The potential for underutilized crops to improve security of food production.

Staple crops face major challenges in the near future and a diversification away from over-reliance on staples will be important as part of the progress towards the goal of achieving security of food production. Underutilized or neglected crops species are often indigenous ancient crop species which are still used at some level within the local, national or even international communities, but have the potential to contribute further to the mix of food sources than they currently do. The most cost-effective and easily disseminated changes that can be made to a crop are changes to the genetics, as these are contained within the seed itself and, for many species, the seed is a pure breeding, self-replicating, resource. This article focuses on the potential of underutilized crops to contribute to food security and, in particular, whether genetics and breeding can overcome some of the constraints to the enhanced uptake of these species in the future. The focus here is on overview rather than detail and subsequent articles will examine the current evidence base.

Intraguild interactions between the entomopathogenic fungus Pandora neoaphidis and an aphid predator and parasitoid at the population scale.

The interactions that occur between the entomopathogenic fungus Pandora neoaphidis and a predator (Coccinella septempunctata) and a parasitoid (Aphidius ervi) were assessed in microcosm and polytunnel experiments. Transmission of P. neoaphidis to the pea aphid, Acyrthosiphon pisum, was enhanced in the presence of both C. septempunctata and A. ervi in microcosm experiments done under fixed abiotic conditions. In contrast, the reproductive success of A. ervi was reduced in the presence of P. neoaphidis. Despite the increased fungal transmission in the presence of C. septempunctata, there was no additional decrease in the aphid population indicating that P. neoaphidis is functionally redundant in the presence of the coccinellid. In polytunnel experiments the reproductive success of A. ervi was not affected by P. neoaphidis. These results do not support those of the microcosm and may be due to the more natural abiotic conditions in the polytunnel reducing the competitive advantage of the fungus. Microcosms therefore provide an arena in which the interactions between fungal pathogens and other aphid-natural enemies can be assessed however, further assessments at increased spatial scales under more natural abiotic conditions are also required to accurately determine the outcome of these interactions.

Phenolic content and antioxidant capacity of hybrid variety cocoa beans.

Cocoa (Theobroma cacao L.) is a major, economically important, international crop and has been associated with several nutritional benefits including high antioxidant capacity. New cocoa hybrids have been developed in Ghana that exhibit resistance to pest damage during storage. The aim of this work was to assess the phenolic content and antioxidant capacity of these new hybrids in comparison to more traditional cocoa varieties. Total extractable phenolics were similar in all the four hybrids tested ranging from 69.9 to 81.6FAEg(-1). These levels were very similar to that extracted from traditional beans (73.8±2.5FAEg(-1)). The "phenolic profile" was determined by HPLC. A total of 25 peaks was observed but there were only minor differences in this profile between traditional and hybrid bean extracts. Antioxidant capacity was determined using the FRAP assay and traditional beans were found to possess 12.4µmolTEg(-1). In comparison the hybrid beans had antioxidant capacities ranging from 21.6 to 45.5µmolTEg(-1), and these were significantly higher than in the traditional beans for three out of the four hybrids. Since the phenolic and antioxidant levels and in these hybrid varieties were either similar to, or higher than, that obtained from traditional beans, the introduction of these new varieties would be unlikely to impact detrimentally on these nutritional components of the beans.

Behavioural responses of Tribolium castaneum (Herbst) to volatiles identified from dry cocoa beans.

New hybrid types of cocoa beans are attractive to insects in storage, however some of the insects feed little, if at all, on these beans compared to those of the traditional type (mixed genotypes). Based on a sniffing test using GCMS, differences in flavour volatiles in these types of beans have been determined and from these, six major volatiles of cocoa beans were selected for olfactometric analysis using a Pettersson olfactometer to determine which of them contributed to the attraction of Tribolium castaneum (Herbst) to the cocoa beans. The behaviour of Tribolium was affected by dose of 2-phenyl ethanol, acetophenone, 3-methyl butyraldehyde, ethyl butyrate, ethyl 3-hydroxybutyrate and butyl 2-methacrylate. Compared to beans of the new hybrid varieties, beans of the traditional type cocoa contained less 3-methylbutyraldehyde but more ethyl butyrate and acetophenone. In future breeding programmes, reducing the amount of acetophenone and ethyl butyrate but increasing the amount of 3-methylbutyraldehyde in cocoa beans may deter Tribolium from feeding on beans in storage.

The growth of Tribolium castaneum (Herbst) and Lasioderma serricorne (Fabricius) on feed media dosed with flavour volatiles found in dry cocoa beans.

Acetophenone, ethyl butyrate and 2-phenyl ethanol were used as additives to media on which Tribolium castaneum (Herbst) and Lasioderma serricorne (Fabricius) were cultured for 65 days. Acetophenone and ethyl butyrate had a positive impact on the weight and multiplication of the insects. The weight and rate of multiplication was higher in feed media containing acetophenone alone. The amount of the flavour volatiles added was high enough to deter the Lasioderma from feeding well, which affected their growth. This attraction to specific flavour volatiles in some stored products could be used in the formulation of effective control measures for insect pests.

Effect of fungal infection on the reproductive potential of aphids and their progeny.

The effect of infection by Pandora neoaphidis and Beauveria bassiana on the reproductive potential of the pea aphid, Acyrthosiphon pisum, and their progeny was assessed. Infection by either P. neoaphidis or B. bassiana reduced the number of nymphs produced within 24 h of inoculation and over the entire infection period compared to uninfected aphids. However, infection by either P. neoaphidis or B. bassiana for 24 or 72 h did not alter the intrinsic rate of increase of the host aphid's progeny. Therefore, fungal infection appears to have no indirect effects on the fitness of the host's progeny.

Pandora neoaphidis transmission and aphid foraging behaviour.

Pandora neoaphidis is an aphid-specific entomopathogen that produces infective conidia. As aphid movement increases, so does the likelihood of contact with conidia. Volatile distress signals released in response to aphid infestation as an indirect defence against herbivory may affect aphid foraging and, therefore, the fungus-aphid interaction. In this study, two different methods were used to investigate the effect of plant volatiles and P. neoaphidis-sporulating cadavers on (1) the colonisation of Vicia faba plants by Acyrthosiphon pisum and (2) P. neoaphidis transmission. This study indicates that A. pisum does not avoid bean plants containing P. neoaphidis and that transmission of conidia occurs during plant colonisation and, to a lesser extent, during in situ feeding. Although significantly more aphids were recovered from damaged plants compared to undamaged plants, the likelihood of infection was not affected by previous infestation by aphids.

Response of the entomopathogenic fungus Pandora neoaphidis to aphid-induced plant volatiles.

We used a model plant-aphid system to investigate whether the aphid-specific entomopathogenic fungus Pandora neoaphidis responds to aphid-induced defence by the broad-bean plant, Vicia faba. Laboratory experiments indicated that neither in vivo sporulation, conidia size nor the in vitro growth of P. neoaphidis was affected by Acyrthosiphon pisum-induced V. faba volatiles. The proportion of conidia germinating on A. pisum feeding on previously damaged plants was significantly greater than on aphids feeding on undamaged plants, suggesting a direct functional effect of the plant volatiles on the fungus. However, there were no significant differences in the infectivity of P. neoaphidis towards A. pisum feeding on either undamaged V. faba plants or plants previously infested with A. pisum. Therefore, these results provide no evidence to suggest that P. neoaphidis contributes to plant indirect defence strategies.

Effect of spatial heterogeneity on the role of Coccinella septempunctata as an intra-guild predator of the aphid pathogen Pandora neoaphidis.

The foraging behavior of starved and non-starved adult and larval Coccinella septempunctata on groups of plants in the presence of Pandora neoaphidis-infected Acyrthosiphon pisum, uninfected aphids or a mixture of these two prey types was compared. In general results of these studies confirmed the results of previous work comparing foraging behavior on a smaller spatial scale in Petri dishes. However, behaviors were modified in response to spatial complexity, prey quality, and the host plant. Starved C. septempunctata adults and larvae fed for longer and consumed more aphids than non-starved coccinellids. Both larvae and adults fed on infected aphids and in some cases entirely consumed them. This was thought to be due to the ease of capture of infected (dead) aphids and the feeding stimuli provided by the presence of the host plant and, where there was a choice of prey, uninfected aphids in the environment. Both larvae and adults spent the majority of the time foraging in the upper regions of plants and visited more plants when they were not starved or when they were in the presence of less suitable, infected aphid prey.

Implications of predator foraging on aphid pathogen dynamics.

The foraging behavior of starved and nonstarved second and fourth instar Coccinella septempunctata larvae on dead Acyrthosiphon pisum aphids, either infected with the entomopathogenic fungus Erynia neoaphidis (sporulating) or uninfected, was examined. Larvae searched for longer and fed less when presented with infected rather than uninfected A. pisum. Although no sporulating infected aphids were completely consumed, both adult and larval ladybirds can still be considered as intraguild predators. In a further study, fourth instar larvae fed on dying infected, dead infected (not sporulating), and dead uninfected aphids for similar periods of time but again the infected aphids were seldom entirely consumed. Live uninfected aphids were fed upon for significantly longer than any other prey. Infected aphids which were damaged at an early stage of infection (0, 1, or 2 days after inoculation) did not sporulate, whereas damaged moribund aphids (3 days after inoculation) did subsequently sporulate. Damaged sporulating cadavers continued to sporulate. However, damage to moribund and sporulating infected aphids, both mechanical or due to C. septempunctata feeding, reduced the number of conidia subsequently produced. Larval feeding caused the most significant reduction. Under laboratory conditions, C. septempunctata foraging on infected aphids did, therefore, reduce the pathogen density. However, conidia produced from a damaged cadaver resulted in levels of transmission to healthy aphids comparable to that resulting from an intact cadaver. Furthermore, the presence of a foraging adult ladybird resulted in a significant increase in transmission of the fungus to healthy aphids. Preliminary studies to assess the potential of other aphid natural enemies as intraguild predators illustrated that adults of the generalist carabid, Pterostichus madidus, entirely consumed sporulating cadavers. Third instar lacewing, Chrysoperla carnea, and hoverfly, Episyrphus balteatus, larvae never fed on sporulating cadavers. The ecological implications of these results are discussed.