Transgressive Overyielding Research Articles

Coordinating lodging incidence and grain yield through wheat genetic diversity

ContextLodging is a complex phenomenon that affects wheat grain yield worldwide. There is a contradiction between lodging incidence and continuous increase in grain yield in a single-cultivar pattern. However, few studies have explored the coordination between lodging incidence and grain yield with an increase in genetic diversity. ObjectiveThis study tested the hypothesis that positive intra-population interactions among wheat cultivars in mixtures may improve lodging resistance and grain yield. MethodsA field experiment was conducted on the North China Plain, wherein wheat cultivars were grown in a single-planting pattern and cultivar mixtures comprising two (mixture-2), four (mixture-4) and eight (mixture-8) wheat cultivars, to explore the interaction between lodging resistance and grain yield. ResultsLodging was not observed in all the cultivar mixtures; however, some lodging-susceptible cultivars underwent a partial reduction in grain yield owing to lodging in single-cultivar patterns. Lodging stress resistance was significantly (P< 0.01) and positively correlated with the breaking strength of the second basal internode at the dough stage. The stem morphological characteristics and lignin and cellulose contents of the second internode contributed to the enhanced breaking strength of the cultivar mixtures. The cultivar mixtures showed increased grain yield compared with the weighted mean of the single-component cultivars. A transgressive over-yielding effect of the wheat cultivar mixtures was observed in 2019 and 2021. ConclusionsOur findings demonstrate that the cultivar mixtures exhibited improved lodging resistance and grain yield at the population level, whereas partial yield loss caused by lodging was evident in single-cultivar patterns. The coordinating effect can be attributed to the higher proportion of lodging-resistant cultivars and the greater mechanical strength of the stem in the cultivar mixtures. SignificanceThe findings imply that wheat cultivar mixture, as an ecological approach, is an effective strategy for reducing yield loss caused by lodging in field production.

Optimization of N Fertilizer and Synergistic Intercropping to Enhance the Productivity Advantage of Faba Bean and Effective Control of Chocolate Spot Epidemics.

Intercropping of wheat/faba bean is a common practice within the legume-cereal family. However, the benefits of nitrogen (N) fertilizer-optimized synergistic intercropping in improving faba bean productivity while controlling the prevalence of chocolate spot disease have not been established. This study conducted continuous field experiments spanning two planting seasons to investigate two key findings: (i) Optimizing N fertilizer application can enhance the productivity of intercropped faba bean. (ii) The percentage severity index (PSI) during the period of maximum prevalence rate (Rmax) of faba bean chocolate spot disease poses a substantial challenge to faba bean yield. The results indicated that the land equivalent ratio and transgressive overyielding index for each intercropping treatment increased with higher N fertilizer application, exceeding a value of 1, and the land saving proportion also exceeded 0. Intercropping primarily enhances productivity, as measured by the harvest index (HI), by amplifying the complementary effect rather than the selection effect, thus improving the net benefit of intercropping. The HI of single and intercropped faba bean increased with the N1 and N2 treatments in both planting seasons. However, the HI of single and intercropped faba bean at the N3 level decreased significantly, ranging from 17.85 to 29.62%. Furthermore, a notable negative correlation was established between the PSI during critical epidemic (initial epidemic, maximum epidemic rate, and late epidemic) periods and observed and expected faba bean yields. As the PSI increased, faba bean yields decreased, and the PSI of intercropping at different periods was lower than that observed in the single cropping. Additionally, intercropping with the optimized N fertilizer treatment (N2 treatment) exhibited an enhanced relative control effect on chocolate spot disease in faba bean, ranging from 35.21 to 52.36%. This finding confirmed the productivity advantage of intercropping faba bean. In conclusion, this study suggested that optimizing N fertilizer application can enhance the productivity of intercropped faba bean. Wheat/faba bean intercropping effectively controlled the PSI during the period of Rmax, which would otherwise threaten faba bean yield. Consequently, this practice ensured sustained advantages of wheat/faba bean intercropping.

Microbe-induced phenotypic variation leads to overyielding in clonal plant populations.

Overyielding, the high productivity of multispecies plant communities, is commonly seen as the result of plant genetic diversity. Here we demonstrate that biodiversity-ecosystem functioning relationships can emerge in clonal plant populations through interaction with microorganisms. Using a model clonal plant species, we found that exposure to volatiles of certain microorganisms led to divergent plant phenotypes. Assembling communities out of plants associated with different microorganisms led to transgressive overyielding in both biomass and seed yield. Our results highlight the importance of belowground microbial diversity in plant biodiversity research and open new avenues for precision ecosystem management.

Annual yields of multispecies grassland mesocosms outperformed monocultures across a drought gradient due to complementarity effects and rapid recovery

AbstractBackgroundMore frequent and severe drought events due to climate change pose a major challenge for sustainable forage production in managed grasslands. This study investigated whether multispecies grassland communities can provide greater resistance to and/or recovery from drought compared to monoculture communities.MethodsMesocosms of Lolium perenne L., Cichorium intybus L., Trifolium repens L. and Trifolium pratense L. were established as monocultures, and a four‐species mixture. A drought gradient with five levels of water supply ranging from a mild to a severe treatment was applied for 10 weeks, in each of 2 years. Shoot biomass was harvested to assess drought resistance, drought recovery and annual yields. Root mass density and specific root length were measured in Year 2.ResultsAcross the drought gradient, four‐species communities had significantly larger annual yields than each of the four monocultures, indicating transgressive overyielding. This was despite relatively low drought resistance for four‐species communities compared with L. perenne and C. intybus monocultures. Recovery of yields following drought was high for all communities.ConclusionsMultispecies swards with complementary traits can provide a viable adaptation option across a wide range of drought severities. Application of a stress gradient methodology allowed a more detailed understanding of stress responses.

Positive effects of plant diversity on dry matter yield while maintaining a high level of forage digestibility in intensively managed grasslands across two contrasting environments

AbstractThe local abiotic and environmental conditions of a grass‐based farming system may influence the agricultural benefits of mixtures in comparison to pure stands. We investigated the effects of species identities and interactions between grass, legume, and herb species on dry matter yield and sward digestibility and explored how contrasting environments may affect these relationships. We established experimental plots across 11 mixtures and 4 monocultures of perennial ryegrass (Lolium perenne L.), chicory (Cichorium intybus L.), white clover (Trifolium repens L.), and red clover (Trifolium pratense L.) established at two seeding densities in two sites Brody and Szelejewo (Poland) and managed them over a 3‐year period. The two sites were close geographically and had similar climates, but differed in soil types (Luvisols and Cambisols, respectively). We confirmed that the annual DM yields were generally higher on Cambisols soil as compared to Luvisols soil; both individual species effects and species interaction strengths differed across the two contrasting environments. The predicted annual DM yield for the 4‐species mixtures in general gave comparable or higher yields compared to monocultures across the 3 years, with the mixtures giving around 10%–30% higher yields than the weighted average of the monocultures of the constituent species. Analysing the transgressive overyielding it turned out that there was no significant difference between the 4‐species mixtures and best performing monoculture. The benefits of species diversity in our study was greater on less productive site because we found larger overyielding of the 4‐species sward on the Luvisols soil than on the Cambisols soil. Additionally, the effect of mixtures composed by 4‐species increased the yield stability compared to monocultures in the 3‐year period of our study, particularly on Cambisols soil. Sward digestibility applied to average values did not differ much between sites with mixtures performing similarly to monocultures. The reason for that could have been the dynamics of sward botanical composition during study years shifted towards increased perennial ryegrass and decreased proportions of chicory in the sward as well as the observed phenomenon that the species in mixed swards progressed to successive growth stages more slowly than in pure sowing.

Yield of lucerne-grass mixtures did not differ from lucerne pure stands in a multi-site field experiment

Changing climatic conditions in north-western Europe require adaptations in management of ley grassland, requiring new grass-legume mixture designs in response to increased variability in precipitation and warmer summers. Evaluation of lucerne (Medicago sativa L.)-grass swards with the grasses cocksfoot (Dactylis glomerata L.) and tall fescue (Festuca arundinacea Schreb.), which are potentially more tolerant to summer water shortage, have yet to be evaluated under the agro-climatic conditions of north-western Europe. We tested these grasses in three-species swards and binary lucerne grass swards and compared them against pure stands and a standard lucerne-grass mixture. The study was conducted in eight environments resulting from the combination of four soil sites during the two full main production years 2020 and 2021 with an annual precipitation range of 416–839 mm. Swards were harvested three to five times annually and the accumulated herbage dry matter (DM) and nitrogen (N) yields, as well as species proportions were recorded. We found an average yield of 11.1 t DM ha-1 and no significant differences in herbage DM yields between lucerne pure stands and the different mixture swards. This pattern was stable irrespective of environmental conditions, but in general herbage yield and aboveground N yield were affected by precipitation amount. Swards with cocksfoot had a greater grass proportion than those with tall fescue. The N yield of lucerne-grass swards was lower and declined as the grass proportion increased, and reached on average 353 kg N ha-1. The maximum N yield of lucerne pure stands was 547 kg N ha-1. Species mixtures yielded higher than expected from the pure stands (net mixture effect, NE) pointing at overyielding. Consequently, synergistic interspecific interactions occurred but did not lead to transgressive overyielding. The results indicate that the inclusion of grass species with tolerance to water shortage into lucerne-based swards offers the potential to diversify grasslands without losses of agronomic potential, at least during the first two main production years.

Crop Diversity Experiment: towards a mechanistic understanding of the benefits of species diversity in annual crop systems

Abstract 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.

Scaling‐up the biodiversity–ecosystem functioning relationship: the effect of environmental heterogeneity on transgressive overyielding

Knowledge of how biodiversity sustains ecosystem function comes predominantly from studies focused on small spatial scales. Thus, we know relatively little about the role of biodiversity at larger scales of space and time where habitats become increasingly heterogeneous. Efforts to upscale the relationship between biodiversity and function have yielded inconclusive results. Given that increasing habitat heterogeneity is a ubiquitous consequence of increasing spatial scale, we asked: as habitat heterogeneity increases, can single species continue to maintain ecosystem function? Or, does transgressive overyielding (functioning of species mixture divided by the functioning of the highest functioning single species) change with habitat heterogeneity? We addressed this using a combination of computer simulations, an experiment and a meta‐analysis. The three parts followed the same rationale: habitat heterogeneity was increased by aggregating local habitats with different conditions into larger and more heterogeneous landscapes. The computer simulations showed that, on average, transgressive overyielding increased with habitat heterogeneity because monoculture functioning decreased with habitat heterogeneity. We tested this expectation experimentally by varying the strain richness from one to five species across 10800 bacterial communities in five different habitats defined by sub‐inhibitory concentrations of antibiotics. On average, the experimental results concurred with the simulations. We tested the generality of this result using a meta‐analysis of 26 published experiments that manipulated habitat conditions and species richness. This confirmed that transgressive overyielding tended to increase with habitat heterogeneity but only when species were specialised to different habitats and were not inhibited in mixtures by negative species interactions. This was not the case in several experiments used in our meta‐analysis where one species maximised functioning across all habitats, contrary to the assumptions of many ecological models. Our results illustrate the importance of biodiversity at larger spatial scales with more heterogeneity but also highlights contingencies that this pattern depends on.

The productive performance of intercropping

Crop diversification has been put forward as a way to reduce the environmental impact of agriculture without penalizing its productivity. In this context, intercropping, the planned combination of two or more crop species in one field, is a promising practice. On an average, intercropping saves land compared with the component sole crops, but it remains unclear whether intercropping produces a higher yield than the most productive single crop per unit area, i.e., whether intercropping achieves transgressive overyielding. Here, we quantified the performance of intercropping for the production of grain, calories, and protein in a global meta-analysis of several production indices. The results show that intercrops outperform sole crops when the objective is to achieve a diversity of crop products on a given land area. However, when intercropping is evaluated for its ability to produce raw products without concern for diversity, intercrops on average generate a small loss in grain or calorie yield compared with the most productive sole crop (-4%) but achieve similar or higher protein yield, especially with maize/legume combinations grown at moderate N supply. Overall, although intercropping does not achieve transgressive overyielding on average, our results show that intercropping performs well in producing a diverse set of crop products and performs almost similar to the most productive component sole crop to produce raw products, while improving crop resilience, enhancing ecosystem services, and improving nutrient use efficiency. Our study, therefore, confirms the great interest of intercropping for the development of a more sustainable agricultural production, supporting diversified diets.

Comparison of transgressive overyielding effect and plant diversity effects of annual and perennial legume-grass mixtures

Comparison of transgressive overyielding effect and plant diversity effects of annual and perennial legume-grass mixtures

Identification of species traits enhancing yield in wheat-faba bean intercropping: development and sensitivity analysis of a minimalist mixture model

AimCereal-legume intercropping can result in yield gains compared to monocrops. We aim to identify the combination of crop traits and management practices that confer a yield advantage in strip intercropping.MethodsWe developed a novel, parameter-sparse process-based crop growth model (Minimalist Mixture Model, M3) that can simulate strip intercrops under well-watered but nitrogen limited growth conditions. It was calibrated and validated for spring wheat (Triticum aestivum) and spring faba bean (Vicia faba) grown as monocrops and intercrops, and used to identify the most suitable trait combinations in these intercrops via sensitivity analyses.ResultsThe land equivalent ratio of intercrops was greater than one over a wide range of nitrogen fertilizer levels, but transgressive overyielding, with total yield in the intercrop greater than that of either sole crop, was only obtained at intermediate nitrogen applications. We ranked the local sensitivities of the individual yields of wheat and faba bean of the whole intercrop under various nitrogen input levels to various crop traits.ConclusionsThe total intercrop yield can be improved by selecting specific traits related to phenology of both species, as well as light use efficiency of faba bean and, under high nitrogen applications, of wheat. Changes in height-related crop traits affected individual yields of species in intercrops but not the total intercrop yield.

Using forest gap models and experimental data to explore long-term effects of tree diversity on the productivity of mixed planted forests

Key messageIn this exploratory study, we show how combining the strength of tree diversity experiment with the long-term perspective offered by forest gap models allows testing the mixture yielding behavior across a full rotation period. Our results on a SW France example illustrate how mixing maritime pine with birch may produce an overyielding (i.e., a positive net biodiversity effect).ContextUnderstanding the link between tree diversity and stand productivity is a key issue at a time when new forest management methods are investigated to improve carbon sequestration and climate change mitigation. Well-controlled tree diversity experiments have been set up over the last decades, but they are still too young to yield relevant results from a long-term perspective. Alternatively, forest gap models appear as appropriate tools to study the link between diversity and productivity as they can simulate mixed forest growth over an entire forestry cycle.AimsWe aimed at testing whether a forest gap model could first reproduce the results from a tree diversity experiment, using its plantation design as input, and then predict the species mixing effect on productivity and biomass in the long term.MethodsHere, we used data from different forest experimental networks to calibrate the gap model ForCEEPS for young pine (Pinus pinaster) and birch (Betula pendula) stands. Then, we used the refined model to compare the productivity of pure and mixed pine and birch stands over a 50-year cycle. The mixing effect was tested for two plantation designs, i.e., species substitution and species addition, and at two tree densities.ResultsRegarding the comparison with the experiment ORPHEE (thus on the short term), the model well reproduced the species interactions observed in the mixed stands. Simulations showed an overyielding (i.e., a positive net biodiversity effect) in pine-birch mixtures in all cases and during the full rotation period. A transgressive overyielding was detected in mixtures resulting from birch addition to pine stands at low density. These results were mainly due to a positive mixing effect on pine growth being larger than the negative effect on birch growth.ConclusionAlthough this study remains explorative, calibrating gap models with data from monospecific stands and validating with data from the manipulative tree diversity experiment (ORPHEE) offers a powerful tool for further investigation of the productivity of forest mixtures. Improving our understanding of how abiotic and biotic factors, including diversity, influence the functioning of forest ecosystems should help to reconsider new forest managements optimizing ecosystem services.

What influences the long-term development of mixtures in British forests?

Abstract Six experiments were established between 1955 and 1962 in different parts of northern and western Britain which used replicated randomized block designs to compare the performance of two species 50:50 mixtures with pure stands of the component species. The species involved were variously lodgepole pine (Pinus contorta Dougl.), Japanese larch (Larix kaempferi Lamb. Carr.), Scots pine (Pinus sylvestris L.), silver birch (Betula pendula Roth.), Sitka spruce (Picea sitchensis Bong. Carr.) and western hemlock (Tsuga heterophylla Raf. Sarg.). The first four species are light demanding, while Sitka spruce is of intermediate shade tolerance and western hemlock is very shade tolerant: only Scots pine and silver birch are native to Great Britain. In three experiments (Bickley, Ceannacroc, Hambleton), the mixtures were of two light-demanding species, while at the other three sites, the mixture tested contained species of different shade tolerance. The experiments were followed for around 50 years, similar to a full rotation of even-aged conifer stands in Britain. Five experiments showed a tendency for one species to dominate in mixture, possibly reflecting differences in the shade tolerance or other functional traits of the component species. In the three experiments, the basal area of the mixtures at the last assessment was significantly higher than predicted based on the performance of the pure stands (i.e. the mixture ‘overyielded’). In two of these cases, the mixture had had a higher basal area than found in the more productive pure stand indicating ‘transgressive overyielding’. Significant basal area differences were generally more evident at the later assessment date. The exception was in a Scots pine: western hemlock mixture where greater overyielding at the earlier date indicated a nursing (‘facilitation’) effect. In the remaining experiments, the performance of the mixture conformed to predictions from the growth of the component species in pure stands. Taken overall, the results suggest that functional traits can be used to interpret the performance of mixtures but prediction of the outcome will require better understanding of the interplay between species and site characteristics plus the influence of silvicultural interventions.

Mixing effect on stand yield of Pinus tabulaeformis and Quercus liaotungensis is modulated by site quality and stand density in the Loess Plateau, China

ABSTRACT Interactions between plants are complicated: they can change resource availability, the proportion of resources acquired and the efficiency of resource use. This can eventually lead to changes in stand yield, but the driving mechanisms remain controversial. This study aimed to clarify the effects of mixing on stand yield. We established 120 mixed and pure plots of Chinese pine (Pinus tabulaeformis) and Liaodong oak (Quercus liaotungensis) in the Loess Plateau, China. Based on the inventory data in 2009 and 2016, we compared the differences in yields between mixed and pure stands. Our results indicated that the mixing of pine and oak resulted in over-yielding, but there was no transgressive over-yielding and pure pine stands consistently produced the highest yields. The over-yielding was due to facilitation between pine and oak, especially the improvement of light conditions in the understory, and an increase in the light use-efficiency of young pine and the availability of light to young oak. Mixing did not reduce competition: pine was dominant in interspecific competition and inhibited the growth of oak. The mixing effect was affected by site quality and stand density: improvements in site quality reduced over-yielding, while increases in stand density increased over-yielding.

Interspecific interactions contribute to higher forage yield and are affected by phosphorus application in a fully-mixed perennial legume and grass intercropping system

Although intercropping is a common technique that has been widely applied, the effect of phosphorus (P) fertilization on productivity of a fully-mixed perennial intercropping system composed of four species, including two legumes and two grasses, has rarely been studied. The objective of this study was to determine the effect of interspecific interactions on forage yield and the influence of P application on interspecific competitiveness and yield in a fully-mixed intercropping system of alfalfa (Medicago sativa L.), sainfoin (Onobrychis viciifolia Scop.), orchardgrass (Dactylis glomerata L.) and tall fescue (Festuca arundinacea Schreb.). A three-year (2016–2018) field experiment was conducted in central China with an annual rainfall and temperature of 456 mm and 9.5 ℃, respectively. Five legume and grass seeding ratios (L: G) of 5:5, 4:6, 3:7, 6:4, and 7:3 were included, with four monocultures used as the control. The total forage yield was measured when 20% of the alfalfa flowers had opened (BBCH 62). Competitiveness indices, including the land equivalent ratio (LER), aggressivity (A), relative crowding coefficient (K values), competitive ratio (CR), and actual yield loss (AYL), were assessed. The total forage yield increased under fully-mixed intercropping in 2017–2018 when a transgressive overyielding advantage was attained, which was contributed by the interspecific interactions and P application. Moreover, the maximum yield and highest intercropping advantage appeared in L: G 3:7 and L: G 5:5 under 80 kg ha−1 P application in 2017–2018. Additionally, adequate P (80 kg ha−1) contributed to higher A values, K values, CR, and AYL of alfalfa and orchardgrass than in sainfoin and tall fescue, which suggested greater interspecific benefits and land-use efficiency. The fully-mixed intercropping system of alfalfa, sainfoin, orchardgrass, and tall fescue realized these potential advantages by coordinating interspecific interactions and environmental resources, which were manipulated by P application.

Functional similarity and competitive symmetry control productivity in mixtures of Mediterranean perennial grasses.

Competition is a major factor structuring plant communities and controlling their productivity. The functional similarity between the interacting species and the context resource availability are assumed to be most critical factors that modulate the strength, sign, and outcome of plant competition, yet their roles and interactions are subjected to debate. In a glasshouse experiment, we constructed monocultures and bi-specific cultures of three common perennial grasses of Mediterranean drylands, the short grass Brachypodium retusum and the tussock grasses Stipa tenacissima and Lygeum spartum, and investigated how the functional similarity between these species modulate their interactions and culture productivity under contrasting levels of water availability. Regardless the degree of functional similarity between the interacting species, B. retusum consistently exhibited a greater competitive ability than the other two species, followed by L. spartum, and with S. tenacissima behaving as the weakest competitor. Bi-specific cultures of B. retusum and either L. spartum or S. tenacissima produced higher biomass than the average biomass of the respective monocultures (i.e. overyielding), whereas the combination of the most similar species, L. spartum—S. tenacissima, which exhibited the highest competition symmetry (i.e., the more similar mutual impact), did not show any significant overyielding. Higher water availability increased productivity and promoted transgressive overyielding for the most dissimilar species, B. retusum and L. spartum, which however exhibited intermediate competition asymmetry. This study calls attention to the thin line between differences in functional traits and competition asymmetry that could eventually lead to either competitive exclusion or resource partitioning and coexistence.

Soil microbiome mediates positive plant diversity‐productivity relationships in late successional grassland species

Which processes drive the productivity benefits of biodiversity remain a critical, but unanswered question in ecology. We tested whether the soil microbiome mediates the diversity-productivity relationships among late successional plant species. We found that productivity increased with plant richness in diverse soil communities, but not with low-diversity mixtures of arbuscular mycorrhizal fungi or in pasteurised soils. Diversity-interaction modelling revealed that pairwise interactions among species best explained the positive diversity-productivity relationships, and that transgressive overyielding resulting from positive complementarity was only observed with the late successional soil microbiome, which was both the most diverse and exhibited the strongest community differentiation among plant species. We found evidence that both dilution/suppression from host-specific pathogens and microbiome-mediated resource partitioning contributed to positive diversity-productivity relationships and overyielding. Our results suggest that re-establishment of a diverse, late successional soil microbiome may be critical to the restoration of the functional benefits of plant diversity following anthropogenic disturbance.

Effect of species identity and diversity on biomass production and its stability in cover crop mixtures

Thanks to positive interactions between species, growing mixtures of cover crops allows improving the ecosystem services provided by cover crop cultivation. In this study, the influence of species diversity but also of species identity and mixture composition on cover crop biomass production and its stability in diverse growing conditions was studied. Several field experiments (varying soil type, preceding crop, soil tillage, sowing density, nitrogen fertilization and spatial replication) were set up in Switzerland during the period 2013−2016. In these experiments, the performance of cover crop species grown as sole crops was compared to that of multispecific mixtures. Part of these experiments followed a simplex design in which four cover crop species were combined together with different proportions, producing a total of 25 mixtures of varied diversity. The other experiments compared sole crop and mixture biomass production in standard randomised block or split plot experiments.Globally, mixtures tended to produce slightly more biomass than the sole crops, with an average between 2 t/ha and 3.2 t/ha for sole crops and of about 3.5 t/ha for mixtures. Overyielding as well as transgressive overyielding were observed, in 81% and 37% of the cases on average, respectively. However no effect of the level of species diversity within mixtures could be found. Biomass production of cover crops was highly influenced by their growing conditions and by the identity of the species involved, especially for sole crops and bispecific mixtures. The analyses of the simplex experiments allowed to show that species interactions played an important role in biomass production in 7 out of 15 growing conditions, even for a short growing period of about three months. Most of the cover crop mixtures with the highest biomass production had a rather low diversity, i.e. about two species on average, but the identity of the species involved in these mixtures depended on the growing conditions. Our results do not show a strong diversity effect on the biomass production of cover crop mixtures cultivated for a short growing period, but a stronger effect of species identity and of the growing conditions. Mixtures with low diversity generally outcompete more diverse mixtures, but more diverse mixtures offer an insurance effect given the unpredictability of growing conditions during cover crop cultivation.

Transgressive overyielding in mixed compared with monospecific Scots pine (Pinus sylvestris L.) and oak (Quercus robur L., Quercus petraea (Matt.) Liebl.) stands – Productivity gains increase with annual water supply

Tree species mixing has become a crucial tool in European forest management as positive interactions between species have been found to promote the provision of multiple ecosystem services, while at the same time reducing the risks associated with climate change. However, mixing effects have proven to be strongly context-dependant and some species combinations have still not been studied in detail. Here we focus on mixed forests of Scots pine and oak, which are likely to become increasingly popular for balancing wood production and other ecosystem services under climate change. Using 20-year growth data from newly established triplets in Germany and Denmark, this study investigates how mean tree and stand characteristics as well as productivity in mixed Scots pine-oak stands compare with adjacent monocultures and how stable the observed productivity relation is, considering inter-annual variations in local climate. Species mixing on average resulted in 15% higher standing volume and 14% higher volume productivity compared with the weighted mean of the adjacent monocultures. Oak profited most in mixture, showing overyielding of 19%. Overyielding on the stand and species level increased in years with higher water supply. In mixture, standing volume of Scots pine was 25% higher than in monocultures. Both species were found to modify their morphology in mixture. Oak in mixture showed a significantly higher inequality in stem volume compared with monocultures. We hypothesise that the observed overyielding of Scots pine-oak mixtures mainly results from complementary light use, where differences in shade tolerance, crown architecture and leaf phenology may be contributing factors.

Levels of forest ecosystem services depend on specific mixtures of commercial tree species.

Global and local ecosystem change resulting in diversity loss has motivated efforts to understand relationships between species diversity and ecosystem services. However, it is unclear how such a general understanding can inform policies for the management of ecosystem services in production systems, because these systems are primarily used for food or fibre, and are rarely managed for the conservation of species diversity. Here, using data from a nationwide forest inventory covering an area of 230,000 km2, we show that relative abundances of commercial tree species in mixed stands strongly influence the potential to provide ecosystem services. The mixes provided higher levels of ecosystem services compared to respective plant monocultures (overyielding or transgressive overyielding) in 35% of the investigated cases, and lower (underyielding) in 9% of the cases. We further show that relative abundances, not just species richness per se, of specific tree-species mixtures affect the potential of forests to provide multiple ecosystem services, which is crucial information for policy and sustainable forest management.