity, and whether these changes subsequently affect crop yield. Experiments were carried out in mesocosms under natural conditions in Switzerland and in Spain, two countries with drastically different soils and climate, and our crop communities included either one, two or four species. We sampled and sequenced soil microbial DNA to assess soil microbial diversity, and measured soil basal respiration as a proxy for soil activity. Results indicate that in Switzerland, increasing crop diversity led to shifts in soil microbial community composition, and in particular to an increase of several plant-growth promoting microbes, such as members of the bacterial phylum Actinobacteria. These shifts in community composition subsequently led to a 15 and 35% increase in crop yield in 2 and 4-species mixtures, respectively. This suggests that the positive effects of crop diversity on crop productivity can partially be explained by changes in soil microbial composition. However, the effects of crop diversity on soil microbes were relatively small compared to the effects of abiotic factors such as fertilization (three times larger) or soil moisture (three times larger). Furthermore, these processes were context-dependent: in Spain, where resources were limited, soil microbial communities did not respond to crop diversity, and their effect on crop yield was less strong. This research highlights the potential beneficial role of soil microbial communities in intercropping systems, while also reflecting on the relative importance of crop diversity compared to abiotic drivers of microbiomes and emphasizing the context-dependence of crop-microbe relationships.

Diversity increases yield but reduces harvest index in crop mixtures.

Resource allocation to reproduction is a critical trait for plant fitness1,2. This trait, called harvest index in the agricultural context3-5, determines how plant biomass is converted to seed yield and consequently financial revenue from numerous major staple crops. While plant diversity has been demonstrated to increase plant biomass6-8, plant diversity effects on seed yield of crops are ambiguous9 and dependent on the production syndrome10. This discrepancy might be explained through changes in the proportion of resources invested in reproduction in response to changes in plant diversity, namely through changes in species interactions and microenvironmental conditions11-14. Here, we show that increasing crop plant diversity from monocultures over two- to four-species mixtures increased annual primary productivity, resulting in overall higher plant biomass and, to a lesser extent, higher seed yield in mixtures compared with monocultures. The difference between the two responses to diversity was due to a reduced harvest index of the eight tested crop species in mixtures, possibly because their common cultivars have been bred for maximum performance in monoculture. While crop diversification provides a sustainable measure of agricultural intensification15, the use of currently available cultivars may compromise larger gains in seed yield. We therefore advocate regional breeding programmes for crop varieties to be used in mixtures that should exploit complementarity16 among crop species.

Cadmium accumulation and allocation in different cacao cultivars.

Cadmium (Cd) is a biologically non-essential heavy metal that can cause toxic effects in plants, animals and humans already at low concentrations compared to other metals. After Cd concentrations in cacao beans of various provenances, particularly from Latin America, were found to exceed the new regulations enforced by the European Union in 2019, there is an urgent need to find measures to lower Cd accumulation in cacao beans to acceptable values. In this research, the long-term cacao cultivar trial CEDEC-JAS in northern Honduras was used to investigate differences between 11 cultivars in Cd uptake and translocation. Sampling of various plant parts, including rootstocks, scions, leaves and beans, from three replicate trees per cultivar and the soil around each tree was conducted at this site. Results indicate that concentrations of available soil Cd were more closely correlated with Cd concentrations of the rootstocks (R2 = 0.56), scions (R2 = 0.59) and leaves (R2 = 0.46) than with bean Cd concentrations (R2 = 0.26). In addition, Cd concentrations of rootstocks, scions and leaves showed close relationships to available soil Cd concentrations, with no significant differences between the cultivars. In contrast, bean Cd concentrations showed only weak correlations to available soil Cd and Cd concentrations in the vegetative plant parts, but significant variation among cultivars. Three cultivars, which were analysed in more detail, showed significant differences in Cd concentrations of mature beans, but not of immature beans. These results suggest that cultivar-related differences in bean Cd concentrations primarily result from differences in Cd loading during bean maturation, possibly due to cultivar-specific differences in the xylem-to-phloem transfer of Cd. The results show that selection of cultivars with low Cd transfer from vegetative parts into the beans has high potential to keep Cd accumulation in cacao beans at levels that are safe for consumption.