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

Farmers' perception and evaluation of brachiaria Grass (brachiaria spp.) genotypes for smallholder cereal-livestock production in East Africa

Brachiaria (Urochloa) is a genus, common name brachiaria, of forage grasses that is increasingly transforming integrated crop-livestock production systems in East Africa. A study was undertaken to (i) assess smallholder farmers’ perception on benefits of brachiaria in cereal-livestock production, (ii) identify brachiaria production constraints, and (iii) identify farmer preferred brachiaria genotypes. A multi-stage sampling technique was adopted for sample selection. Data were collected through semi-structured individual questionnaire and focus group discussions (FGDs). The study areas included Bondo, Siaya, Homabay and Mbita sub-counties in Western Kenya and the Lake zone of Tanzania. A total of 223 farmers participated in individual response questionnaires while 80 farmers participated in the FGDs. The respondents considered brachiaria mainly important in management of cereal pests (70.4% of respondents) and as an important fodder (60.8%). The major production constraint perceived by both male and female respondents is attacks by arthropods pests (49.2% and 63%, respectively). Spider smites had been observed on own farms by 50.8% of men and 63.1% of women, while sorghum shoot flies had been observed by 58.1% of men and 67.9% of women. These pests were rated as a moderate to severe problem. Xaraes was the most preferred genotype, followed by Mulato II and Piata. These genotypes are important in developing new crop pest management strategies, such as push-pull, and for relatively rapid improvements in crop management and yield increases, particularly in developing countries

Push-pull farming system controls fall armyworm: lessons from Africa

Fall armyworm (FAW) Spodoptera frugiperda invaded Africa, with the first detections being reported in Central and Western Africa in early 2016, and now affects at least 40 countries in Africa, causing up to total crop loss amounting to over $6.2 billion p.a. FAW is an invasive polyphagous pest that causes damage to economically important crops and has recently been reported in the Indian sub-continent. Effective control of FAW through use of synthetic chemical pesticides and genetically modified crops such as Bt maize faces challenges including improper use, unaffordability by smallholder farmers and development of resistance by the pest. Additionally, dispersal of FAW larvae into the lower maize plant canopy keeps them out of reach of topical insecticide applications. Integrated pest management (IPM) packages like the push-pull technology which eliminate pesticide use, and deploy natural processes are more suitable and cost-effective. Push-pull is a farming system intensification approach that involves attracting insect pests with trap plants (pull) such as Napier grass (Pennisetum purpureum) or Brachiaria grass, while driving them away from the main crop using a repellent intercrop (push), Desmodium spp., commonly known as desmodium, and attracting natural parasitoids and predators to the field. In the rhizosphere, chemicals secreted by desmodium roots inhibit attachment of germinated striga to maize or sorghum roots and abort germination of striga seeds which are rapidly depleted in the soil. Moreover, it improves soil fertility by fixing nitrogen, improving carbon sequestration, organic matter, moisture retention, and soil biota, and prevents further degradation of soil. The climate-adapted push-pull technology significantly reduces plant damage by FAW and is the first IPM management tool for the pest in Africa, and is well suited to agro-ecosystem intensification needs of smallholder mixed farming systems in Africa and beyond. Both the African and the Asian continents provide favourable climatic conditions for sustained reproduction of the FAW, which is expected to result in severe damage to crops; and being a new pest in both continents, it might have few natural enemies. Conventional control methods have limited effectiveness, as explained above. Therefore, an IPM approach that is compatible with mixed cropping farming systems of small and medium scale farmers is necessary. The climate adapted push-pull is the first demonstrated IPM management tool for the FAW in Africa. The technology combines this with other concomitant benefits including control of stemborers and the parasitic striga weeds, improvement of soil health through factors such as nitrogen fixation by desmodium, natural mulching, moisture retention and improvement of soil organic matter and soil biota. The combined benefits, including control of FAW, result in ecologically sustainable higher crop yields, and well suited to agro-ecosystem intensification needs of smallholder mixed farming systems in Africa and beyond

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, N2O. 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

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

Field evaluation of a new third generation push-pull technology for control of striga weed, stemborers, and fall armyworm in western Kenya

Production of cereal crops in sub-Saharan Africa is threatened by parasitic striga weeds and attack by stemborers and the invasive fall armyworm (FAW), compounded by increasing hot and dry conditions. A climate-smart push-pull technology (PPT) significantly reduces effects of these biotic challenges. To improve further resilience of the system to climate change, more adapted and suitable companion plants were identified and integrated in a new version of PPT, termed ‘third generation PPT’. Our study evaluates field performance and farmer opinions of this new version in comparison with the earlier version, climate-smart PPT, and farmers’ own practices of growing maize in controlling stemborers, FAW, and striga weeds. Trials were conducted across five locations in western Kenya for two cropping seasons in the year 2019 following a one-farm one-replicate completely randomized design. We assessed infestation on striga, stemborers, and FAW, and yield performance of the three cropping systems. We also sought the opinions of the hosting farmers through semi-structured questionnaires that were administered through individual interviews. Both PPT plots recorded significantly (P < 0.05) lower striga count, FAW, and stemborer damage, and higher grain yield than in plots that followed farmers’ own practices. There was no statistically significant difference between the two PPT plots except for stemborer damage for which the third generation PPT recorded higher damage than the climate-smart PPT. However, farmers preferred the third generation PPT for important traits possessed by its companion plants which their counterparts in climate-smart PPT are deficient. The cultivar Xaraes was rated as ‘very good’ for resistance to spider mites, biomass yield, and drought tolerance while Desmodium incanum was rated ‘very good’ for seed production and drought tolerance. The third generation PPT is based on companion crops that are more resilient to hot and dry conditions which are increasing rapidly in prevalence with climate change. This version therefore presents a better option to upscale the technology and meet different needs of farmers especially in arid and semi-arid conditions

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

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

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

The dominant Anopheles vectors of human malaria in Africa, Europe and the Middle East: occurrence data, distribution maps and bionomic précis

<p>Abstract</p> <p>Background</p> <p>This is the second in a series of three articles documenting the geographical distribution of 41 dominant vector species (DVS) of human malaria. The first paper addressed the DVS of the Americas and the third will consider those of the Asian Pacific Region. Here, the DVS of Africa, Europe and the Middle East are discussed. The continent of Africa experiences the bulk of the global malaria burden due in part to the presence of the <it>An. gambiae </it>complex. <it>Anopheles gambiae </it>is one of four DVS within the <it>An. gambiae </it>complex, the others being <it>An. arabiensis </it>and the coastal <it>An. merus </it>and <it>An. melas</it>. There are a further three, highly anthropophilic DVS in Africa, <it>An. funestus</it>, <it>An. moucheti </it>and <it>An. nili</it>. Conversely, across Europe and the Middle East, malaria transmission is low and frequently absent, despite the presence of six DVS. To help control malaria in Africa and the Middle East, or to identify the risk of its re-emergence in Europe, the contemporary distribution and bionomics of the relevant DVS are needed.</p> <p>Results</p> <p>A contemporary database of occurrence data, compiled from the formal literature and other relevant resources, resulted in the collation of information for seven DVS from 44 countries in Africa containing 4234 geo-referenced, independent sites. In Europe and the Middle East, six DVS were identified from 2784 geo-referenced sites across 49 countries. These occurrence data were combined with expert opinion ranges and a suite of environmental and climatic variables of relevance to anopheline ecology to produce predictive distribution maps using the Boosted Regression Tree (BRT) method.</p> <p>Conclusions</p> <p>The predicted geographic extent for the following DVS (or species/suspected species complex*) is provided for Africa: <it>Anopheles </it>(<it>Cellia</it>) <it>arabiensis</it>, <it>An. </it>(<it>Cel.</it>) <it>funestus*</it>, <it>An. </it>(<it>Cel.</it>) <it>gambiae</it>, <it>An. </it>(<it>Cel.</it>) <it>melas</it>, <it>An. </it>(<it>Cel.</it>) <it>merus</it>, <it>An. </it>(<it>Cel.</it>) <it>moucheti </it>and <it>An. </it>(<it>Cel.</it>) <it>nili*</it>, and in the European and Middle Eastern Region: <it>An. </it>(<it>Anopheles</it>) <it>atroparvus</it>, <it>An. </it>(<it>Ano.</it>) <it>labranchiae</it>, <it>An. </it>(<it>Ano.</it>) <it>messeae</it>, <it>An. </it>(<it>Ano.</it>) <it>sacharovi</it>, <it>An. </it>(<it>Cel.</it>) <it>sergentii </it>and <it>An. </it>(<it>Cel.</it>) <it>superpictus*</it>. These maps are presented alongside a bionomics summary for each species relevant to its control.</p