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

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

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

A climate-adapted push-pull system effectively controls fall armyworm, Spodoptera frugiperda (J E Smith), in maize in East Africa

Fall armyworm, Spodoptera frugiperda (J E Smith), an economically important pest native to tropical and subtropical America has recently invaded Africa, causing substantial damage to maize and other crops. We evaluated functionality of a companion cropping system, ‘climate-adapted push-pull’, developed for control of cereal stemborers in drier agro-ecologies, as an added tool for the management of fall armyworm. The technology comprises intercropping maize with drought-tolerant greenleaf desmodium, Desmodium intortum (Mill.) Urb., and planting Brachiaria cv Mulato II as a border crop around this intercrop. Protection to maize is provided by semiochemicals that are emitted by the intercrop that repel (push) stemborer moths while those released by the border crop attract (pull) them. 250 farmers who had adopted the technology in drier areas of Kenya, Uganda and Tanzania were randomly selected for the study during the long rainy season (March-August) of 2017. Each farmer had a set of two plots, a climate-adapted push–pull and a maize monocrop. Data were collected in each plot on the number of fall armyworm larvae on maize, percentage of maize plants damaged by the larvae and maize grain yields. Similarly, farmers' perceptions of the impact of the technology on the pest were assessed using a semi-structured questionnaire. Reductions of 82.7% in average number of larvae per plant and 86.7% in plant damage per plot were observed in climate-adapted push-pull compared to maize monocrop plots. Similarly, maize grain yields were significantly higher, 2.7 times, in the climate-adapted push-pull plots. Farmers rated the technology significantly superior in reducing fall armyworm infestation and plant damage rates. These results demonstrate that the technology is effective in controlling fall armyworm with concomitant maize grain yield increases, and represent the first documentation of a technology that can be immediately deployed for management of the pest in East Africa and beyond

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

Qualitative Distribution of Candidatus Phytoplasma Oryzae in Roots, Stems and Leave of Napier Grass (Pennisetum purpureum)

For sustainable production of Napier grass (Pennisetum purpureum), screening for the absence of phytoplasma on propagation material (stem) is carried by Nested-PCR in detection of 16S rDNA on leaves.  However, the aim of this study was to investigate if there is uneven distribution of the pathogen in roots, stems and leaves of the infected plants that could influence screening results. A total of 294 Napier grasses infected with candidatus phytoplasma oryzae were sampled from western Kenya for detection of Napier Stunt Disease (Phytoplasma Oryzae) by nested PCR. There was significant difference on distribution of phytoplasma in roots, stems and leaves (F=36.26, df=2, p<0.001). The results show that high number of positive samples were detected from stem (94.1 %), leaves (77 %) and roots (44.1 %) samples, respectively. Conclusively, the study revealed uneven distribution thus inclusion of stem samples should be considered during screening procedures. Other factors like seasonal variation are also essential to be studied. Keywords: Phytoplasma, Napier grass, nested PCR, Distribution of the pathogen, Propagatio

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

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

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