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

Abstracts 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

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

Climate-adapted companion cropping increases agricultural productivity in East Africa

AbstractProduction of cereals, the main staple and cash crops for millions of farmers in sub-Saharan Africa (SSA) is severely constrained by parasitic striga weed Striga hermonthica, stemborers and poor soil fertility. A companion cropping system known as ‘push–pull’ overcomes these constraints while providing additional soil fertility and forage grass benefits to smallholder farmers. To ensure the technology's long-term sustainability in view of the current and further potential aridification as a consequence of climate change, drought-tolerant crops, Brachiaria cv mulato (border crop) and greenleaf desmodium (intercrop), have been identified and incorporated into a ‘climate-adapted push–pull’. The aims of the current study were to evaluate effectiveness of the new system (i) in integrated control of striga and stemborer pests and (ii) in improving maize grain yields, and to evaluate farmers’ perceptions of the technology to assess potential for further adoption. 395 farmers who had adopted the technology in drier areas of Kenya, Uganda and Tanzania were randomly selected for the study. Each farmer had a set of two plots, a climate-adapted push–pull and a maize monocrop. Seasonal data were collected in each plot on the number of emerged striga plants, percentage of maize plants damaged by stemborers, plant height and grain yields. Similarly, farmers’ perceptions of the benefits of the technology were assessed using a semi-structured questionnaire. There were highly significant reductions in striga and stemborer damage to maize plants in the climate-adapted push–pull compared to the maize monocrop plots: striga levels were 18 times lower and stemborer levels were 6 times lower. Similarly, maize plant height and grain yields were significantly higher. Mean yields were 2.5 times higher in companion planting plots. Farmers rated the climate-adapted push–pull significantly superior in reducing striga infestation and stemborer damage rates, and in improving soil fertility and maize grain yields. These results demonstrate that the technology is effective in controlling both weeds and pests with concomitant yield increases under farmers’ conditions. It thus provides an opportunity to improve food security, stimulate economic growth, and alleviate poverty in the region while making agriculture more resilient to climate change

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

Biosynthesis of natural and novel C-glycosylflavones utilising recombinant Oryza sativa C-glycosyltransferase (OsCGT) and Desmodium incanum root proteins

The rice C-glycosyltransferase (OsCGT) is one of only a small number of characterised plant C-glycosyltransferases (CGT) known. The enzyme C-glucosylates a 2-hydroxyflavanone substrate with UDP-glucose as the sugar donor to produce C-glucosyl-2-hydroxyflavanones. We tested substrate specificity of the enzyme, using synthetic 2-hydroxyflavanones, and showed it has the potential to generate known natural CGFs that have been isolated from rice and also other plants. In addition, we synthesised novel, unnatural 2-hydroxyflavanone substrates to test the B-ring chemical space of substrate accepted by the OsCGT and demonstrated the OsCGT capacity as a synthetic reagent to generate significant quantities of known and novel CGFs. Many B-ring analogues are tolerated within a confined steric limit. Finally the OsCGT was used to generate novel mono-C-glucosyl-2-hydroxyflavanones as putative biosynthetic intermediates to examine the potential of Desmodium incanum biosynthetic CGTs to produce novel di-C-glycosylflavones, compounds implicated in the allelopathic biological activity of Desmodium against parasitic weeds from the Striga genus

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 fourarm 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

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

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 formillions 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

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