Spatial Variation in Throughfall, Soil, and Plant Water Isotopes in a Temperate Forest

Studies of stable isotopes of water in the environment have been fundamental to advancing our understanding of how water moves through the soil‐plant‐atmosphere continuum; however, much of this research focuses on how water isotopes vary in time, rather than in space. We examined the spatial variation in the δ18O and δ2H of throughfall and bulk soil water, as well as branch xylem and bulk leaf water of Picea abies (Norway Spruce) and Fagus sylvatica (Beech), in a 1 ha forest plot in the northern Alps of Switzerland. Means and ranges of water isotope ratios varied considerably among throughfall, soil, and xylem samples. Soil water isotope ratios were often poorly explained by soil characteristics and often not predictable from proximal samples. Branch xylem water isotope values varied less than either soil water or bulk leaf water. The isotopic range observed within an individual tree crown was often similar to that observed among different crowns. As a result of the heterogeneity in isotope ratios, inferences about the depth of plant root water uptake drawn from a two end‐member mixing model were highly sensitive to the soil sampling location. Our results clearly demonstrate that studies using water isotopes to infer root water uptake must explicitly consider how to characterize soil water, incorporating measures of both vertical and lateral variation. By accounting for this spatial variation and the processes that shape it, we can improve the application of water isotopes to studies of plant ecophysiology, ecohydrology, soil hydrology, and paleoclimatology

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 pheno-types 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 there-fore 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.ISSN:2050-084

Using spatially-explicit plant competition models to optimise crop productivity in intercropped systems

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.ISSN:1439-1791ISSN:1618-008

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.ISSN:1051-0761ISSN:1939-558

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.ISSN:1051-0761ISSN:1939-558

Ecological and evolutionary effects of crop diversity decrease yield variability

1. Higher plant species diversity decreases variability of plant community productivity. The stabilizing effect of plant diversity can result from species-specific responses to environmental fluctuations and from shifts in competitive hierarchies. Evolutionary adaptation of species to surrounding plant diversity could further decrease productivity variability. 2. We used a three-year dataset from a crop diversity experiment with seven species to assess the effect of crop diversity and selection history on temporal variability of yield. 3. We found contrasting patterns of temporal variability: Yield of species varied more in mixtures than in monocultures over years. However, at community-level, we found lower yield variability in crop mixtures compared to monocultures, although only in combination with fertilizer application under Mediterranean climate. Furthermore, we found that a mixture selection history can increase yield productivity and decrease its variability, although only in monocultures. This suggests that the interspecific interactions among crops in mixtures act as an evolutionary selective force, promoting niche complementarity. 4. Synthesis. Our results highlight the ecological and evolutionary role of plant interactions in crop mixtures, which can affect yield stability, while also reflecting on the importance of climate and resource availability in modifying the diversity-stability relationship.ISSN:0022-047

Temporal dynamics of biodiversity effects and light-use-related traits in two intercropping systems

Introduction: Intercropping systems can be more productive than their respective monocultures and this positive net biodiversity effect is caused by complementarity and selection effects. While the complementarity effect is caused through resource partitioning or facilitation, the selection effect operates via the greater probability that a more diverse community contains a dominant and high-yielding species which will account for the majority of productivity in that community. Here, we investigated how light-use-related traits contribute to the net biodiversity effect via complementarity or selection effects and how these qrelationships change throughout an annual growing season. Materials and Methods: We conducted weekly destructive harvests to examine temporal dynamics of biodiversity effects in two crop mixtures (oat–lupin and oat–camelina) and their respective monocultures. We linked the biodiversity effects to traits related to light use (i.e., light interception, plant height, photosynthetic efficiency and photosynthetic capacity) and investigated how these relationships changed over time. Results: We found that the net biodiversity and selection effect increased over time in both mixtures, while complementarity effects increased only in the oat–lupin mixture. More intercepted light and taller plants in mixtures compared to monocultures positively contributed to biodiversity effects in both mixtures. Strategies for shade tolerance differed between the mixtures, that is, increased photosynthetic capacity and increased photosynthetic efficiency contributed to a positive net biodiversity effect in the oat–lupin and oat–camelina mixture, respectively. Conclusion: By linking the temporal dynamics of the net biodiversity effect and its two additive components to light-use-related traits in two different crop mixtures, this study demonstrates that complementary light use contributes to overyielding in intercropping systems. Such understanding is important for the design of effective intercropping systems and developing new crop cultivars suited to these environments.ISSN:2767-035

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

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.ISSN:1664-462