Diversity increases yield but reduces harvest index in crop mixtures.
ABSTRACT: 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.
Project description: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.
Project description:Cover crop benefits include nitrogen accumulation and retention, weed suppression, organic matter maintenance, and reduced erosion. Organic farmers need region-specific information on winter cover crop performance to effectively integrate cover crops into their crop rotations. Our research objective was to compare cover crop seeding mixtures, planting dates, and termination dates on performance of rye (Secale cereale L.) and hairy vetch (Vicia villosa Roth) monocultures and mixtures in the maritime Pacific Northwest USA. The study included four seed mixtures (100% hairy vetch, 25% rye-75% hairy vetch, 50% rye-50% hairy vetch, and 100% rye by seed weight), two planting dates, and two termination dates, using a split-split plot design with four replications over six years. Measurements included winter ground cover; stand composition; cover crop biomass, N concentration, and N uptake; and June soil NO3(-)-N. Rye planted in mid-September and terminated in late April averaged 5.1 Mg ha(-1) biomass, whereas mixtures averaged 4.1 Mg ha(-1) and hairy vetch 2.3 Mg ha(-1). Delaying planting by 2.5 weeks reduced average winter ground cover by 65%, biomass by 50%, and cover crop N accumulation by 40%. Similar reductions in biomass and N accumulation occurred for late March termination, compared with late April termination. Mixtures had less annual biomass variability than rye. Mixtures accumulated 103 kg ha(-1) N and had mean C:N ratio <17:1 when planted in mid-September and terminated in late April. June soil NO3(-)-N (0 to 30 cm depth) averaged 62 kg ha(-1) for rye, 97 kg ha(-1) for the mixtures, and 119 kg ha(-1) for hairy vetch. Weeds comprised less of the mixtures biomass (20% weeds by weight at termination) compared with the monocultures (29%). Cover crop mixtures provided a balance between biomass accumulation and N concentration, more consistent biomass over the six-year study, and were more effective at reducing winter weeds compared with monocultures.
Project description:Cover crop mixtures have the potential to provide more ecosystem services than cover crop monocultures. However, seeding rates that are typically recommended (i.e. seeding rate of monoculture divided by the number of species in the mixture) are non-optimized and often result in the competitive species dominating the mixture, and therefore limiting the amount of ecosystem services that are provided. We created an analytical framework for selecting seeding rates for cover crop mixtures that maximize multifunctionality while minimizing seed costs. The framework was developed using data from a field experiment, which included six response surface designs of two-species mixtures, as well as a factorial replacement design of three-species and four-species mixtures. We quantified intraspecific and interspecific competition among two grasses and two legume cover crop species with grass and legume representing two functional groups: pearl millet [Pennisetum glaucum (L.) R.Br.], sorghum sudangrass [Sorghum bicolor (L.) Moench × Sorghum sudanense (Piper) Stapf], sunn hemp (Crotalaria juncea L.), and cowpea [Vigna unguiculata (L.) Walp]. Yield-density models were fit to estimate intraspecific and interspecific competition coefficients for each species in biculture. The hierarchy from most to least competitive was sorghum sudangrass > sunn hemp > pearl millet > cowpea. Intraspecific competition of a less competitive species was the greatest when the biculture was composed of two species in the same functional group. Competition coefficients were used to build models that estimated the biomass of each cover crop species in three-species and four-species mixtures. The competition coefficients and models were validated with an additional nine site-years testing the same cover crop mixtures. The biomass of a species in a site-year was accurately predicted 69% of the time (low root mean square error, correlation > 0.5, not biased, r<sup>2</sup> > 0.5). Applying the framework, we designed three-species and four-species mixtures by identifying relative seeding rates that produced high biomass with high species evenness (i.e. high multifunctionality) at low seed costs based on a Pareto front analysis of 10,418 mixtures. Accounting for competition when constructing cover crop mixtures can improve the ecosystem services provided, and such an advancement is likely to lead to greater farmer adoption.
Project description:Soil microorganisms are key to biological diversity and many ecosystem processes in terrestrial ecosystems. Despite the current alarming loss of plant diversity, it is unclear how plant species diversity affects soil microorganisms. By conducting a global meta-analysis with paired observations of plant mixtures and monocultures from 106 studies, we show that microbial biomass, bacterial biomass, fungal biomass, fungi:bacteria ratio, and microbial respiration increase, while Gram-positive to Gram-negative bacteria ratio decrease in response to plant mixtures. The increases in microbial biomass and respiration are more pronounced in older and more diverse mixtures. The effects of plant mixtures on all microbial attributes are consistent across ecosystem types including natural forests, planted forests, planted grasslands, croplands, and planted containers. Our study underlines strong relationships between plant diversity and soil microorganisms across global terrestrial ecosystems and suggests the importance of plant diversity in maintaining belowground ecosystem functioning.
Project description:Compared to monocultures, multi-species swards have demonstrated numerous positive diversity effects on aboveground plant performance, such as yield, N concentration, and even legacy effects on a following crop. Whether such diversity effects are seen in the soil microbiome is currently unclear. In a field experiment, we analyzed the effect that three plant species (a grass, forb, and legume), and mixtures of these, had on soil fungal and bacterial community structures, as well as their associated legacy effects under a following crop, the grass Lolium multiflorum. We utilized six sward types, three monocultures (Lolium perenne, Cichorium intybus and Trifolium pratense), two bi-species mixtures, and a mixture of the three species. Soil samples were taken from these swards in March (at the end of a three year conditioning phase) and in June, August, and September after L. multiflorum was established, that is, the legacy samplings. When present, the differing monocultures had a significant effect on various aspects of the fungal community: structure, OTU richness, the relative abundance of the phylum Glomeromycota, and indicator OTUs. The effect on bacterial community structure was not as strong. In the multi-species swards, a blending of individual plant species monoculture effects (identity effect) was seen in (a) fungal and bacterial community structure and (b) fungal OTU richness and the relative abundance of the Glomeromycota. This would indicate that plant species identity, rather than diversity effects (i.e., the interactions among the plant species), was the stronger determinant. During the legacy samplings, structural patterns in the fungal and bacterial communities associated with the previous swards were retained, but the effect faded with time. These results highlight that plant species identity can be a strong driver of soil microbial community structures. They also suggest that their legacy effect on the soil microbiome may play a crucial role in following crop performance.
Project description:Grassland diversity can support sustainable intensification of grassland production through increased yields, reduced inputs and limited weed invasion. We report the effects of diversity on weed suppression from 3 years of a 31-site continental-scale field experiment.At each site, 15 grassland communities comprising four monocultures and 11 four-species mixtures based on a wide range of species' proportions were sown at two densities and managed by cutting. Forage species were selected according to two crossed functional traits, "method of nitrogen acquisition" and "pattern of temporal development".Across sites, years and sown densities, annual weed biomass in mixtures and monocultures was 0.5 and 2.0 t DM ha<sup>-1</sup> (7% and 33% of total biomass respectively). Over 95% of mixtures had weed biomass lower than the average of monocultures, and in two-thirds of cases, lower than in the most suppressive monoculture (transgressive suppression). Suppression was significantly transgressive for 58% of site-years. Transgressive suppression by mixtures was maintained across years, independent of site productivity.Based on models, average weed biomass in mixture over the whole experiment was 52% less (95% confidence interval: 30%-75%) than in the most suppressive monoculture. Transgressive suppression of weed biomass was significant at each year across all mixtures and for each mixture.Weed biomass was consistently low across all mixtures and years and was in some cases significantly but not largely different from that in the equiproportional mixture. The average variability (standard deviation) of annual weed biomass within a site was much lower for mixtures (0.42) than for monocultures (1.77). <i>Synthesis and applications</i>. Weed invasion can be diminished through a combination of forage species selected for complementarity and persistence traits in systems designed to reduce reliance on fertiliser nitrogen. In this study, effects of diversity on weed suppression were consistently strong across mixtures varying widely in species' proportions and over time. The level of weed biomass did not vary greatly across mixtures varying widely in proportions of sown species. These diversity benefits in intensively managed grasslands are relevant for the sustainable intensification of agriculture and, importantly, are achievable through practical farm-scale actions.
Project description:Legumes rely on soil mineral nitrogen (N) and biological N fixation (BNF). The interplay between these two sources is biologically interesting and agronomically relevant as the crop can accommodate the cost of BNF by five non-mutually exclusive mechanisms, whereby BNF: reduces shoot growth and seed yield, or maintains shoot growth and seed yield by enhanced photosynthesis, or reduced root:shoot ratio, or maintains shoot growth but reduces seed yield by reducing the fraction of shoot biomass allocated to seed (harvest index), or reducing concentration of oil and protein in seed. We explore the impact of N application on the seasonal dynamics of BNF, and its consequences for seed yield with emphasis on growth and shoot allocation mechanisms. Trials were established in 23 locations across the US Midwest under four N conditions. Fertilizer reduced the peak of BNF up to 16% in applications at the full flowering stage. Seed yield declined 13?kg?ha<sup>-1</sup> per % increase in RAU<sub>R6</sub>. Harvest index accounted for the decline in seed yield with increasing BNF. This indicates the cost of BNF was met by a relative change in dry matter allocation against the energetically rich seed, and in favor of energetically cheaper vegetative tissue.
Project description:<h4>Background and aims</h4>Plant genotypic mixtures have the potential to increase yield stability in variable, often unpredictable environments, yet knowledge of the specific mechanisms underlying enhanced yield stability remains limited. Field studies are constrained by environmental conditions which cannot be fully controlled and thus reproduced. A suitable model system would allow reproducible experiments on processes operating within crop genetic mixtures.<h4>Methods</h4>Phenotypically dissimilar genotypes of Arabidopsis thaliana were grown in monocultures and mixtures under high levels of competition for abiotic resources. Seed production, flowering time and rosette size were recorded.<h4>Key results</h4>Mixtures achieved high yield stability across environments through compensatory interactions. Compensation was greatest when plants were under high levels of heat and nutrient stress. Competitive ability and mixture performance were predictable from above-ground phenotypic traits even though below-ground competition appeared to be more intense.<h4>Conclusions</h4>This study indicates that the mixing ability of plant genotypes can be predicted from their phenotypes expressed in a range of relevant environments, and implies that a phenotypic screen of genotypes could improve the selection of suitable components of genotypic mixtures in agriculture intended to be resilient to environmental stress.
Project description:Numerous experiments, mostly performed in particular environments, have shown positive diversity-productivity relationships. Although the complementary use of resources is discussed as an important mechanism explaining diversity effects, less is known about how resource availability controls the strength of diversity effects and how this response depends on the functional composition of plant communities. We studied aboveground biomass production in experimental monocultures, two- and four-species mixtures assembled from two independent pools of four perennial grassland species, each representing two functional groups (grasses, forbs) and two growth statures (small, tall), and exposed to different combinations of light and nutrient availability. On average, shade led to a decrease in aboveground biomass production of 24% while fertilization increased biomass production by 36%. Mixtures were on average more productive than expected from their monocultures (relative yield total, RYT>1) and showed positive net diversity effects (NE: +34% biomass increase; mixture minus mean monoculture biomass). Both trait-independent complementarity effects (TICE: +21%) and dominance effects (DE: +12%) positively contributed to net diversity effects, while trait-dependent complementarity effects were minor (TDCE: +1%). Shading did not alter diversity effects and overyielding. Fertilization decreased RYT and the proportion of biomass gain through TICE and TDCE, while DE increased. Diversity effects did not increase with species richness and were independent of functional group or growth stature composition. Trait-based analyses showed that the dominance of species with root and leaf traits related to resource conservation increased TICE. Traits indicating the tolerance of shade showed positive relationships with TDCE. Large DE were associated with the dominance of species with tall growth and low diversity in leaf nitrogen concentrations. Our field experiment shows that positive diversity effects are possible in grass-forb mixtures irrespective of differences in light availability, but that the chance for the complementary use of resources increases when nutrients are not available at excess.
Project description:Biodiversity experiments show that increases in plant diversity can lead to greater biomass production, and some researchers suggest that high diversity plantings should be used for bioenergy production. However, many methods used in past biodiversity experiments are impractical for bioenergy plantings. For example, biodiversity experiments often use intensive management such as hand weeding to maintain low diversity plantings and exclude unplanted species, but this would not be done for bioenergy plantings. Also, biodiversity experiments generally use high seeding densities that would be too expensive for bioenergy plantings. Here we report the effects of biodiversity on biomass production from two studies of more realistic bioenergy crop plantings in southern Michigan, USA. One study involved comparing production between switchgrass (Panicum virgatum) monocultures and species-rich prairie plantings on private farm fields that were managed similarly to bioenergy plantings. The other study was an experiment where switchgrass was planted in monoculture and in combination with increasingly species-rich native prairie mixtures. Overall, we found that bioenergy plantings with higher species richness did not produce more biomass than switchgrass monocultures. The lack of a positive relationship between planted species richness and production in our studies may be due to several factors. Non-planted species (weeds) were not removed from our studies and these non-planted species may have competed with planted species and also prevented realized species richness from equaling planted species richness. Also, we found that low seeding density of individual species limited the biomass production of these individual species. Production in future bioenergy plantings with high species richness may be increased by using a high density of inexpensive seed from switchgrass and other highly productive species, and future efforts to translate the results of biodiversity experiments to bioenergy plantings should consider the role of seeding density.