RSS 1.0Crop Science
Oak Park is home to the National Centre for Arable Crops Research. Situated on 225 hectare research farm, the main objective of the work programme at Oak Park is to support the arable crops sector of Irish agriculture. The task ahead is to adapt new challenges in developing sustainable and environmentally sensitive agricultural systems which will provide farmers with adequate incomes and consumers with quality food at reasonable prices.
Recent Submissions
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Environmental and cultivar variability in composition, content and biological activity of phenolic acids and alkylresorcinols of winter wheat grains from a multi-site field trial across EuropeDifferent factors such as the genotype, environmental conditions, temperature stress, solar radiation and others can influence the phytochemical status of plants. The concentration of phenolic acids and alkylresorciols (ARs) as well as their chemical composition and biological activity have been determined in twelve winter wheat cultivars grown at eight European locations. This was the first winter wheat multi-location field trial of the European Consortium for Open Field Experimentation (ECOFE). Extracts from grain were analyzed using a UPLC-PDA-ESI-MS system (phenolic acids), UPLC-PDA-MS/MS (alkylresorcinols) and TLC-DPPH• test with ImageJ program (antiradical activity). The phenolic acid profile consisted of five hydroxybenzoic acid and four hydroxycinnamic acid derivatives, among which ferulic and sinapic acids were predominated. The ARs profile consisted of nine AR derivatives, among which 5-n-heneicosylresorcinol (C21:0) and 5-n-nonadecanylresorcinol (C19:0) were predominated. Our study showed significant differences in phenolic acids and AR content between wheat cultivars, as well as between locations. We observed a positive correlation between the biological activity of extracts and the total amount of phenolic acids and ARs. Two cultivars, Chambo and Julius (average of all sites) and samples from the Spanish site (average of all cultivars) showed the highest content and composition of nutritional substances.
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A Major Facilitator Superfamily Peptide Transporter From Fusarium oxysporum Influences Bioethanol Production From Lignocellulosic MaterialFusarium oxysporum is a leading microbial agent in the emerging consolidated bioprocessing (CBP) industry owing to its capability to infiltrate the plant’s lignin barrier and degrade complex carbohydrates to value-added chemicals such as bioethanol in a single step. Membrane transport of nutrients is a key factor in successful microbial colonization of host tissue. This study assessed the impact of a peptide transporter on F. oxysporum’s ability to convert lignocellulosic straw to ethanol. We characterized a novel F. oxysporum peptide transporter (FoPTR2) of the dipeptide/tripeptide transporter (PTR) class. FoPTR2 represents a novel transporter with high homology to the Trichoderma sp. peptide transporters ThPTR2 and TrEST-AO793. Its expression level was highly activated in nitrogen-poor environments, which is a characteristic of PTR class peptide transporters. Overexpression and post-translational gene silencing of the FoPTR2 in F. oxysporum affected the peptide transport capacity and ethanol yielded from a both a wheat straw/bran mix and glucose. Thus, we conclude that it FoPTR2 plays a role in the nutrient acquisition system of F. oxysporum which serves to not only enhance fungal fitness but also CBP efficacy.
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Communicating the risks of genetically modified organisms: lessons learnt from an Irish field of cisgenic potatoesAs plant scientists we are all too familiar with the generic commentary that is often associated with the development or use of genetically modified organisms (GMOs) in agricultural systems, but through fact-driven communication, constructive engagement can be achieved. The EU-funded ‘AMIGA’ project, one element of which involved assessing the impact of a GM potato (previously engineered for late blight resistance using cisgenics) with field trials in Ireland, provides a valuable case study in how this can come about. The experiences detailed highlight important lessons learnt relating to the presentation of scientific evidence in a non-scientific format and the necessity for greater integration of biological and social sciences to support the participation of biological researchers in public engagement exercises.
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Integrating a newly developed BAC-based physical mapping resource for Lolium perenne with a genome-wide association study across a L. perenne European ecotype collection identifies genomic contexts associated with agriculturally important traitsBackground and Aims Lolium perenne (perennial ryegrass) is the most widely cultivated forage and amenity grass species in temperate areas worldwide and there is a need to understand the genetic architectures of key agricultural traits and crop characteristics that deliver wider environmental services. Our aim was to identify genomic regions associated with agriculturally important traits by integrating a bacterial artificial chromosome (BAC)-based physical map with a genome-wide association study (GWAS). Methods BAC-based physical maps for L. perenne were constructed from ~212 000 high-information-content fingerprints using Fingerprint Contig and Linear Topology Contig software. BAC clones were associated with both BAC-end sequences and a partial minimum tiling path sequence. A panel of 716 L. perenne diploid genotypes from 90 European accessions was assessed in the field over 2 years, and genotyped using a Lolium Infinium SNP array. The GWAS was carried out using a linear mixed model implemented in TASSEL, and extended genomic regions associated with significant markers were identified through integration with the physical map. Key Results Between ~3600 and 7500 physical map contigs were derived, depending on the software and probability thresholds used, and integrated with ~35 k sequenced BAC clones to develop a resource predicted to span the majority of the L. perenne genome. From the GWAS, eight different loci were significantly associated with heading date, plant width, plant biomass and water-soluble carbohydrate accumulation, seven of which could be associated with physical map contigs. This allowed the identification of a number of candidate genes. Conclusions Combining the physical mapping resource with the GWAS has allowed us to extend the search for candidate genes across larger regions of the L. perenne genome and identified a number of interesting gene model annotations. These physical maps will aid in validating future sequence-based assemblies of the L. perenne genome.
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Geography is essential for reproductive isolation between florally diversified morning glory species from Amazon canga savannahsThe variety, relative importance and eco-evolutionary stability of reproductive barriers are critical to understanding the processes of speciation and species persistence. Here we evaluated the strength of the biotic prezygotic and postzygotic isolation barriers between closely related morning glory species from Amazon canga savannahs. The flower geometry and flower visitor assemblage analyses supported pollination by the bees in lavender-flowered Ipomoea marabaensis and recruitment of hummingbirds as pollinators in red-flowered Ipomoea cavalcantei. Nevertheless, native bee species and alien honeybees foraged on flowers of both species. Real-time interspecific hybridization underscored functionality of the overlap in flower visitor assemblages, questioning the strength of prezygotic isolation underpinned by diversification in flower colour and geometry. Interspecific hybrids were fertile and produced offspring in nature. No significant asymmetry in interspecific hybridization and hybrid incompatibilities among offspring were found, indicating weak postmating and postzygotic isolation. The results suggested that despite floral diversification, the insular-type geographic isolation remains a major barrier to gene flow. Findings set a framework for the future analysis of contemporary evolution of plant-pollinator networks at the population, community, and ecosystem levels in tropical ecosystems that are known to be distinct from the more familiar temperate climate models.
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Growth under cold conditions in a wide perennial ryegrass panel is under tight physiological controlBackground. Perennial ryegrass is a cool-season grass species from the family Poaceae and is widely cultivated in temperate regions because it exhibits rapid growth and establishment, and possesses high forage quality. The extension of the growing season in Ireland in spring and autumn is a breeding target to make farming more profitable since a grass-fed diet based on grazing is the cheapest way of nutrition for ruminants. Methods. Fifty-seven perennial ryegrass accessions were screened for their ability to grow under typical Irish spring conditions as taken from long term temperature records in controlled climate chambers. They were grown in low temperature (8 ◦C/2 ◦C day/night) and control conditions (15 ◦C/8 ◦C day/night) in three consecutive independent experiments. Fresh weight, height, chlorophyll content and electrolyte leakage were measured, and these parameters were used to rank plant performance under low temperature growth conditions. Results. The results showed that height, yield and electrolyte leakage are excellent measures for the impact of cold stress tolerance. Little variation in growth was seen under cold stress, but a wide variety of responses were observed under control conditions. Discussion. Our results suggest that cold stress is under tight physiological control. Interestingly, the various genotypes responded differentially to more amenable control conditions, indicating that a quick response to more amenable growth conditions is a better target for breeding programmes.
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Optimized Use of Low-Depth Genotyping-by-Sequencing for Genomic Prediction Among Multi-Parental Family Pools and Single Plants in Perennial Ryegrass (Lolium perenne L.)Ryegrass single plants, bi-parental family pools, and multi-parental family pools are often genotyped, based on allele-frequencies using genotyping-by-sequencing (GBS) assays. GBS assays can be performed at low-coverage depth to reduce costs. However, reducing the coverage depth leads to a higher proportion of missing data, and leads to a reduction in accuracy when identifying the allele-frequency at each locus. As a consequence of the latter, genomic relationship matrices (GRMs) will be biased. This bias in GRMs affects variance estimates and the accuracy of GBLUP for genomic prediction (GBLUP-GP). We derived equations that describe the bias from low-coverage sequencing as an effect of binomial sampling of sequence reads, and allowed for any ploidy level of the sample considered. This allowed us to combine individual and pool genotypes in one GRM, treating pool-genotypes as a polyploid genotype, equal to the total ploidy-level of the parents of the pool. Using simulated data, we verified the magnitude of the GRM bias at different coverage depths for three different kinds of ryegrass breeding material: individual genotypes from single plants, pool-genotypes from F2 families, and pool-genotypes from synthetic varieties. To better handle missing data, we also tested imputation procedures, which are suited for analyzing allele-frequency genomic data. The relative advantages of the bias-correction and the imputation of missing data were evaluated using real data. We examined a large dataset, including single plants, F2 families, and synthetic varieties genotyped in three GBS assays, each with a different coverage depth, and evaluated them for heading date, crown rust resistance, and seed yield. Cross validations were used to test the accuracy using GBLUP approaches, demonstrating the feasibility of predicting among different breeding material. Bias-corrected GRMs proved to increase predictive accuracies when compared with standard approaches to construct GRMs. Among the imputation methods we tested, the random forest method yielded the highest predictive accuracy. The combinations of these two methods resulted in a meaningful increase of predictive ability (up to 0.09). The possibility of predicting across individuals and pools provides new opportunities for improving ryegrass breeding schemes.
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Genomic prediction of starch content and chipping quality in tetraploid potato using genotyping-by-sequencingGenomic prediction models for starch content and chipping quality show promising results, suggesting that genomic selection is a feasible breeding strategy in tetraploid potato. Genomic selection uses genome-wide molecular markers to predict performance of individuals and allows selections in the absence of direct phenotyping. It is regarded as a useful tool to accelerate genetic gain in breeding programs, and is becoming increasingly viable for crops as genotyping costs continue to fall. In this study, we have generated genomic prediction models for starch content and chipping quality in tetraploid potato to facilitate varietal development. Chipping quality was evaluated as the colour of a potato chip after frying following cold induced sweetening. We used genotyping-by-sequencing to genotype 762 offspring, derived from a population generated from biparental crosses of 18 tetraploid parents. Additionally, 74 breeding clones were genotyped, representing a test panel for model validation. We generated genomic prediction models from 171,859 single-nucleotide polymorphisms to calculate genomic estimated breeding values. Cross-validated prediction correlations of 0.56 and 0.73 were obtained within the training population for starch content and chipping quality, respectively, while correlations were lower when predicting performance in the test panel, at 0.30-0.31 and 0.42-0.43, respectively. Predictions in the test panel were slightly improved when including representatives from the test panel in the training population but worsened when preceded by marker selection. Our results suggest that genomic prediction is feasible, however, the extremely high allelic diversity of tetraploid potato necessitates large training populations to efficiently capture the genetic diversity of elite potato germplasm and enable accurate prediction across the entire spectrum of elite potatoes. Nonetheless, our results demonstrate that GS is a promising breeding strategy for tetraploid potato.
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Quantification of In Planta Zymoseptoria tritici Progression Through Different Infection Phases and Related Association with Components of AggressivenessIn planta growth of Zymoseptoria tritici, causal agent of Septoria tritici blotch of wheat, during the infection process has remained an understudied topic due to the long symptomless latent period before the emergence of fruiting bodies. In this study, we attempted to understand the relationship between in planta growth of Z. tritici relative to the primary components of aggressiveness, i.e., latent period and pycnidia coverage in regard to contrasting host resistance. We tested isolates collected from Ireland against the susceptible cultivar Gallant and cultivar Stigg, which has strong partial resistance. A clear isolate−host interaction effect (F = 3.018; P = 0.005, and F = 6.008; P < 0.001) for latent period and pycnidia coverage, respectively, was identified. Furthermore, during the early infection phase of latency from 5 to 11 days postinoculation (dpi), in planta growth rate of fungal biomass was significantly (F = 30.06; P < 0.001) more affected by host resistance than isolate specificity (F = 1.27; P = 0.27), indicating the importance of host resistance in the early infection phase. In planta Z. tritici growth rates in cultivar Gallant spiked between 11 and 16 dpi followed by a continuous fall onward, whereas in cultivar Stigg it was slowly progressive in nature. From correlation and regression analysis, we found that the in planta growth rate preceding the average latent period of cultivar Gallant has more influence on latency duration and pycnidia production. Likewise, correlation between component of aggressiveness and in planta growth rate of pathogen supports our understanding of aggressiveness to be driven by the pathogen’s multiplication capacity within host tissue.
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The occurrence of herbicide-resistant Avena fatua (wild oats) populations to ACCase-inhibiting herbicides in IrelandFollowing growers’ reports of herbicide control problems, populations of 30 wild oats, Avena fatua, were collected from the south-east main arable counties of Ireland in 2016 and investigated for the occurrence and potential for herbicide resistance to acetyl-CoA carboxylase (ACCase) inhibitors pinoxaden, propaquizafop and cycloxydim, as well as acetolactate synthase (ALS) inhibitor mesosulfuron + iodosulfuron. Plant survival ≥20% was considered as the discriminating threshold between resistant and susceptible populations, when plants were treated with full recommended field rates of ACCase/ALS inhibitors. Glasshouse sensitivity screens revealed 2 out of 30 populations were cross-resistant to all three ACCase inhibitors. While three populations were cross-resistant to both pinoxaden and propaquizafop, and additionally, two populations were resistant to propaquizafop only. Different degree of resistance and cross-resistance between resistant populations suggest the involvement of either different point mutations or more than one resistance mechanism. Nevertheless, all populations including the seven ACCase-resistant populations were equally susceptible to ALS inhibitor. An integrated weed management (cultural/non-chemical control tactics and judicious use of herbicides) approach is strongly recommended to minimize the risk of herbicide resistance evolution.
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A note on the early transcriptional response in leaves and root of potato plants to cadmium exposurePotato plants can accumulate a high amount of cadmium (Cd) in the tuber when grown in soils rich in Cd. The molecular mechanisms governing Cd accumulation in the potato plant are poorly understood. Here we performed an RNA-sequencing experiment to identify genes differentially expressed in the leaf and root of potato during early stages of Cd exposure. Results did not identify any significant transcriptional response in leaves under 1 or 5 mg kg−1 Cd after 72 h. However, in the roots we did identify 2,846 genes that were significantly differentially expressed after 72 h between plants grown in 5 mg kg−1 Cd and controls. These included genes involved in photosynthesis and autophagy being up-regulated, and genes involved in intracellular transport being down-regulated. This study is the first report on the transcriptome-wide response of potato to Cd stress, providing insight into the molecular mechanisms involved in the response.
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Experimental comparison of two methods to study barley responses to partial submergenceBackground Crop yield is dependent on climate conditions, which are becoming both more variable and extreme in some areas of the world as a consequence of global climate change. Increased precipitation and flooding events are the cause of important yield losses due to waterlogging or (partial) submergence of crops in the field. Our ability to screen efficiently and quickly for varieties that have increased tolerance to waterlogging or (partial) submergence is important. Barley, a staple crop worldwide, is particularly sensitive to waterlogging. Screening for waterlogging tolerant barley varieties has been ongoing for many years, but methods used to screen vary greatly, from the type of soil used to the time at which the treatment is applied. This variation makes it difficult to cross-compare results. Results Here, we have devised a scoring system to assess barley tolerance to waterlogging and compare two different methods when partial submergence is applied with either water or a starch solution at an early developmental stage, which is particularly sensitive to waterlogging or partial submergence. The use of a starch solution has been previously shown to result in more reducing soil conditions and has been used to screen for waterlogging tolerance. Conclusions Our results show that the two methods provide similar results to qualitatively rank varieties as tolerant or sensitive, while also affecting plants differently, in that application of a starch solution results in stronger and earlier symptoms than applying partial submergence with water.
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Physiological and transcriptional response to drought stress among bioenergy grass Miscanthus speciesBackground Miscanthus is a commercial lignocellulosic biomass crop owing to its high biomass productivity, resilience and photosynthetic capacity at low temperature. These qualities make Miscanthus a particularly good candidate for temperate marginal land, where yields can be limited by insufficient or excessive water supply. Differences in response to water stress have been observed among Miscanthus species, which correlated to origin. In this study, we compared the physiological and molecular responses among Miscanthus species under excessive (flooded) and insufficient (drought) water supply in glasshouse conditions. Results A significant biomass loss was observed under drought conditions in all genotypes. M. x giganteus showed a lower reduction in biomass yield under drought conditions compared to the control than the other species. Under flooded conditions, biomass yield was as good as or better than control conditions in all species. 4389 of the 67,789 genes (6.4%) in the reference genome were differentially expressed during drought among four Miscanthus genotypes from different species. We observed the same biological processes were regulated across Miscanthus species during drought stress despite the DEGs being not similar. Upregulated differentially expressed genes were significantly involved in sucrose and starch metabolism, redox, and water and glycerol homeostasis and channel activity. Multiple copies of the starch metabolic enzymes BAM and waxy GBSS-I were strongly up-regulated in drought stress in all Miscanthus genotypes, and 12 aquaporins (PIP1, PIP2 and NIP2) were also up-regulated in drought stress across genotypes. Conclusions Different phenotypic responses were observed during drought stress among Miscanthus genotypes from different species, supporting differences in genetic adaption. The low number of DEGs and higher biomass yield in flooded conditions supported Miscanthus use in flooded land. The molecular processes regulated during drought were shared among Miscanthus species and consistent with functional categories known to be critical during drought stress in model organisms. However, differences in the regulated genes, likely associated with ploidy and heterosis, highlighted the value of exploring its diversity for breeding.
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In vitro screening of different Pseudomonas fluorescens isolates to study lytic enzyme production and growth inhibition during antagonism of Fusarium oxysporum f. sp. cumini, wilt causing pathogen of cuminLand plants exist in close association with bacterial and fungal microbes, where some associations can be pathogenic and others can be mutualistic/beneficial. One such relation exists between host plant, Cuminum cyminum L. (Cumin) and Fusarium oxysporum f. sp. cumini (Foc), the causal pathogen of cumin wilt and Pseudomonas fluorescens (Pf), where Pf acts as a bio-agent for inhibiting Foc and promoting plant growth of cumin. In this study, antagonism by 10 different Pf isolates against Foc was studied under laboratory conditions through percent growth inhibition and biochemical mechanisms. Among these Pf isolates, Pf-5 exhibited the highest in vitro growth inhibition (82.51%). A positive correlation was observed between percent growth inhibition and specific activities of hydrolytic enzymes, chitinase, β-1, 3 glucanase, and protease, where a negative correlation was observed with cell wall degrading enzymes, cellulase and polygalacturonase. To conclude, isolate Pf-5 could be a potential biocontrol agent for Fusarium wilt disease of cumin.
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Plant Sciences and Crop Husbandry: Research Report 1989This report summarizes the research undertaken by Teagasc in the areas of Plant Sciences and Crop Husbandry in 1989
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A Response to the Draft Climate Change Adaptation Sectoral Plan for Agriculture, Forest and Seafood SectorTeagasc is pleased to have the opportunity to contribute to this Draft Climate Change Adaptation Sectoral Plan for Agriculture, Forest and Seafood Sectors, although our contribution will largely be limited to the agriculture and forestry sectors. We have also taken the liberty to contribute in the form of ‘submissions, observations and comments’ as indicated in the call for contributions rather than in the formal questionnaire which appears to be more appropriate for an individual submission rather than an organisational contribution.
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The Double Spade Method: a ‘mini-profile’ visual soil evaluation techniqueVisual Soil Evaluation (VSE) methods are established for soil quality assessment and focus on the examination of soil structure and associated anthropogenic impacts. VSE techniques, of which numerous types exist, are successfully used internationally both in soil research and as sustainable soil management tools. Techniques are generally categorised into profile and spade methods. Profile methods examine entire soil profiles in soil-pits to depths of ~ 1.5 m, exploring interactions between inherent soil features and anthropic management at specific sample points. Spade methods examine the upper soil profile, often by extracting sample blocks of topsoil by spade and focus on anthropic impacts. The VESS method (Guimarães et al., 2011) is a widely used spade method and involves assessment of soil sample blocks to 25 cm depth. However, in arable soils, important structural features may occur just below this depth such as plough pans, which VESS may not capture. The SubVESS method (Ball et al., 2015) follows principles of VESS but allows assessment to ~ 1 m depth. However, the later involves soil-pit excavation by mechanical means, which may be destructive, costly, time consuming and limit replication. When used in on-farm situations by farmers or advisors, full soil-pit excavation may not be desirable. Here we describe a method previously outlined (Emmet-Booth et al. 2018) called the Double Spade Method (DS) designed to examine miniprofiles in soil pits to 40 cm depth, therefore capturing potential structural features below the VESS assessment depth, without requiring full soil-pit excavation.
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Beyond ruminants: discussing opportunities for alternative pasture uses in New ZealandThe New Zealand government has set ambitious goals for primary sector growth and of zero net carbon emissions by 2050. This presents an opportunity and obligation to develop new ideas for grassland production systems to increase export value and generate new job opportunities, while reducing environmental impacts. The aim of this paper is to draw on recent research in Europe to investigate some of the alternative and complementary uses for pasture as a feedstock for a green biorefinery. A biorefinery is a facility, or a series of processes, that convert biomass into a spectrum of value-added products. For example, protein can be extracted mechanically from green biomass once harvested. The residual fibre fraction could be used as a low-nitrogen feed for ruminants to reduce urinary nitrogen, while the liquid protein fraction could be processed to make it suitable for mono-gastric or human consumption. Enzymes can promote protein extraction and controlled conversion of insoluble plant fibres and oligosaccharides to foster gut-health promoting prebiotic food ingredients. Anaerobic digestion of residues can then be used to create energy and soilimproving products. Research and demonstration of these approaches in practice, along with the results of feasibility studies, will be required to see which of these opportunities is a good fit for New Zealand pasture systems.
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Parthenium hysterophorus Herbage Mulching: a Potential Source of Weeds Control in Soybean (Glycine max)Weeds have indirect effects on crop plants. Crop development is affected by allelopathy from certain weed species. Allelochemicals from allelopathic weeds can disturb the root and shoot growth of emerging crop seedlings, as well as cause several other types of damage. A study was carried out to investigate the allelopathic potential of Parthenium hysterophorus for weed response in soybean. The experiment was laid out in Randomized Complete Block Design (RCBD) with split plot arrangements and replicated thrice. Sowing methods (broadcast and line sowing) were kept in the main plot and mulching treatments (surface mulching and soil incorporation) were kept in the sub-plots. Mulching of Parthenium hysterophorus was applied at the rate of 1.0 t ha-1, 2.5 t ha-1, 5 t ha-1 with control (no parthenium). Manual weed control was also used as treatments. The results revealed that significantly higher shoot length, shoot fresh weight, shoot dry weight, root length, root fresh weigh, root dry weight, number of nodules per plant, nodules fresh and dry weight, number of branches, number of pods per plant, thousand seed weight biological yield, economic yield, dry matter yield and harvest index were recorded with the soil incorporation of Parthenium herbage at the rate of 2.5 t ha-1. Maximum weed density and weed dry biomass were recorded in control plots while weed control efficiency was seen greater in plots where Parthenium herbage was applied to surface at the rate of 5 t ha-1. The results suggested that the use of Parthenium hysterophorus herbage mulching can reduce infestation of weeds by its allelopathic effects and increase the yield of soybean under sub-humid agro-climatic conditions.
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Grain aphids (Sitobion avenae) with knockdown resistance (kdr) to insecticide exhibit fitness trade-offs, including increased vulnerability to the natural enemy Aphidius erviThe development of insecticide-resistance mechanisms in aphids has been associated with inhibitory, pleiotropic fitness costs. Such fitness costs have not yet been examined in the UK’s most damaging cereal aphid, Sitobion avenae (grain aphid) (Hemiptera: Aphididae). This study aimed to evaluate the fitness trade-offs of the insecticide-resistant S. avenae clone versus an insecticide-susceptible S. avenae clone. Additionally, the parasitoid, Aphidius ervi (Hymenoptera: Braconidae), was introduced to examine its potential as a biological control agent. This study found that insecticide-resistant clones had significantly lower population growth and individual relative growth rate. Furthermore, insecticide-resistant clones suffered from a significantly greater rate of parasitisation (mummification) compared to their insecticide-susceptible counterparts. The successfulness of the parasitoid as a biological control agent could prevent the spread of the insecticide-resistant genotype. However, for this to be possible, insecticide spraying regimes need to be moderated, and habitat modification and parasitoid manipulation must be considered.
