Simultaneously enhancing ecosystem services provided by biodiversity below and above ground is recommended to reduce dependence on chemical pesticides and mineral fertilisers in agriculture. However, consequences for crop yield have been poorly evaluated. Above ground, increased landscape complexity is assumed to enhance biological pest control, whereas below ground, soil organic carbon is a proxy for several yield-supporting services. In a field experiment replicated in 114 fields across Europe, we found that fertilisation had the strongest positive effect on yield, but hindered simultaneous harnessing of below- and above-ground ecosystem services. We furthermore show that enhancing natural enemies and pest control through increasing landscape complexity can prove disappointing in fields with low soil services or in intensively cropped regions. Thus, understanding ecological interdependences between land use, ecosystem services and yield is necessary to promote more environmentally friendly farming by identifying situations where ecosystem services are maximised and agrochemical inputs can be reduced.

Soil organic matter (SOM) is declining in most agricultural ecosystems, impacting multiple ecosystem services including erosion and flood prevention, climate and greenhouse gas regulation as well as other services that underpin crop production, such as nutrient cycling and pest control. Ecological intensification aims to enhance crop productivity by including regulating and supporting ecosystem service management into agricultural practices. We investigate the potential for increased SOM to support the ecological intensification of arable systems by reducing the need for nitrogen fertiliser application and pest control. Using a large-scale European field trial implemented across 84 fields in 5 countries, we tested whether increased SOM (using soil organic carbon as a proxy) helps recover yield in the absence of conventional nitrogen fertiliser and whether this also supports crops less favourable to key aphid pests. Greater SOM increased yield by 10%, but did not offset nitrogen fertiliser application entirely, which improved yield by 30%. Crop pest responses depended on species: Metopolophium dirhodum were more abundant in fertilised plots with high crop biomass, and although population growth rates of Sitobion avenae were enhanced by nitrogen fertiliser application in a cage trial, field populations were not affected. We conclude that under increased SOM and reduced fertiliser application, pest pressure can be reduced, while partially compensating for yield deficits linked to fertiliser reduction. If the benefits of reduced fertiliser application and increased SOM are considered in a wider environmental context, then a yield cost may become acceptable. Maintaining or increasing SOM is critical for achieving ecological intensification of European cereal production.

Summary1. Above-ground and below-ground environmental conditions influence crop yield by pollination, pest pressure and resource supply. However, little is known about how interactions between these factors contribute to yield. Here, we used oilseed rape Brassica napus to test their effects on crop yield. 2. We exposed potted plants to all combinations of high and low levels of soil organic matter (SOM) and fertilizer supply, and placed all treatments at a variety of field sites representing a gradient in pollinator visitation rate and pest exposure. We determined the relative contribution of pollinators and pests, SOM and fertilizer supply to yield. We also tested whether SOM can moderate effects of fertilizer on yield and whether soil conditions influence the relationship between above-ground conditions and yield. 3. Increases in pollinator visitation rate and decreases in pest pressure enhanced yield more than increase in fertilizer supply. Although higher SOM content resulted in plants with more biomass and flowers, under our experimental conditions SOM neither enhanced yield, nor influenced effects of fertilizer, pollinators or pests on yield. 4. The relationships between yield, pollinator visitation rate and pest pressure did not depend on the level of fertilization, suggesting that the effects of fertilizer application and aboveground (dis)services on yield were additive. In contrast, pollinator visitation rate was more strongly related to yield at low pest pressure than at high pest pressure indicating trade-offs between above-ground services and disservices. 5. Synthesis and applications. Our results show that it is possible to increase oilseed rape yield by enhancing pollination, irrespective of supplying mineral fertilizer. Moreover, the fact that belowground conditions did not alter the effect of above-ground conditions suggests that farmers may obtain even higher yields by maximizing both above-ground ecosystem services and external inputs. Further studies are needed to understand at which point the positive relationships between pollinator visitation and yield, as well as between fertilizer and yield, will level off. Considering above-ground and below-ground services and inputs in agro-ecosystems in conjunction is crucial in order to optimize external inputs for crop yield from an economic and ecological perspective.

to design landscapes that accommodate beneficial aboveground communities with respect to their required resources. For successful adoption of above-belowground interactions in agriculture there is a need for context-specific solutions, as well as sound socio-economic embedding.

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There is increasing evidence showing that microbes can influence plant-insect interactions. In addition, various studies have shown that aboveground pathogens can alter the interactions between plants and insects. However, little is known about the role of soil-borne pathogens in plant-insect interactions. It is also not known how environmental conditions, that steer the performance of soil-borne pathogens, might influence these microbe-plant-insect interactions. Here, we studied effects of the soil-borne pathogen Rhizoctonia solani on aphids (Sitobion avenae) using wheat (Triticum aestivum) as a host.In a greenhouse experiment, we tested how different levels of soil organic matter (SOM) and fertilizer addition influence the interactions between plants and aphids. To examine the influence of the existing soil microbiome on the pathogen effects, we used both unsterilized field soil and sterilized field soil.In unsterilized soil with low SOM content, R. solani addition had a negative effect on aphid biomass, whereas it enhanced aphid biomass in soil with high SOM content. In sterilized soil, however, aphid biomass was enhanced by R. solani addition and by high SOM content. Plant biomass was enhanced by fertilizer addition, but only when SOM content was low, or in the absence of R. solani.We conclude that belowground pathogens influence aphid performance and that the effect of soil pathogens on aphids can be more positive in the absence of a soil microbiome. This implies that experiments studying the effect of pathogens under sterile conditions might not represent realistic interactions. Moreover, pathogen-plant-aphid interactions can be more positive for aphids under high SOM conditions. We recommend that soil conditions should be taken into account in the study of microbe-plant-insect interactions.

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