Enhanced weathering in the US Corn Belt delivers carbon removal with agronomic benefits.

Terrestrial enhanced weathering (EW) of silicate rocks, such as crushed basalt, on farmlands is a promising scalable atmospheric carbon dioxide removal (CDR) strategy that urgently requires performance assessment with commercial farming practices. We report findings from a large-scale replicated EW field trial across a typical maize-soybean rotation on an experimental farm in the heart of the United Sates Corn Belt over 4 y (2016 to 2020). We show an average combined loss of major cations (Ca2+ and Mg2+) from crushed basalt applied each fall over 4 y (50 t ha-1 y-1) gave a conservative time-integrated cumulative CDR potential of 10.5 ± 3.8 t CO2 ha-1. Maize and soybean yields increased significantly (P < 0.05) by 12 to 16% with EW following improved soil fertility, decreased soil acidification, and upregulation of root nutrient transport genes. Yield enhancements with EW were achieved with significantly (P < 0.05) increased key micro- and macronutrient concentrations (including potassium, magnesium, manganese, phosphorus, and zinc), thus improving or maintaining crop nutritional status. We observed no significant increase in the content of trace metals in grains of maize or soybean or soil exchangeable pools relative to controls. Our findings suggest that widespread adoption of EW across farming sectors has the potential to contribute significantly to net-zero greenhouse gas emissions goals while simultaneously improving food and soil security.

Virus-induced changes in root volatiles attract soil nematode vectors to infected plants.

Plant-derived volatiles mediate interactions among plants, pathogenic viruses, and viral vectors. These volatile-dependent mechanisms have not been previously demonstrated belowground, despite their likely significant role in soil ecology and agricultural pest impacts. We investigated how the plant virus, tobacco rattle virus (TRV), attracts soil nematode vectors to infected plants. We infected Nicotiana benthamiana with TRV and compared root growth relative to that of uninfected plants. We tested whether TRV-infected N. benthamiana was more attractive to nematodes 7 d post infection and identified a compound critical to attraction. We also infected N. benthamiana with mutated TRV strains to identify virus genes involved in vector nematode attraction. Virus titre and associated impacts on root morphology were greatest 7 d post infection. Tobacco rattle virus infection enhanced 2-ethyl-1-hexanol production. Nematode chemotaxis and 2-ethyl-1-hexanol production correlated strongly with viral load. Uninfected plants were more attractive to nematodes after the addition of 2-ethyl-1-hexanol than were untreated plants. Mutation of TRV RNA2-encoded genes reduced the production of 2-ethyl-1-hexanol and nematode attraction. For the first time, this demonstrates that virus-driven alterations in root volatile emissions lead to increased chemotaxis of the virus's nematode vector, a finding with implications for sustainable management of both nematodes and viral pathogens in agricultural systems.

Growth-defence trade-off in rice: fast-growing and acquisitive genotypes have lower expression of genes involved in immunity.

Plant ecologists and molecular biologists have long considered the hypothesis of a trade-off between plant growth and defence separately. In particular, how genes thought to control the growth-defence trade-off at the molecular level relate to trait-based frameworks in functional ecology, such as the slow-fast plant economics spectrum, is unknown. We grew 49 phenotypically diverse rice genotypes in pots under optimal conditions and measured growth-related functional traits and the constitutive expression of 11 genes involved in plant defence. We also quantified the concentration of silicon (Si) in leaves to estimate silica-based defences. Rice genotypes were aligned along a slow-fast continuum, with slow-growing, late-flowering genotypes versus fast-growing, early-flowering genotypes. Leaf dry matter content and leaf Si concentrations were not aligned with this axis and negatively correlated with each other. Live-fast genotypes exhibited greater expression of OsNPR1, a regulator of the salicylic acid pathway that promotes plant defence while suppressing plant growth. These genotypes also exhibited greater expression of SPL7 and GH3.2, which are also involved in both stress resistance and growth. Our results do not support the hypothesis of a growth-defence trade-off when leaf Si and leaf dry matter content are considered, but they do when hormonal pathway genes are considered. We demonstrate the benefits of combining ecological and molecular approaches to elucidate the growth-defence trade-off, opening new avenues for plant breeding and crop science.

Associational resistance through intercropping reduces yield losses to soil-borne pests and diseases.

Associational resistance to herbivore and pathogen attack is a well documented ecological phenomenon and, if applied to agriculture, may reduce impact of pests and diseases on crop yields without recourse to pesticides. The value of associational resistance through intercropping, planting multiple crops alongside each other, as a sustainable control method remains unclear, due to variable outcomes reported in the published literature. We performed a meta-analysis to provide a quantitative assessment of benefits of intercropping for target plant resistance to plant-parasitic nematodes and soil-borne diseases. We found that intercropping reduced damage to focal crops from nematodes by 40% and disease incidence by 55%. Intercropping efficacy varied with biological variables, such as field fertilisation status and intercrop family, and methodology, including whether study samples were potted or in fields. Nematode control using intercropping was sufficient to offset reductions in focal crop yield from intercrop presence, making intercropping a viable agricultural tool. We identify key drivers for underpinning the success of intercropping and indicate areas for future research to improve efficacy. This study also highlights the potential benefits of harnessing ecological knowledge on plant-enemy interactions for improving agricultural and landscape sustainability.

Plant silicon application alters leaf alkaloid concentrations and impacts parasitoids more adversely than their aphid hosts.

Grasses accumulate large amounts of silicon (Si) which acts as a highly effective physical defence against insect herbivory, however recent evidence shows that Si supplementation also modifies plant secondary metabolite concetrations. Changes in plant secondary metabolites concentrations can have cascading effects on higher trophic levels, such as parasitoids, as they are dependent on the host herbivore for growth and development. However, relatively little is known about how Si application affects higher trophic levels. We examined the effects of Si addition on alkaloid content in leaves of Phalaris aquatica (Poaceae) and the effect on interactions between an aphid (Rhopalosiphum padi) and its parasitoid (Aphidius colemani). Si supplementation had no effect on aphid abundance or parasitism rate. Adult aphids, aphid mummies (parasitised aphids) and the emergent parasitoids were, however, significantly smaller on Si+ plants. Parasitoid traits (size and emergence) were correlated with aphid mummy size. Si addition reduced parasitoid emergence rate and size due to reduced host mummy size, in addition, significantly fewer females emerged from mummies on Si+ plants. Aphid infestation significantly altered alkaloids concentrations, reducing gramine by 80% while increasing tryptamine by 91% in Si- plants. Si addition reduced aphid-induced tryptamine concentrations by 64% and increased 5-MeO-tryptamine by over 800% in control and 142% in aphid infested plants. Our results show that while Si addition has modest impacts on the herbivore, it significantly alters secondary metabolites and has stronger effects on the higher trophic level through changes in the quality of the parasitised host.

High silicon concentrations in grasses are linked to environmental conditions and not associated with C4 photosynthesis.

The uptake and deposition of silicon (Si) as silica phytoliths is common among land plants and is associated with a variety of functions. Among these, herbivore defense has received significant attention, particularly with regard to grasses and grasslands. Grasses are well known for their high silica content, a trait which has important implications ranging from defense to global Si cycling. Here, we test the classic hypothesis that C4 grasses evolved stronger mechanical defenses than C3 grasses through increased phytolith deposition, in response to extensive ungulate herbivory ("C4 -grazer hypothesis"). Despite mixed support, this hypothesis has received broad attention, even outside the realm of plant biology. Because C3 and C4 grasses typically dominate in different climates, with the latter more abundant in hot, dry regions, we also investigated the effects of water availability and temperature on Si deposition. We compiled a large dataset of grasses grown under controlled environmental conditions. Using phylogenetically informed generalized linear mixed models and character evolution models, we evaluated whether photosynthetic pathway or growth condition influenced Si concentration. We found that C4 grasses did not show consistently elevated Si concentrations compared with C3 grasses. High temperature treatments were associated with increased concentration, especially in taxa adapted to warm regions. Although the effect was less pronounced, reduced water treatment also promoted silica deposition, with slightly stronger response in dry habitat species. The evidence presented here rejects the "C4 -grazer hypothesis." Instead, we propose that the tendency for C4 grasses to outcompete C3 species under hot, dry conditions explains previous observations supporting this hypothesis. These findings also suggest a mechanism via which anthropogenic climate change may influence silica deposition in grasses and, by extension, alter the important ecological and geochemical processes it affects.

Increased yield and CO2 sequestration potential with the C4 cereal Sorghum bicolor cultivated in basaltic rock dust-amended agricultural soil.

Land-based enhanced rock weathering (ERW) is a biogeochemical carbon dioxide removal (CDR) strategy aiming to accelerate natural geological processes of carbon sequestration through application of crushed silicate rocks, such as basalt, to croplands and forested landscapes. However, the efficacy of the approach when undertaken with basalt, and its potential co-benefits for agriculture, require experimental and field evaluation. Here we report that amending a UK clay-loam agricultural soil with a high loading (10 kg/m2 ) of relatively coarse-grained crushed basalt significantly increased the yield (21 ± 9.4%, SE) of the important C4 cereal Sorghum bicolor under controlled environmental conditions, without accumulation of potentially toxic trace elements in the seeds. Yield increases resulted from the basalt treatment after 120 days without P- and K-fertilizer addition. Shoot silicon concentrations also increased significantly (26 ± 5.4%, SE), with potential benefits for crop resistance to biotic and abiotic stress. Elemental budgets indicate substantial release of base cations important for inorganic carbon removal and their accumulation mainly in the soil exchangeable pools. Geochemical reactive transport modelling, constrained by elemental budgets, indicated CO2 sequestration rates of 2-4 t CO2 /ha, 1-5 years after a single application of basaltic rock dust, including via newly formed soil carbonate minerals whose long-term fate requires assessment through field trials. This represents an approximately fourfold increase in carbon capture compared to control plant-soil systems without basalt. Our results build support for ERW deployment as a CDR technique compatible with spreading basalt powder on acidic loamy soils common across millions of hectares of western European and North American agriculture.

Arbuscular Mycorrhizal Fungi and Plant Chemical Defence: Effects of Colonisation on Aboveground and Belowground Metabolomes.

Arbuscular mycorrhizal fungal (AMF) colonisation of plant roots is one of the most ancient and widespread interactions in ecology, yet the systemic consequences for plant secondary chemistry remain unclear. We performed the first metabolomic investigation into the impact of AMF colonisation by Rhizophagus irregularis on the chemical defences, spanning above- and below-ground tissues, in its host-plant ragwort (Senecio jacobaea). We used a non-targeted metabolomics approach to profile, and where possible identify, compounds induced by AMF colonisation in both roots and shoots. Metabolomics analyses revealed that 33 compounds were significantly increased in the root tissue of AMF colonised plants, including seven blumenols, plant-derived compounds known to be associated with AMF colonisation. One of these was a novel structure conjugated with a malonyl-sugar and uronic acid moiety, hitherto an unreported combination. Such structural modifications of blumenols could be significant for their previously reported functional roles associated with the establishment and maintenance of AM colonisation. Pyrrolizidine alkaloids (PAs), key anti-herbivore defence compounds in ragwort, dominated the metabolomic profiles of root and shoot extracts. Analyses of the metabolomic profiles revealed an increase in four PAs in roots (but not shoots) of AMF colonised plants, with the potential to protect colonised plants from below-ground organisms.

Mapping regional risks from climate change for rainfed rice cultivation in India.

Global warming is predicted to increase in the future, with detrimental consequences for rainfed crops that are dependent on natural rainfall (i.e. non-irrigated). Given that many crops grown under rainfed conditions support the livelihoods of low-income farmers, it is important to highlight the vulnerability of rainfed areas to climate change in order to anticipate potential risks to food security. In this paper, we focus on India, where ~ 50% of rice is grown under rainfed conditions, and we employ statistical models (climate envelope models (CEMs) and boosted regression trees (BRTs)) to map changes in climate suitability for rainfed rice cultivation at a regional level (~ 18 × 18 km cell resolution) under projected future (2050) climate change (IPCC RCPs 2.6 and 8.5, using three GCMs: BCC-CSM1.1, MIROC-ESM-CHEM, and HadGEM2-ES). We quantify the occurrence of rice (whether or not rainfed rice is commonly grown, using CEMs) and rice extent (area under cultivation, using BRTs) during the summer monsoon in relation to four climate variables that affect rice growth and yield namely ratio of precipitation to evapotranspiration (PER), maximum and minimum temperatures (Tmax and Tmin ), and total rainfall during harvesting. Our models described the occurrence and extent of rice very well (CEMs for occurrence, ensemble AUC = 0.92; BRTs for extent, Pearson's r = 0.87). PER was the most important predictor of rainfed rice occurrence, and it was positively related to rainfed rice area, but all four climate variables were important for determining the extent of rice cultivation. Our models project that 15%-40% of current rainfed rice growing areas will be at risk (i.e. decline in climate suitability or become completely unsuitable). However, our models project considerable variation across India in the impact of future climate change: eastern and northern India are the locations most at risk, but parts of central and western India may benefit from increased precipitation. Hence our CEM and BRT models agree on the locations most at risk, but there is less consensus about the degree of risk at these locations. Our results help to identify locations where livelihoods of low-income farmers and regional food security may be threatened in the next few decades by climate changes. The use of more drought-resilient rice varieties and better irrigation infrastructure in these regions may help to reduce these impacts and reduce the vulnerability of farmers dependent on rainfed cropping.

A Zoospore Inoculation Method with Phytophthora sojae to Assess the Prophylactic Role of Silicon on Soybean Cultivars.

The objective of this study was to evaluate whether silicon (Si) amendments, known to have a prophylactic role against biotrophic and hemibiotrophic pathogens, could protect soybean against Phytophthora sojae. To fulfill this objective, the initial challenge was to develop a method of inoculation that reproduced the natural infection process while allowing regular Si feeding to the plants. In a first set of experiments, inoculation of P. sojae zoospores directly into hydroponic solutions led to reproducible infections and expected phenotypes when using 'Williams' (rps), 'L75-6141' (Rps1a), 'haro15' (Rps1k), and 'L77-1863' (Rps1b) soybean challenged to races 3 and 7 of P. sojae. This approach offers the advantage of testing simultaneously many soybean cultivars against different races of P. sojae in a controlled environment, and the expression of partial and root resistance. In a second set of experiments aimed at testing the effect of Si, our results clearly showed that Si amendments had a significant effect on disease reduction and plant yield. The effect was particularly noticeable when combined with a cultivar displaying a certain level of resistance to the disease. These results demonstrate a useful method of direct inoculation of soybean plants with P. sojae zoospores through a hydroponic system and show that Si amendments can represent an alternative method of control of P. sojae against soybean.

Why do plants silicify?

Despite seminal papers that stress the significance of silicon (Si) in plant biology and ecology, most studies focus on manipulations of Si supply and mitigation of stresses. The ecological significance of Si varies with different levels of biological organization, and remains hard to capture. We show that the costs of Si accumulation are greater than is currently acknowledged, and discuss potential links between Si and fitness components (growth, survival, reproduction), environment, and ecosystem functioning. We suggest that Si is more important in trait-based ecology than is currently recognized. Si potentially plays a significant role in many aspects of plant ecology, but knowledge gaps prevent us from understanding its possible contribution to the success of some clades and the expansion of specific biomes.

Delayed induced silica defences in grasses and their potential for destabilising herbivore population dynamics.

Some grass species mount a defensive response to grazing by increasing their rate of uptake of silica from the soil and depositing it as abrasive granules in their leaves. Increased plant silica levels reduce food quality for herbivores that feed on these grasses. Here we provide empirical evidence that a principal food species of an herbivorous rodent exhibits a delayed defensive response to grazing by increasing silica concentrations, and present theoretical modelling that predicts that such a response alone could lead to the population cycles observed in some herbivore populations. Experiments performed under greenhouse conditions revealed that the rate of deposition of silica defences in the grass Deschampsia caespitosa is a time-lagged, nonlinear function of grazing intensity and that, upon cessation of grazing, these defences take around one year to decay to within 5 % of control levels. Simple coupled grass-herbivore population models incorporating this functional response, and parameterised with empirical data, consistently predict population cycles for a wide range of realistic parameter values for a (Microtus) vole-grass system. Our results support the hypothesis that induced silica defences have the potential to strongly affect the population dynamics of their herbivores. Specifically, the feedback response we observed could be a driving mechanism behind the observed population cycles in graminivorous herbivores in cases where grazing levels in the field become sufficiently large and sustained to trigger an induced silica defence response.

Nitrogen availability and plant-plant interactions drive leaf silicon concentration in wheat genotypes.

Estimating plasticity of leaf silicon (Si) in response to abiotic and biotic factors underpins our comprehension of plant defences and stress resistance in natural and agroecosystems. However, how nitrogen (N) addition and intraspecific plant-plant interactions affect Si concentration remains unclear.We grew 19 durum wheat genotypes (Triticum turgidum ssp. durum) in pots, either alone or in intra- or intergenotypic cultures of two individuals, and with or without N. Above-ground biomass, plant height and leaf [Si] were quantified at the beginning of the flowering stage.Nitrogen addition decreased leaf [Si] for most genotypes, proportionally to the biomass increase. Si plasticity to plant-plant interactions varied significantly among genotypes, with both increases and decreases in leaf [Si] when mixed with a neighbour, regardless of the mixture type (intra-/intergenotype). Besides, increased leaf [Si] in response to plant-plant interactions was associated with increased plant height.Our results suggest the occurrence of both facilitation and competition for Si uptake from the rhizosphere in wheat mixtures. Future research should identify which leaf and root traits characterise facilitating neighbours for Si acquisition. We also show that Si could be involved in height gain in response to intraspecific competition, possibly for increasing light capture. This important finding opens up new research directions on Si and plant-plant interactions in both natural ecosystems and agroecosystems. More generally, our results stress the need to explore leaf Si plasticity in responses to both abiotic and biotic factors to understand plant stress resistance. Read the free Plain Language Summary for this article on the Journal blog.

Herbivory of tropical rain forest tree seedlings correlates with future mortality.

Tree seedlings in tropical rain forests are subject to both damage from natural enemies and intense interspecific competition. This leads to a trade-off in investment between defense and growth, and it is likely that tree species specialized to particular habitats tailor this balance to correspond with local resource availability. It has also been suggested that differential herbivore impacts among tree species may drive habitat segregation, favoring species adapted to particular resource conditions. In order to test these predictions, a reciprocal transplant experiment in Sabah, Malaysia, was established with seedlings of five species of Dipterocarpaceae. These were specialized to either alluvial (Hopea nervosa, Parashorea tomentella) or sandstone soils (Shorea multiflora, H. beccariana), or were locally absent (S. fallax). A total of 3000 seedlings were planted in paired gap and understory plots in five sites on alluvial and sandstone soils. Half of all seedlings were fertilized. Seedling growth and mortality were recorded in regular samples over 3.5 years, and rates of insect herbivore damage were estimated from censuses of foliar tissue loss on marked mature leaves and available young leaves. Greater herbivory rates on mature leaves had no measurable effects on seedling growth but were associated with a significantly increased likelihood of mortality during the following year. In contrast, new-leaf herbivory rates correlated with neither growth nor mortality. There were no indications of differential impacts of herbivory among the five species, nor between experimental treatments. Herbivory was not shown to influence segregation of species between soil types, although it may contribute toward differential survival among light habitats. Natural rates of damage were substantially lower than have been shown to influence tree seedling growth and mortality in previous manipulative studies.

Impact of osmotic stress on the growth and root architecture of introgression lines derived from a wild ancestor of rice and a modern cultivar.

Many modern rice varieties have been intensively selected for high-yielding performance under irrigated conditions, reducing their genetic diversity and potentially increasing their susceptibility to abiotic stresses such as drought. In this study, we tested benefits for stress tolerance of introducing DNA segments from wild ancestor Oryza rufipogon to the modern cultivar O. sativa cv Curinga (CUR) by applying a gradient of osmotic stress to both parents and seven introgressed lines. Shoot growth of O. rufipogon had a high tolerance to osmotic stress, and the number of total root tips increased under mild osmotic stress. One introgression line showed greater shoot growth, root growth, and higher number of total root tips than the parent line CUR under osmotic stress. Abscisic acid (ABA) is a key hormone mediating plant responses to abiotic stresses. Both root and shoot growth of O. rufipogon were much more sensitive to ABA than CUR. Introgression lines varied in the extent to which the sensitivity of their growth responses to ABA and some lines correlated with their sensitivity to osmotic stress. Our results suggest that rice responses to ABA and osmotic stress are genotype dependent, and growth responses of rice to ABA are not a consistent indicator of resilience to abiotic stress in introgression lines.

Physical defences wear you down: progressive and irreversible impacts of silica on insect herbivores.

1. Silica in the leaves of grasses can act as a defence against both vertebrate and invertebrate herbivores. The mechanisms by which silica affects herbivore performance are not well characterized. Here we expose an insect herbivore Spodoptera exempta to high-silica diets and test two mechanisms by which silica has been proposed to act as a defence. First, that silica reduces the digestibility of leaves and second, that silica causes wear to insect mandibles, both of which could potentially impact on herbivore performance. 2. Silica reduced the efficiency with which S. exempta converted ingested food to body mass and the amount of nitrogen absorbed from their food, leading to reduced insect growth rates. The measure of how efficiently herbivores utilize digested food (ECD) was unaffected by silica. 3. These effects occurred even with short-term exposure to silica-rich diets, but they also increased markedly with the duration of exposure and affected late instars more than early instar larvae. This appears to be due to the progressive impacts of silica with longer exposure times and suggests that herbivores cannot adapt to silica defences, nor do they develop a tolerance for silica with age. 4. Exposure to silica-rich diets caused increased mandible wear in S. exempta. This effect was extremely rapid, occurring within a single instar, further reducing feeding efficiency and growth rates. These effects on insect growth and feeding efficiency are nonreversible, persisting after the herbivore has switched diets. Up to a third of this residual impact can be explained by the degree of mandible wear caused by previous silica-rich diets. 5. The impacts of silica on S. exempta larvae were progressive with exposure time and could not be compensated for, even by switching to a different diet. Thus, herbivores cannot easily adapt to physical defences such as silica, suggesting this defence will have major implications for herbivore fitness.

Aphids Influence Soil Fungal Communities in Conventional Agricultural Systems.

Arbuscular mycorrhizal fungi (AMF) form symbioses with the roots of most plant species, including cereals. AMF can increase the uptake of nutrients including nitrogen (N) and phosphorus (P), and of silicon (Si) as well as increase host resistance to various stresses. Plants can simultaneously interact with above-ground insect herbivores such as aphids, which can alter the proportion of plant roots colonized by AMF. However, it is unknown if aphids impact the structure of AMF communities colonizing plants or the extent of the extraradical mycelium produced in the soil, both of which can influence the defensive and nutritional benefit a plant derives from the symbiosis. This study investigated the effect of aphids on the plant-AMF interaction in a conventionally managed agricultural system. As plants also interact with other soil fungi, the non-AMF fungal community was also investigated. We hypothesized that aphids would depress plant growth, and reduce intraradical AMF colonization, soil fungal hyphal density and the diversity of AM and non-AM fungal communities. To test the effects of aphids, field plots of barley enclosed with insect proof cages were inoculated with Sitobion avenae or remained uninoculated. AMF specific and total fungal amplicon sequencing assessed root fungal communities 46 days after aphid addition. Aphids did not impact above-ground plant biomass, but did increase the grain N:P ratio. Whilst aphid presence had no impact on AMF intraradical colonization, soil fungal hyphal length density, or AMF community characteristics, there was a trend for the aphid treatment to increase vesicle numbers and the relative abundance of the AMF family Gigasporaceae. Contrary to expectations, the aphid treatment also increased the evenness of the total fungal community. This suggests that aphids can influence soil communities in conventional arable systems, a result that could have implications for multitrophic feedback loops between crop pests and soil organisms across the above-below-ground interface.

A Plant-Feeding Nematode Indirectly Increases the Fitness of an Aphid.

Plants suffer multiple, simultaneous assaults from above and below ground. In the laboratory, pests and/or pathogen attack are commonly studied on an individual basis. The molecular response of the plant to attack from multiple organisms and the interaction of different defense pathways is unclear. The inducible systemic responses of the potato (Solanum tuberosum L.) host plant were analyzed to characterize the plant-mediated indirect interactions between a sedentary, endoparasitic nematode (Globodera pallida), and a phloem-sucking herbivore (Myzus persicae). The reproductive success of M. persicae was greater on potato plants pre-infected with G. pallida compared to control plants. Salicylic acid (SA) increased systemically in the leaves of potato plants following nematode and aphid infection singly with a corresponding increase in expression of SA-mediated marker genes. An increase in jasmonic acid associated with aphid infection was suppressed when plants were co-infected with nematodes. Our data suggests a positive, asymmetric interaction between a sedentary endoparasitic nematode and a sap-sucking insect. The systemic response of the potato plant following infection with G. pallida indirectly influences the performance of M. persicae. This work reveals additional secondary benefits of controlling individual crop pests.