Associational Effects of Desmodium Intercropping on Maize Resistance and Secondary Metabolism.

Intercropping is drawing increasing attention as a strategy to increase crop yields and manage pest pressure, however the mechanisms of associational resistance in diversified cropping systems remain controversial. We conducted a controlled experiment to assess the impact of co-planting with silverleaf Desmodium (Desmodium uncinatum) on maize secondary metabolism and resistance to herbivory by the spotted stemborer (Chilo partellus). Maize plants were grown either in the same pot with a Desmodium plant or adjacent to it in a separate pot. Our findings indicate that co-planting with Desmodium influences maize secondary metabolism and herbivore resistance through both above and below-ground mechanisms. Maize growing in the same pot with a Desmodium neighbor was less attractive for oviposition by spotted stemborer adults. However, maize exposed only to above-ground Desmodium cues generally showed increased susceptibility to spotted stemborer herbivory (through both increased oviposition and larval consumption). VOC emissions and tissue secondary metabolite titers were also altered in maize plants exposed to Desmodium cues, with stronger effects being observed when maize and Desmodium shared the same pot. Specifically, benzoxazinoids were strongly suppressed in maize roots by direct contact with a Desmodium neighbor while headspace emissions of short-chain aldehydes and alkylbenzenes were increased. These results imply that direct root contact or soil-borne cues play an important role in mediating associational effects on plant resistance in this system.

Molasses Grass Induces Direct and Indirect Defense Responses in Neighbouring Maize Plants.

Plants have evolved intricate defence strategies against herbivore attack which can include activation of defence in response to stress-related volatile organic compounds (VOCs) emitted by neighbouring plants. VOCs released by intact molasses grass (Melinis minutiflora), have been shown to repel stemborer, Chilo partellus (Swinhoe), from maize and enhance parasitism by Cotesia sesamiae (Cameron). In this study, we tested whether the molasses grass VOCs have a role in plant-plant communication by exposing different maize cultivars to molasses grass for a 3-week induction period and then observing insect responses to the exposed plants. In bioassays, C. partellus preferred non-exposed maize landrace plants for egg deposition to those exposed to molasses grass. Conversely, C. sesamiae parasitoid wasps preferred volatiles from molasses grass exposed maize landraces compared to volatiles from unexposed control plants. Interestingly, the molasses grass induced defence responses were not observed on hybrid maize varieties tested, suggesting that the effect was not simply due to absorption and re-emission of VOCs. Chemical and electrophysiological analyses revealed strong induction of bioactive compounds such as (R)-linalool, (E)-4,8-dimethyl-1,3,7-nonatriene and (E,E)-4,8,12-trimethyl-1,3,7,11-tridecatetraene from maize landraces exposed to molasses grass volatiles. Our results suggest that constitutively emitted molasses grass VOCs can induce direct and indirect defence responses in neighbouring maize landraces. Plants activating defences by VOC exposure alone could realize enhanced levels of resistance and fitness compared to those that launch defence responses upon herbivore attack. Opportunities for exploiting plant-plant signalling to develop ecologically sustainable crop protection strategies against devastating insect pests such as stemborer C. partellus are discussed.

Agro-ecological options for fall armyworm (Spodoptera frugiperda JE Smith) management: Providing low-cost, smallholder friendly solutions to an invasive pest.

Fall armyworm (FAW), a voracious agricultural pest native to North and South America, was first detected on the African continent in 2016 and has subsequently spread throughout the continent and across Asia. It has been predicted that FAW could cause up to $US13 billion per annum in crop losses throughout sub-Saharan Africa, thereby threatening the livelihoods of millions of poor farmers. In their haste to respond to FAW governments may promote indiscriminate use of chemical pesticides which, aside from human health and environmental risks, could undermine smallholder pest management strategies that depend to a large degree on natural enemies. Agro-ecological approaches offer culturally appropriate low-cost pest control strategies that can be readily integrated into existing efforts to improve smallholder incomes and resilience through sustainable intensification. Such approaches should therefore be promoted as a core component of integrated pest management (IPM) programmes for FAW in combination with crop breeding for pest resistance, classical biological control and selective use of safe pesticides. Nonetheless, the suitability of agro-ecological measures for reducing FAW densities and impact need to be carefully assessed across varied environmental and socio-economic conditions before they can be proposed for wide-scale implementation. To support this process, we review evidence for the efficacy of potential agro-ecological measures for controlling FAW and other pests, consider the associated risks, and draw attention to critical knowledge gaps. The evidence indicates that several measures can be adopted immediately. These include (i) sustainable soil fertility management, especially measures that maintain or restore soil organic carbon; (ii) intercropping with appropriately selected companion plants; and (iii) diversifying the farm environment through management of (semi)natural habitats at multiple spatial scales. Nevertheless, we recommend embedding trials into upscaling programmes so that the costs and benefits of these interventions may be determined across the diverse biophysical and socio-economic contexts that are found in the invaded range.

Genetic variation and host-parasite specificity of Striga resistance and tolerance in rice: the need for predictive breeding.

The parasitic weeds Striga asiatica and Striga hermonthica cause devastating yield losses to upland rice in Africa. Little is known about genetic variation in host resistance and tolerance across rice genotypes, in relation to virulence differences across Striga species and ecotypes. Diverse rice genotypes were phenotyped for the above traits in S. asiatica- (Tanzania) and S. hermonthica-infested fields (Kenya and Uganda) and under controlled conditions. New rice genotypes with either ecotype-specific or broad-spectrum resistance were identified. Resistance identified in the field was confirmed under controlled conditions, providing evidence that resistance was largely genetically determined. Striga-resistant genotypes contributed to yield security under Striga-infested conditions, although grain yield was also determined by the genotype-specific yield potential and tolerance. Tolerance, the physiological mechanism mitigating Striga effects on host growth and physiology, was unrelated to resistance, implying that any combination of high, medium or low levels of these traits can be found across rice genotypes. Striga virulence varies across species and ecotypes. The extent of Striga-induced host damage results from the interaction between parasite virulence and genetically determined levels of host-plant resistance and tolerance. These novel findings support the need for predictive breeding strategies based on knowledge of host resistance and parasite virulence.

Development of field-applicable tests for rapid and sensitive detection of Candidatus Phytoplasma oryzae.

Napier grass Stunt Disease (NSD) is a severe disease of Napier grass (Pennisetum purpureum) in Eastern Africa, caused by the leafhopper-transmitted bacterium Candidatus Phytoplasma oryzae. The pathogen severely impairs the growth of Napier grass, the major fodder for dairy cattle in Eastern Africa. NSD is associated with biomass losses of up to 70% of infected plants. Diagnosis of NSD is done by nested PCR targeting the phytoplasma DNA, which is difficult to perform in developing countries with little infrastructure. We report the development of an easy to use, rapid, sensitive and specific molecular assay for field diagnosis of NSD. The procedure is based on recombinase polymerase amplification and targets the imp gene encoding a pathogen-specific immunodominant membrane protein. Therefore we followed a two-step process. First we developed an isothermal DNA amplification method for real time fluorescence application and then transferred this assay to a lateral flow format. The limit of detection for both procedures was estimated to be 10 organisms. We simplified the template preparation procedure by using freshly squeezed phloem sap from Napier grass. Additionally, we developed a laboratory serological assay with the potential to be converted to a lateral flow assay. Two murine monoclonal antibodies with high affinity and specificity to the immunodominant membrane protein IMP of Candidatus Phytoplasma oryzae were generated. Both antibodies specifically reacted with the denatured or native 17 kDa IMP protein. In dot blot experiments of extracts from infected plant, phytoplasmas were detected in as little as 12,5 µg of fresh plant material.

Drought-tolerant Desmodium species effectively suppress parasitic striga weed and improve cereal grain yields in western Kenya.

The parasitic weed Striga hermonthica Benth. (Orobanchaceae), commonly known as striga, is an increasingly important constraint to cereal production in sub-Saharan Africa (SSA), often resulting in total yield losses in maize (Zea mays L.) and substantial losses in sorghum (Sorghum bicolor (L.) Moench). This is further aggravated by soil degradation and drought conditions that are gradually becoming widespread in SSA. Forage legumes in the genus Desmodium (Fabaceae), mainly D. uncinatum and D. intortum, effectively control striga and improve crop productivity in SSA. However, negative effects of climate change such as drought stress is affecting the functioning of these systems. There is thus a need to identify and characterize new plants possessing the required ecological chemistry to protect crops against the biotic stress of striga under such environmental conditions. 17 accessions comprising 10 species of Desmodium were screened for their drought stress tolerance and ability to suppress striga. Desmodium incanum and D. ramosissimum were selected as the most promising species as they retained their leaves and maintained leaf function for longer periods during their exposure to drought stress conditions. They also had desirable phenotypes with more above ground biomass. The two species suppressed striga infestation, both under controlled and field conditions, and resulted in significant grain yield increases, demonstrating the incremental capability of Desmodium species in striga suppression. These results demonstrate beneficial effects of Desmodium species in enhancing cereal productivity in dry areas.

Push-Pull: Chemical Ecology-Based Integrated Pest Management Technology.

Lepidopterous stemborers, and parasitic striga weeds belonging to the family Orobanchaceae, attack cereal crops in sub-Saharan Africa causing severe yield losses. The smallholder farmers are resource constrained and unable to afford expensive chemicals for crop protection. The push-pull technology, a chemical ecology- based cropping system, is developed for integrated pest and weed management in cereal-livestock farming systems. Appropriate plants were selected that naturally emit signaling chemicals (semiochemicals). Plants highly attractive for stemborer egg laying were selected and employed as trap crops (pull), to draw pests away from the main crop. Plants that repelled stemborer females were selected as intercrops (push). The stemborers are attracted to the trap plant, and are repelled from the main cereal crop using a repellent intercrop (push). Root exudates of leguminous repellent intercrops also effectively control the parasitic striga weed through an allelopathic mechanism. Their root exudates contain flavonoid compounds some of which stimulate germination of Striga hermonthica seeds, such as Uncinanone B, and others that dramatically inhibit their attachment to host roots, such as Uncinanone C and a number of di-C-glycosylflavones (di-CGFs), resulting in suicidal germination. The intercrop also improves soil fertility through nitrogen fixation, natural mulching, improved biomass, and control of erosion. Both companion plants provide high value animal fodder, facilitating milk production and diversifying farmers' income sources. The technology is appropriate to smallholder mixed cropping systems in Africa. Adopted by about 125,000 farmers to date in eastern Africa, it effectively addresses major production constraints, significantly increases maize yields, and is economical as it is based on locally available plants, not expensive external inputs.

Detection, Identification, and Significance of Phytoplasmas in Wild Grasses in East Africa.

Plant-pathogenic phytoplasmas found in wild grasses in East Africa could pose a serious threat to the cultivation of Napier grass, Pennisetum purpureum, the most important livestock fodder in the region. To asses this threat, leaves from plants of 33 grass species were sampled from Mbita, Bungoma, and Busia districts in western Kenya; Tarime district in northern Tanzania; and Busia and Bugiri districts in the eastern Uganda to determine which species host phytoplasmas, the identity of the phytoplasmas, and their relationship with disease symptoms. Phytoplasmas were detected using universal primers based on conserved phytoplasma-specific 16S rDNA sequences from 11 grass species collected. Sequence and phylogenetic analysis revealed the presence of Napier grass stunt-related phytoplasmas in 11 grass species, 'Candidatus Phytoplasma cynodontis' in three, and goosegrass white leaf phytoplasma in 2 wild grass species. This study showed that the geographical distribution, diversity of phytoplasmas, and their grass host species in East Africa is greater than antecedently thought and that typical disease symptoms, including white leaf or stunting alone, are not reliable indicators of the presence of phytoplasma. It also shows the need to identify insect vectors responsible for phytoplasma transmission from native grasses to Napier grass or other cereals present in the region.

The biosynthesis of allelopathic di-C-glycosylflavones from the roots of Desmodium incanum (G. Mey.) DC.

The allelopathic root exudate of the drought-tolerant subsistence cereal intercrop D. incanum, protecting against the parasitic weed Striga hermonthica, comprises a number of di-C-glycosylflavones specifically containing C-glucosyl, C-galactosyl and C-arabinosyl moieties. Here we demonstrate that the biosynthesis of all compounds containing a C-glucose involves C-glucosylation of 2-hydroxynaringenin with subsequent C-galactosylation, C-glucosylation or C-arabinosylation. In addition, the crude soluble enzyme extract converts two fluorinated 2-hydroxyflavanone analogues to corresponding mono- and di-C-glycosylflavones demonstrating that some differences in C-ring substitution can be tolerated by the plant enzymes. Elucidating the biosynthesis of these C-glycosylflavones (CGFs) has the potential to open up opportunities for transferring the enzymic and genetic basis for the S. hermonthica inhibiting allelopathic trait to food crop plants.

Responses of parasitoids to volatiles induced by Chilo partellus oviposition on teosinte, a wild ancestor of maize.

Maize, a genetically diverse crop, is the domesticated descendent of its wild ancestor, teosinte. Recently, we have shown that certain maize landraces possess a valuable indirect defense trait not present in commercial hybrids. Plants of these landraces release herbivore-induced plant volatiles (HIPVs) that attract both egg [Trichogramma bournieri Pintureau & Babault (Hymenoptera: Trichogrammatidae)] and larval [Cotesia sesamiae Cameron (Hymenoptera: Braconidae)] parasitoids in response to stemborer egg deposition. In this study, we tested whether this trait also exists in the germplasm of wild Zea species. Headspace samples were collected from plants exposed to egg deposition by Chilo partellus Swinhoe (Lepidoptera: Crambidae) moths and unexposed control plants. Four-arm olfactometer bioassays with parasitic wasps, T. bournieri and C. sesamiae, indicated that both egg and larval parasitoids preferred HIPVs from plants with eggs in four of the five teosinte species sampled. Headspace samples from oviposited plants released higher amounts of EAG-active compounds such as (E)-4,8-dimethyl-1,3,7-nonatriene. In oviposition choice bioassays, plants without eggs were significantly preferred for subsequent oviposition by moths compared to plants with prior oviposition. These results suggest that this induced indirect defence trait is not limited to landraces but occurs in wild Zea species and appears to be an ancestral trait. Hence, these species possess a valuable trait that could be introgressed into domesticated maize lines to provide indirect defense mechanisms against stemborers.

Ecological management of cereal stemborers in African smallholder agriculture through behavioural manipulation.

1. Africa faces serious challenges in feeding its rapidly growing human population owing to the poor productivity of maize and sorghum, the most important staple crops for millions of smallholder farmers in the continent, with yields being among the lowest in the world. 2. A complex of lepidopterous stemborers attack cereals in Africa. However, their effective control is difficult, largely as a result of the cryptic and nocturnal habits of moths, and protection provided by host stem for immature pest stages. Moreover, current control measures are uneconomical and impractical for resource-poor farmers. 3. An ecological approach, based on companion planting, known as 'push-pull', provides effective management of these pests, and involves combined use of inter- and trap cropping systems where stemborers are attracted and trapped on trap plants with added economic value ('pull'), and are driven away from the cereal crop by antagonistic intercrops ('push'). 4. Novel defence strategies inducible by stemborer oviposition have recently been discovered, leading to the attraction of egg and larval parasitoids, in locally adapted maize lines but not in elite hybrids. We also established that landscape complexity did not improve the ecosystem service of biological control, but rather provided a disservice by acting as a 'source' of stemborer pests colonising the crop. 5. Here we review and provide new data on the direct and indirect effects of the push-pull approach on stemborers and their natural enemies, including the mechanisms involved, and highlight opportunities for exploiting intrinsic plant defences and natural ecosystem services in pest management in smallholder farming systems in Africa.

Do NERICA rice cultivars express resistance to Striga hermonthica (Del.) Benth. and Striga asiatica (L.) Kuntze under field conditions?

The parasitic weeds Striga asiatica and Striga hermonthica cause high yield losses in rain-fed upland rice in Africa. Two resistance classes (pre- and post-attachment) and several resistant genotypes have been identified among NERICA (New Rice for Africa) cultivars under laboratory conditions (in vitro) previously. However, little is known about expression of this resistance under field conditions. Here we investigated (1) whether resistance exhibited under controlled conditions would express under representative Striga-infested field conditions, and (2) whether NERICA cultivars would achieve relatively good grain yields under Striga-infested conditions. Twenty-five rice cultivars, including all 18 upland NERICA cultivars, were screened in S. asiatica-infested (in Tanzania) and S. hermonthica-infested (in Kenya) fields during two seasons. Additionally, a selection of cultivars was tested in vitro, in mini-rhizotron systems. For the first time, resistance observed under controlled conditions was confirmed in the field for NERICA-2, -5, -10 and -17 (against S. asiatica) and NERICA-1 to -5, -10, -12, -13 and -17 (against S. hermonthica). Despite high Striga-infestation levels, yields of around 1.8 t ha-1 were obtained with NERICA-1, -9 and -10 (in the S. asiatica-infested field) and around 1.4 t ha-1 with NERICA-3, -4, -8, -12 and -13 (in the S. hermonthica-infested field). In addition, potential levels of tolerance were identified in vitro, in NERICA-1, -17 and -9 (S. asiatica) and in NERICA-1, -17 and -10 (S. hermonthica). These findings are highly relevant to rice agronomists and breeders and molecular geneticists working on Striga resistance. In addition, cultivars combining broad-spectrum resistance with good grain yields in Striga-infested fields can be recommended to rice farmers in Striga-prone areas.

Oviposition induced volatile emissions from African smallholder farmers' maize varieties.

Maize (corn), Zea mays, is a genetically diverse crop, and we have recently shown that certain open pollinated varieties (OPVs) of Latin American origin possess a trait not present in mainstream commercial varieties: they produce volatiles in response to stemborer oviposition that are attractive to stemborer parasitoids. Here, we tested whether a similar tritrophic effect occurs in the African OPVs 'Nyamula' and 'Jowi'. Herbivore induced plant volatiles (HIPVs) were collected from plants exposed to egg deposition by the stemborer Chilo partellus. In a four-arm olfactometer bioassay, the parasitic wasp Cotesia sesamiae preferred samples containing HIPVs from plants with eggs to samples collected from plants without eggs. EAG-active compounds, including (E)-4,8-dimethyl-1,3,7-nonatriene (DMNT), were released in higher amounts from the egg induced headspace samples. Our results suggest that this oviposition trait is not limited to S. American Z. mays germplasm, and that it could be used to increase indirect defense against attack by stemborers.

Maize landraces recruit egg and larval parasitoids in response to egg deposition by a herbivore.

Natural enemies respond to herbivore-induced plant volatiles (HIPVs), but an often overlooked aspect is that there may be genotypic variation in these 'indirect' plant defence traits within plant species. We found that egg deposition by stemborer moths (Chilo partellus) on maize landrace varieties caused emission of HIPVs that attract parasitic wasps. Notably, however, the oviposition-induced release of parasitoid attractants was completely absent in commercial hybrid maize varieties. In the landraces, not only were egg parasitoids (Trichogramma bournieri) attracted but also larval parasitoids (Cotesia sesamiae). This implies a sophisticated defence strategy whereby parasitoids are recruited in anticipation of egg hatching. The effect was systemic and caused by an elicitor, which could be extracted from egg materials associated with attachment to leaves. Our findings suggest that indirect plant defence traits may have become lost during crop breeding and could be valuable in new resistance breeding for sustainable agriculture.

The role of Desmodium intortum, Brachiaria sp. and Phaseolus vulgaris in the management of fall armyworm Spodoptera frugiperda (J. E. Smith) in maize cropping systems in Africa.

BACKGROUND: The fall armyworm (FAW), Spodoptera frugiperda (J.E. Smith) is a serious pest of maize. Farming systems such as push-pull or maize-legume intercropping have been reported to reduce FAW infestations significantly. However, the exact mechanisms involved in FAW management have not been practically elucidated. We therefore assessed larval host preference, feeding and survival rate when exposed to four host plants commonly used in push-pull and legume intercropping. We also compared adult moths' oviposition preference between maize and other grasses used as trap crops in push-pull. RESULTS: The larval orientation and settlement study showed that maize was the most preferred host plant followed by bean, desmodium and Brachiaria brizantha cv Mulato II. The larval arrest and dispersal experiment showed that mean number of larvae was significantly higher on maize than on Desmodium or B. brizantha cv Mulato II. However, no significant differences were found between maize and bean after 24 h. Maize was the most consumed plant, followed by bean, desmodium and finally brachiaria. The mean percentage of survival to the pupation stage was significantly higher on maize. The study on FAW oviposition preference showed no significant differences in egg deposited between maize and other grasses. However, B. brizantha cv Xaraes, which received more eggs than maize, could be a promising alternative to B. brizantha cv Mulato II for the control of FAW. CONCLUSION: The study provides a better understanding of the mechanisms involved in the control of fall armyworm under the push-pull and maize legume intercropping. © 2021 The Authors. Pest Management Science published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.

Managing Fall Armyworm in Africa: Can Bt Maize Sustainably Improve Control?

The recent invasion of Africa by fall armyworm, Spodoptera frugiperda, a lepidopteran pest of maize and other crops, has heightened concerns about food security for millions of smallholder farmers. Maize genetically engineered to produce insecticidal proteins from the bacterium Bacillus thuringiensis (Bt) is a potentially useful tool for controlling fall armyworm and other lepidopteran pests of maize in Africa. In the Americas, however, fall armyworm rapidly evolved practical resistance to maize producing one Bt toxin (Cry1Ab or Cry1Fa). Also, aside from South Africa, Bt maize has not been approved for cultivation in Africa, where stakeholders in each nation will make decisions about its deployment. In the context of Africa, we address maize production and use; fall armyworm distribution, host range, and impact; fall armyworm control tactics other than Bt maize; and strategies to make Bt maize more sustainable and accessible to smallholders. We recommend mandated refuges of non-Bt maize or other non-Bt host plants of at least 50% of total maize hectares for single-toxin Bt maize and 20% for Bt maize producing two or more distinct toxins that are each highly effective against fall armyworm. The smallholder practices of planting more than one maize cultivar and intercropping maize with other fall armyworm host plants could facilitate compliance. We also propose creating and providing smallholder farmers access to Bt maize that produces four distinct Bt toxins encoded by linked genes in a single transgene cassette. Using this novel Bt maize as one component of integrated pest management could sustainably improve control of lepidopteran pests including fall armyworm.

Evaluation of African Maize Cultivars for Resistance to Fall Armyworm Spodoptera frugiperda (J. E. Smith) (Lepidoptera: Noctuidae) Larvae.

The fall armyworm (FAW) has recently invaded and become an important pest of maize in Africa causing yield losses reaching up to a third of maize annual production. The present study evaluated different aspects of resistance of six maize cultivars, cropped by farmers in Kenya, to FAW larvae feeding under laboratory and field conditions. We assessed the arrestment and feeding of FAW neonate larvae in no-choice and choice experiments, development of larvae-pupae, food assimilation under laboratory conditions and plant damage in a field experiment. We did not find complete resistance to FAW feeding in the evaluated maize cultivars, but we detected differences in acceptance and preference when FAW larvae were given a choice between certain cultivars. Moreover, the smallest pupal weight and the lowest growth index were found on 'SC Duma 43' leaves, which suggests an effect of antibiosis of this maize hybrid against FAW larvae. In contrast, the highest growth index was recorded on 'Rachar' and the greatest pupal weight was found on 'Nyamula' and 'Rachar'. The density of trichomes on the leaves of these maize cultivars seems not to be directly related to the preference of neonates for feeding. Plant damage scores were not statistically different between cultivars in the field neither under natural nor artificial infestation. However, plant damage scores in 'Nyamula' and 'Jowi' tended to be lower in the two last samplings of the season compared to the two initial samplings under artificial infestation. Our study provides insight into FAW larval preferences and performance on some African maize cultivars, showing that there are differences between cultivars in these variables; but high levels of resistance to larvae feeding were not found.

Exploiting phytochemicals for developing a 'push-pull' crop protection strategy for cereal farmers in Africa.

Lepidopteran stemborers and parasitic weeds in the genus Striga are major constraints to efficient production of cereals, the most important staple food crops in Africa. Smallholder farmers are resource constrained and unable to afford expensive chemicals for crop protection. Development of a push-pull approach for integrated pest and weed management is reviewed here. Appropriate plants were discovered that naturally emit signalling chemicals (semiochemicals). Plants highly attractive for egg laying by stemborer pests were selected and employed as trap crops (pull), to draw pests away from the main crop. Of these, Napier grass, Pennisetum purpureum (Schumach), despite its attractiveness, supported minimal survival of the pests' immature stages. Plants that repelled stemborer pests, notably molasses grass, Melinis minutiflora P. Beauv., and forage legumes in the genus Desmodium, were selected as intercrops (push). Desmodium intercrops suppress Striga hermonthica (Del.) Benth. through an allelopathic mechanism. Their root exudates contain novel flavonoid compounds, which stimulate suicidal germination of S. hermonthica seeds and dramatically inhibit its attachment to host roots. The companion crops provide valuable forage for farm animals while the leguminous intercrops also improve soil fertility and moisture retention. The system is appropriate as it is based on locally available plants, not expensive external inputs, and fits well with traditional mixed cropping systems in Africa. To date it has been adopted by more than 30,000 smallholder farmers in East Africa where maize yields have increased from ∼1 t ha(-1) to 3.5 t ha(-1). Future directions for semiochemical delivery by plants including biotechnological opportunities are discussed.

Is quality more important than quantity? Insect behavioural responses to changes in a volatile blend after stemborer oviposition on an African grass.

Plants subjected to insect attack usually increase volatile emission which attracts natural enemies and repels further herbivore colonization. Less is known about the capacity of herbivores to suppress volatiles and the multitrophic consequences thereof. In our study, the African forage grass, Brachiaria brizantha, was exposed to ovipositing spotted stemborer, Chilo partellus, moths. A marked reduction in emission of the main volatile, (Z)-3-hexenyl acetate (Z3HA), occurred following oviposition but the ratio of certain other minor component volatiles to Z3HA was increased. While further herbivore colonization was reduced on plants after oviposition, the new volatile profile caused increased attraction of an adapted parasitoid, Cotesia sesamiae. Our results show that insect responses are dependent on the quality of volatile emission rather than merely the quantity in this multitrophic interaction.

Genome wide association analysis of a stemborer egg induced "call-for-help" defence trait in maize.

Tritrophic interactions allow plants to recruit natural enemies for protection against herbivory. Here we investigated genetic variability in induced responses to stemborer egg-laying in maize Zea mays (L.) (Poaceae). We conducted a genome wide association study (GWAS) of 146 maize genotypes comprising of landraces, inbred lines and commercial hybrids. Plants were phenotyped in bioassays measuring parasitic wasp Cotesia sesamiae (Cameron) (Hymenoptera: Braconidae) attraction to volatiles collected from plants exposed to stemborer Chilo partellus (Swinhoe) (Lepidoptera: Crambidae) eggs. Genotyping-by-sequencing was used to generate maize germplasm SNP data for GWAS. The egg-induced parasitoid attraction trait was more common in landraces than in improved inbred lines and hybrids. GWAS identified 101 marker-trait associations (MTAs), some of which were adjacent to genes involved in the JA-defence pathway (opr7, aos1, 2, 3), terpene biosynthesis (fps3, tps2, 3, 4, 5, 7, 9, 10), benzoxazinone synthesis (bx7, 9) and known resistance genes (e.g. maize insect resistance 1, mir1). Intriguingly, there was also association with a transmembrane protein kinase that may function as a receptor for the egg elicitor and other genes implicated in early plant defence signalling. We report maize genomic regions associated with indirect defence and provide a valuable resource for future studies of tritrophic interactions in maize. The markers identified may facilitate selection of indirect defence by maize breeders.