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

Please cite this article as: Amanuel TamiruZeyaur R. KhanToby JA Bruce New directions for improving crop resistance to insects by breeding for egg induced defence (2015), http://dx.

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.

The spotted stemborer, Chilo partellus (Swinhoe) (Lepidoptera: Crambidae), is one of the most important insect pests attacking maize and sorghum in Ethiopia. Recent studies have indicated that the pest is spreading to new locations where it was not reported before. In the current study, laboratory investigations were carried out to determine the combined effect of different levels of relative humidity and temperature regimes on the development and fecundity of C. partellus, as these physical factors are known to play an important role in the life cycle of insects and adaptability to local climate. Developmental time, longevity, potential fecundity and realized fecundity of C. partellus were measured under controlled conditions. Three temperature regimes (22°C, 26°C and 30°C) and three relative humidity levels (40%, 60% and 80%) were tested. It was found that temperature, relative humidity (RH) and their interaction significantly affected the developmental time, adult longevity, potential fecundity and realized fecundity of the pest. Developmental time was inversely related to temperature. Mean duration of C. partellus life cycle was 70.2 days at 22°C and 80% RH, whereas it took only 26.5 days to complete its life cycle at 30°C and 40% RH. Male and female longevity were similar in most cases. The adult life span ranged between 6.9-11.1 days at 22°C and 3.1-7.2 days at 30°C for different levels of relative humidity. The most suitable conditions for C. partellus development and fecundity were 26-30°C temperatures regimes and 60-80% RH levels.

A better understanding of the chemical cues employed by parasitoids to locate their herbivore hosts will contribute towards effective exploitation of parasitoids as biological control agents against damaging crop pests. The current study was conducted to establish key bioactive compounds mediating behavioural response in the parasitic wasp Cotesia sesamiae (Cameron, 1906) (Hymenoptera: Braconidae) by employing electrophysiological techniques and behavioural studies. Electroantennogram (EAG) recording with synthetic equivalents of herbivore-induced plant volatiles (HIPVs) using C. sesamiae confirmed that 13 compounds elicited electrophysiological activity. In an olfactometer bioassay, only (E)-4,8-dimethyl-1,3,7-nonatriene, (E)-β-farnesene, and (E,E)-4,8,-trimethyl-1,3,7-tridecatetraene elicited behavioural responses from the parasitoid when tested individually at a natural dose, while (R)-linalool and (E)-caryophyllene were attractive at the highest test dose. Other EAG-active HIPVs were not attractive either at lower or higher concentrations, suggesting their role as blend components rather than functioning as attractants by themselves. In contrast, nine-component synthetic HIPV blends formulated in the same ratio and concentration as in a natural sample was as attractive as the original headspace sample. By providing further insights into maize–stemborer–parasitoid tritrophic interactions, results from this study will help in selecting crop cultivars emitting appropriate HIPVs to attract natural enemies and perhaps repel damaging crop pests.

Aggregation of the bean flower thrips, Megalurothrips sjostedti (Trybom) (Thysanoptera: Thripidae), has been observed on cowpea, Vigna unguiculata (L.) Walp. To understand the mechanism underpinning this behavior, we studied the responses of M. sjostedti to headspace volatiles from conspecifics in a four-arm olfactometer. Both male and female M. sjostedti were attracted to male, but not to female odor. Gas chromatography/mass spectrometry (GC/MS) analyses revealed the presence of two distinct compounds in male M. sjostedti headspace, namely ( R )-lavandulyl 3-methylbutanoate (major compound) and ( R )-lavandulol (minor compound); by contrast, both compounds were only present in trace amounts in female headspace collections. A behavioral assay using synthetic compounds showed that male M. sjostedti was attracted to both ( R )-lavandulyl 3-methylbutanoate and ( R )-lavandulol, while females responded only to ( R )-lavandulyl 3-methylbutanoate. This is the first report of a male-produced aggregation pheromone in the genus Megalurothrips . The bean flower thrips is the primary pest of cowpea, which is widely grown in sub-Saharan Africa. The attraction of male and female M. sjostedti to these compounds offers an opportunity to develop ecologically sustainable management methods for M. sjostedti in Africa.

Maize lethal necrosis is one of the most devastating diseases of maize causing yield losses reaching up to 90% in sub-Saharan Africa. The disease is caused by a combination of maize chlorotic mottle virus (MCMV) and any one of cereal viruses in the Potyviridae group such as sugarcane mosaic virus. MCMV has been reported to be transmitted mainly by maize thrips (Frankliniella williamsi) and onion thrips (Thrips tabaci). To better understand the role of thrips vectors in the epidemiology of the disease, we investigated behavioral responses of F. williamsi and T. tabaci, to volatiles collected from maize seedlings infected with MCMV in a four-arm olfactometer bioassay. Volatile profiles from MCMV-infected and healthy maize plants were compared by gas chromatography (GC) and GC coupled mass spectrometry analyses. In the bioassays, both sexes of F. williamsi and male T. tabaci were significantly attracted to volatiles from maize plants infected with MCMV compared to healthy plants and solvent controls. Moreover, volatile analysis revealed strong induction of (E)-4,8-dimethyl-1,3,7-nonatriene, methyl salicylate and (E,E)-4,8,12-trimethyltrideca-1,3,7,11-tetraene in MCMV-infected maize seedlings. Our findings demonstrate MCMV induces changes in volatile profiles of host plants to elicit attraction of thrips vectors. The increased vector contact rates with MCMV-infected host plants could enhance virus transmission if thrips feed on the infected plants and acquire the pathogen prior to dispersal. Uncovering the mechanisms mediating interactions between vectors, host plants and pathogens provides useful insights for understanding the vector ecology and disease epidemiology, which in turn may contribute in designing integrated vector management strategies.

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.