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.

Ecological Chemistry of Pest Control in Push-Pull Intercropping Systems: What We Know, and Where to Go?

Push-pull technology (PPT) employs mixed cropping for sustainable intensification: an intercrop repels or suppresses pests of the focal crop (push), while a trap crop attracts pests out of the field (pull), where they may be targeted for control. Underlying chemical-ecological mechanisms have been demonstrated in controlled settings, primarily for some of the best-established cereal PPT systems developed in east Africa. Yet, many questions remain regarding mechanisms, and strategies to adapt PPT for different crops and locations. We conducted a systematic review of scientific literature on PPT and related practices for biological control of pests of food and fodder. Of 3335 results, we identified 45 reporting on chemistry of trap- or intercropping systems for pest control, of which 30 focused on cereals or African pests. Seven of these reported primary chemical data: measurements from glasshouse and laboratory studies (5), or of field-collected samples (2). From these 30, we provide a database of compounds, discussing degrees of evidence for their mediation of push-pull. We depict hypothesized spatial distributions of selected compounds in PPT fields from physical properties and emission/exudation rates, and design of the east African cereal PPT system, and discuss influences on activity in field settings likely to affect success.

Potential roles of selected forage grasses in management of fall armyworm (Spodoptera frugiperda) through companion cropping.

Production of maize, Zea mays L. (Poaceae), in sub-Saharan Africa is threatened by a new invasive pest, fall armyworm (FAW), Spodoptera frugiperda (JE Smith) (Lepidoptera: Noctuidae). To mitigate this threat, push-pull companion cropping, a system originally developed for management of lepidopteran stemborers, may be used to control FAW. The original system involved trap crops that functioned as a 'pull' component to attract moths away from the main crop. How grass species can be used as trap crops in a push-pull system to control FAW is a question that remains to be answered, because maize is already a highly preferred host plant. Therefore, we tested oviposition preference of FAW female moths in no-choice and two-choice experiments and larval performance on six selected grasses (Poaceae) to assess their roles as trap crop 'pull' plants in the system. In no-choice tests, numbers of eggs deposited on Brachiaria brizantha (Hochst. ex A. Rich.) R. Webster cv. 'Piata', cv. 'Mulato II', and cv. 'Xaraes', and Napier grass (Pennisetum purpureum K. Schumach) cv. 'South Africa' were not statistically different from those deposited on maize. In two-choice tests between grasses and maize, there were no significant differences in number of eggs laid when the plants were of the same size. However, in two-choice tests with maize plants half of the size of the grasses, significantly more eggs were laid on B. brizantha cv. Xaraes and P. purpureum cv. South Africa than on maize, suggesting that crop phenology could make a difference. Numbers of larvae arrested on grass leaf cuts were considerably lower than those on maize leaf cuts after 48 h. In two-choice tests with maize, molasses grass (Melinis minutiflora P. Beauv.) was the only grass that was significantly preferred to maize for larval settlement after 24 h. After 48 h in the two-choice test, it was the only grass that retained larvae, although the larval count was significantly lower than on maize. Our data show that none of the grasses tested were strongly preferred to maize, but the results indicate plants attractive to FAW adults and larvae that could be utilized in a multiple trap crop approach to target various stages of the pest. Furthermore, results indicate the importance of planting these companion plants earlier than maize.

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.

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.

Removing constraints to sustainable food production: new ways to exploit secondary metabolism from companion planting and GM.

The entire process of agricultural and horticultural food production is unsustainable as practiced by current highly intensive industrial systems. Energy consumption is particularly intensive for cultivation, and for fertilizer production and its incorporation into soil. Provision of nitrogen contributes a major source of the greenhouse gas, N2 O. All losses due to pests, diseases and weeds are of food for which the carbon footprint has already been committed and so crop protection becomes an even greater concern. The rapidly increasing global need for food and the aggravation of associated problems by the effects of climate change create a need for new and sustainable crop protection. The overall requirement for sustainability is to remove seasonal inputs, and consequently all crop protection will need to be delivered via the seed or other planting material. Although genetic modification (GM) has transformed the prospects of sustainable crop protection, considerably more development is essential for the realisation of the full potential of GM and thereby consumer acceptability. Secondary plant metabolism offers wider and perhaps more robust new crop protection via GM and can be accomplished without associated yield loss because of the low level of photosynthate diverted for plant defence by secondary metabolism. Toxic mechanisms can continue to be targeted but exploiting non-toxic regulatory and signalling mechanisms should be the ultimate objective. There are many problems facing these proposals, both technical and social, and these are discussed but it is certainly not possible to stay where we are in terms of sustainability. The evidence for success is mounting and the technical opportunities from secondary plant metabolism are discussed here. © 2019 The Authors. Pest Management Science published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.

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.

A maize landrace that emits defense volatiles in response to herbivore eggs possesses a strongly inducible terpene synthase gene.

Maize (Zea mays) emits volatile terpenes in response to insect feeding and egg deposition to defend itself against harmful pests. However, maize cultivars differ strongly in their ability to produce the defense signal. To further understand the agroecological role and underlying genetic mechanisms for variation in terpene emission among maize cultivars, we studied the production of an important signaling component (E)-caryophyllene in a South American maize landrace Braz1006 possessing stemborer Chilo partellus egg inducible defense trait, in comparison with the European maize line Delprim and North American inbred line B73. The (E)-caryophyllene production level and transcript abundance of TPS23, terpene synthase responsible for (E)-caryophyllene formation, were compared between Braz1006, Delprim, and B73 after mimicked herbivory. Braz1006-TPS23 was heterologously expressed in E. coli, and amino acid sequences were determined. Furthermore, electrophysiological and behavioral responses of a key parasitic wasp Cotesia sesamiae to C. partellus egg-induced Braz1006 volatiles were determined using coupled gas chromatography electroantennography and olfactometer bioassay studies. After elicitor treatment, Braz1006 released eightfold higher (E)-caryophyllene than Delprim, whereas no (E)-caryophyllene was detected in B73. The superior (E)-caryophyllene production by Braz1006 was positively correlated with high transcript levels of TPS23 in the landrace compared to Delprim. TPS23 alleles from Braz1006 showed dissimilarities at different sequence positions with Delprim and B73 and encodes an active enzyme. Cotesia sesamiae was attracted to egg-induced volatiles from Braz1006 and synthetic (E)-caryophyllene. The variation in (E)-caryophyllene emission between Braz1006 and Delprim is positively correlated with induced levels of TPS23 transcripts. The enhanced TPS23 activity and corresponding (E)-caryophyllene production by the maize landrace could be attributed to the differences in amino acid sequence with the other maize lines. This study suggested that the same analogous genes could have contrasting expression patterns in different maize genetic backgrounds. The current findings provide valuable insight not only into genetic mechanisms underlying variation in defense signal production but also the prospect of introgressing the novel defense traits into elite maize varieties for effective and ecologically sound protection of crops against damaging insect pests.

Plant volatile-mediated signalling and its application in agriculture: successes and challenges.

856 I. 856 II. 857 III. 858 IV. 859 V. 860 VI. 862 VII. 863 VIII. 864 IX. 866 866 References 866 SUMMARY: The mediation of volatile secondary metabolites in signalling between plants and other organisms has long been seen as presenting opportunities for sustainable crop protection. Initially, exploitation of interactions between plants and other organisms, particularly insect pests, foundered because of difficulties in delivering, sustainably, the signal systems for crop protection. We now have mounting and, in some cases, clear practical evidence for successful delivery by companion cropping or next-generation genetic modification (GM). At the same time, the type of plant signalling being exploited has expanded to signalling from plants to organisms antagonistic to pests, and to plant stress-induced, or primed, plant-to-plant signalling for defence and growth stimulation.

An Indirect Defence Trait Mediated through Egg-Induced Maize Volatiles from Neighbouring Plants.

Attack of plants by herbivorous arthropods may result in considerable changes to the plant's chemical phenotype with respect to emission of herbivore-induced plant volatiles (HIPVs). These HIPVs have been shown to act as repellents to the attacking insects as well as attractants for the insects antagonistic to these herbivores. Plants can also respond to HIPV signals from other plants that warn them of impending attack. Recent investigations have shown that certain maize varieties are able to emit volatiles following stemborer egg deposition. These volatiles attract the herbivore's parasitoids and directly deter further oviposition. However, it was not known whether these oviposition-induced maize (Zea mays, L.) volatiles can mediate chemical phenotypic changes in neighbouring unattacked maize plants. Therefore, this study sought to investigate the effect of oviposition-induced maize volatiles on intact neighbouring maize plants in 'Nyamula', a landrace known to respond to oviposition, and a standard commercial hybrid, HB515, that did not. Headspace volatile samples were collected from maize plants exposed to Chilo partellus (Swinhoe) (Lepidoptera: Crambidae) egg deposition and unoviposited neighbouring plants as well as from control plants kept away from the volatile emitting ones. Behavioural bioassays were carried out in a four-arm olfactometer using egg (Trichogramma bournieri Pintureau & Babault (Hymenoptera: Trichogrammatidae)) and larval (Cotesia sesamiae Cameron (Hymenoptera: Braconidae)) parasitoids. Coupled Gas Chromatography-Mass Spectrometry (GC-MS) was used for volatile analysis. For the 'Nyamula' landrace, GC-MS analysis revealed HIPV production not only in the oviposited plants but also in neighbouring plants not exposed to insect eggs. Higher amounts of EAG-active biogenic volatiles such as (E)-4,8-dimethyl-1,3,7-nonatriene were emitted from these plants compared to control plants. Subsequent behavioural assays with female T. bournieri and C. sesamiae parasitic wasps indicated that these parasitoids preferred volatiles from oviposited and neighbouring landrace plants compared to those from the control plants. This effect was absent in the standard commercial hybrid we tested. There was no HIPV induction and no difference in parasitoid attraction in neighbouring and control hybrid maize plants. These results show plant-plant signalling: 'Nyamula' maize plants emitting oviposition-induced volatiles attractive to the herbivore's natural enemies can induce this indirect defence trait in conspecific neighbouring undamaged maize plants. Maize plants growing in a field may thus benefit from this indirect defence through airborne signalling which may enhance the fitness of the volatile-emitting plant by increasing predation pressure on herbivores.

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.

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.

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.

New directions for improving crop resistance to insects by breeding for egg induced defence.

Plant defence responses to insect oviposition, including tritrophic interactions with natural enemies of herbivores, have rarely been targeted in crop breeding programmes. Emission of herbivore induced plant volatiles (HIPVs) that attract natural enemies early on at the egg-laying stage of herbivore attack could provide timely biological control of pests and deter subsequent oviposition. This is needed in an agroecological context where the third trophic level often does not keep pace with the growth rate of pests. Our very recent data, using maize as an example, show that herbivore egg induced volatile emission is very rare in commercial hybrids but common in farmer selected landraces. Strategies for crop genetic improvement to enhance such responses to insect attack are considered.

Delivering sustainable crop protection systems via the seed: exploiting natural constitutive and inducible defence pathways.

To reduce the need for seasonal inputs, crop protection will have to be delivered via the seed and other planting material. Plant secondary metabolism can be harnessed for this purpose by new breeding technologies, genetic modification and companion cropping, the latter already on-farm in sub-Saharan Africa. Secondary metabolites offer the prospect of pest management as robust as that provided by current pesticides, for which many lead compounds were, or are currently deployed as, natural products. Evidence of success and promise is given for pest management in industrial and developing agriculture. Additionally, opportunities for solving wider problems of sustainable crop protection, and also production, are discussed.

Achieving food security for one million sub-Saharan African poor through push-pull innovation by 2020.

Food insecurity is a chronic problem in Africa and is likely to worsen with climate change and population growth. It is largely due to poor yields of the cereal crops caused by factors including stemborer pests, striga weeds and degraded soils. A platform technology, 'push-pull', based on locally available companion plants, effectively addresses these constraints resulting in substantial grain yield increases. It involves intercropping cereal crops with a forage legume, desmodium, and planting Napier grass as a border crop. Desmodium repels stemborer moths (push), and attracts their natural enemies, while Napier grass attracts them (pull). Desmodium is very effective in suppressing striga weed while improving soil fertility through nitrogen fixation and improved organic matter content. Both companion plants provide high-value animal fodder, facilitating milk production and diversifying farmers' income sources. To extend these benefits to drier areas and ensure long-term sustainability of the technology in view of climate change, drought-tolerant trap and intercrop plants are being identified. Studies show that the locally commercial brachiaria cv mulato (trap crop) and greenleaf desmodium (intercrop) can tolerate long droughts. New on-farm field trials show that using these two companion crops in adapted push-pull technology provides effective control of stemborers and striga weeds, resulting in significant grain yield increases. Effective multi-level partnerships have been established with national agricultural research and extension systems, non-governmental organizations and other stakeholders to enhance dissemination of the technology with a goal of reaching one million farm households in the region by 2020. These will be supported by an efficient desmodium seed production and distribution system in eastern Africa, relevant policies and stakeholder training and capacity development.

Push-pull farming systems.

Farming systems for pest control, based on the stimulo-deterrent diversionary strategy or push-pull system, have become an important target for sustainable intensification of food production. A prominent example is push-pull developed in sub-Saharan Africa using a combination of companion plants delivering semiochemicals, as plant secondary metabolites, for smallholder farming cereal production, initially against lepidopterous stem borers. Opportunities are being developed for other regions and farming ecosystems. New semiochemical tools and delivery systems, including GM, are being incorporated to exploit further opportunities for mainstream arable farming systems. By delivering the push and pull effects as secondary metabolites, for example, (E)-4,8-dimethyl-1,3,7-nonatriene repelling pests and attracting beneficial insects, problems of high volatility and instability are overcome and compounds are produced when and where required.

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.