The functional response of Uscana lariophaga under different egg distributions of its host Callosobruchus maculatus

The functional response of the egg parasitoid Uscana lariophaga Steffan (Hymenoptera: Trichogrammatidae) was tested under three different host distributions (even, clumped and random) within clusters of Callosobruchus maculatus Fab. (Coleoptera: Bruchidae) eggs. A Holling Type II functional response was found for all three distributions. Over low host densities, less than 50% of the host clusters was parasitized. At low host densities, U. lariophaga females parasitized significantly fewer eggs in random egg clusters with many beans than in clusters with fewer beans and an even or clumped egg distribution. At higher egg densities, plateau levels of maximum number of hosts parasitized were the same for all three egg distributions. Uscana lariophaga appears to be adapted to search for even or clustered egg distributions, as can be found in the field and under storage conditions.     

Impact of the egg parasitoid Uscana lariophaga and the larval-pupal parasitoid Dinarmus basalis on Callosobruchus maculatus populations and cowpea losses

In West Africa, Uscana lariophaga Steffan (Hymenoptera: Trichogrammatidae) and Dinarmus basalis (Rondani) (Hymenoptera: Pteromalidae) parasitize the eggs and larvae, respectively, of Callosobruchus maculatus Fab. (Coleoptera: Bruchidae), an important pest of stored cowpea. The impact of the parasitoids on pest populations was evaluated in clay pots similar to the ones used in traditional storage in Niger. At the beginning of the storage period, cowpeas were infested with one density of C. maculatus and with either one or both of the parasitoid species. If the parasitoids we inoculated as single species in stores, both D. basalis and U. lariophaga significantly suppressed the bruchid population, but the former did so more effectively than the latter. A combination of D. basalis and U. lariophaga resulted in the same suppression of bruchid populations as when D. basalis was the only parasitoid. Fifteen weeks after storage, the parasitoids reduced the number of grains damaged significantly by 38–56%. The effect of inoculating single or multiple parasitoid species on C. maculatus populations in an augmentation strategy is discussed.     

Uscana lariophaga, egg parasitoid of bruchid beetle storage pests of cowpea in West Africa: the effect of temperature and humidity

The stored-product bruchid pests,Callosobruchus maculatus (Fabricius) andBruchidius atrolineatus (Pic) cause considerable production losses in cowpea in West Africa.Uscana lariophage Steffan parasitizes the eggs of the bruchids both in the field and in storage. As chemical control of bruchids in traditional granaries is not appropriate for poor farmers, enhancement of the efficacy of the parasitoid by environmental manipulation has been investigated. The effect of temperature on the capacity ofU. lariophaga to parasitize eggs has been studied at eleven constant and three fluctuating temperatures within the range 10 to 45°C. Longevity of the female wasp decreased with increasing temperature. The rate of development increased linearly at temperatures from 17.5 to 35°C, but decreased from 35 to 40°C. Mortality of the developing wasp remained below 20% from 20 to 37.5°C, but outside this range, mortality reached 100% at 15 at 42.5°C. Most parasitization occurred at temperatures of 25 and 30°C. Sex ratio (percentage females) increased with temperature in the high temperature range. The intrinsic rate of increase (r_m) forU. lariophaga was highest in the temperature range from 30 to 37.5°C and was higher than that ofC. maculatus at all temperatures. While the r_m value ofC. maculatus did not vary much at temperatures from 25 to 35°C, the r_m value of the wasp doubled. Relative humidity did not effect longevity, egg-laying capacity, mortality, development time and sex ratio of the wasps withC. maculatus as host. However, withB. atrolineatus as the host, development time and mortality increased at lower relative humidity levels. The results indicate that temperature is the major regulating factor on the parasitoid. As the type of storage structure and its location (sun or shade) affects the temperature inside the store, ways are being investigated of manipulating the storage environment through temperature regulation to increase the impact of the parasitoid.     

Uscana lariophaga, West-African egg parasitoid of the cowpea bruchid beetleCallosobruchus maculatus: photoperiod, parasitization and eclosion interactions

The diurnal pattern of parasitization and eclosion of the trichogrammatidUscana lariophaga Steffan, egg parasitoid of the cowpea bruchidCallosobruchus maculatus (Fabricius) was studied for L12:D12 photoperiods. The wasp parasitized eggs throughout each 24-h period with about 70% of the parasitization taking place during the first 12 h regardless whether it was the photophase or the scotophase. More female progeny was produced when the first 12-h period was photophase instead of scotophase. Eclosion of wasps took place over a period of 4 days and occurred during the second half of the scotophase and the first half of the photophase. The number of wasps eclosed during the photophase was similar to that eclosed during the scotophase. Average development time was 8.9 days at 30°C. Male development was completed 6 to 8 h before the females, hence a higher percentage of females emerged in the later eclosion peaks. During scotophases more females eclosed than during photophases. Although the results indicate that the wasp is able to perform under dark storage conditions the effect of permanent low light intensity remains to be studied.     

Resistance to the cowpea weevil (Callosobruchus maculatus) larva in pods of cowpea (Vigna unguiculata)

Thirty cowpea, Vigna unguiculata (L.) Walpers, cultivars were evaluated as intact pods to determine if any possessed resistance to the cowpea weevil, Callosobruchus maculatus (Fabr.). Pod resistance was measured as pre-establishment larval mortality (PreM); those larvae dying after egg hatch but before penetrating into the seeds, and as post-establishment within-seed mortality (PostM); those larvae dying after penetrating into the seeds. Among the 30 varieties examined, PreM ranged from 57.9% to 99.4% and PostM ranged from 6.7% to 82.6%. Ten varieties exhibited total intact pod mortality (mortality from egg hatch to adult emergence from the seed) greater than 95%.Physical measurements were made of several pod and seed characteristics to ascertain whether the observed pod resistance was due to seed factors, pod-wall factors, or to interactions between the pod and seeds. Although resistance to breakage (from handling plus pod dehiscence) is a necessary characteristic for overall pod protection against C. maculatus; other factors are also important. Among the other pod and seed characteristics measured to identify major resistance factors, seed coat thickness was the one most highly correlated with pod resistance. Our results suggest that interactions between pod-wall and seed coat characteristics play a large role in pod resistance of cowpeas to C. maculatus.     

Parasitism by the mite Acarophenax lacunatus on beetle pests of stored products

Recent reports of the biocontrolpotential of the mite species Acarophenaxlacunatus (Cross & Krantz) (Prostigmata:Acarophenacidae) and the lack of biologicalstudies on this regulatory agent led to thepresent study carried out under laboratoryconditions. The objective of the investigationwas to assess the host range of A. lacunatus,so far only reported as egg parasite ofRhyzopertha dominica (F.) (Coleoptera:Bostrichidae). Four Coleoptera species ofstored cereals were used: R. dominica,Tribolium castaneum (Herbst) (Tenebrionidae),Cryptolestes ferrugineus (Stephens)(Laemophloeidae) and Oryzaephilus surinamensis (L.) (Cucujidae). The highest rates of eggparasitism were observed on R. dominica and T.castaneum, leading to a significant decrease ofpopulations of both species and reduced wheatweight loss. A. lacunatus was also able toparasitize eggs of C. ferrugineus, but not ofO. surinamensis. These results indicate abroader host range of A. lacunatus thaninitially suspected and also strengthen itspossibility of use in integrated pestmanagement programs in storage environments.     

Host selection behaviour and specificity of the solitary parasitoid of mealybugs Anagyrus sp. nr. pseudococci (Girault) (Hymenoptera,Encyrtidae)

The host selection behaviour of Anagyrus sp. nr. pseudococci was compared in no-choice tests among five mealybug species of different geographical and phylogenetic origin, including the Mediterranean native host, Planococcus ficus and four exotic mealybug species, one of the same genus, Pl. citri, two Pseudococcus species, Ps. calceolariae and Ps. viburni and a more distant one, Phenacoccus peruvianus. All five studied mealybug species were recognised by the parasitoid as potential hosts and parasitised, but the behavioural pattern of host recognition, host handling and the level of host acceptance of Anagyrus sp. nr. pseudococci significantly varied among the five studied species, indicating a clear preference for the two Planococcus species, Pl. ficus in particular. The results suggest that A. sp. nr. pseudococci has a broader host range and a more generalist behaviour in comparison with other Anagyrus species. Practical implications of the findings are discussed.     

Inheritance of resistance to the cowpea aphid in cowpea

Summary Inheritance of resistance to cowpea aphid, Aphis craccivora Koch, in three resistant cultivars of cowpea, Vigna unguiculata (L.) Walp, was studied. The parents, F₁ and F₂ population were grown in an insect-proof screenhouse. Each 3-day-old seedling was infested with 10 apterous adult aphids. Seedling reaction was recorded when the susceptible check was killed. The segregation data revealed that the resistance of ICV11 and TVU310 is governed by single dominant genes. All the F₂ seedlings of the cross ICV10xTVU310 were resistant, indicating that they have the same gene for resistance. However, the F₂ populations from the crosses ICV10xICV11 and ICV11xTVU310 segregated in a ratio of 151, indicating that the dominant genes in ICV11 and TVU310 are non-allelic and independent of each other. The resistance gene of ICV10 and TVU310 is designated as Ac₁ and that of ICV11 as Ac₂.     

Plasticity in the egg-spacing behavior of a seed beetle: Effects of host deprivation and seed patchiness (Coleoptera: Bruchidae)

Egg-laying females of the seed beetle Callosobruchus maculatusdiscriminate between egg-free and egg-laden seeds and produce a nearly uniform distribution of eggs among seeds. We examined plasticity in this trait in response to both an internal factor (level of host deprivation) and an environmental one (the spatial configuration of available seeds). Responses to each factor were measured in genetically divergent strains that show a relatively strong (S strain) or weak (B strain) tendency to spread eggs evenly among seeds. Following a modest (10-h) period of host deprivation, S-strain females distributed their eggs less uniformly among seeds; the proportion of females committing at least one oviposition mistake increased from 20 to 50%. Similarly, S-strain females distributed their eggs less uniformly if seeds were presented in multiple, discrete patches instead of in a single, large patch. The higher frequency of oviposition mistakes in the multiple-patch arena was caused in part by females maintaining a uniform distribution of eggs within patches but not among patches. In contrast, females from the sloppier B strain were unaffected by either host deprivation or resource dispersion. Responses to seed patchiness are discussed in relation to the role of learning in the egg-spacing behavior of C. maculatus.     

Patch-exploitation patterns in an egg parasite of elm leaf beetle

Eggs of elm beetle,Pyrrhalta luteola (Muller), normally occur in masses which can be viewed as discrete host patches in space and time. Analysis of>800 egg masses from 3 field sites in northern California revealed that the number of eggs/mass varied from 2 to>40 and that the imported egg parasiteTetrastichus gallerucae (Fonsc.) exploited a relatively large proported of the egg masses at certain times. The spatial relationship between parasitization and number of eggs/egg mass was assessed in 2 ways-i.e., for exploited masses only and for all masses combined (exploited+nonexploited). Percent parasitization was density independent in most cases for both data sets. It is suggested that both methods of analysis can provide patterns which are relevant to biological control of insect pests.      

Partitioning of biologically fixed nitrogen in cowpea during pod development

Two days after exposure of roots to¹⁵N labeled N₂, partitioning of biologically fixed N into leaves, stems, peduncles, pods, roots and nodules was measured in the early pod development stage of cowpea (Vigna unguiculata (L.). The experimental objective was to determine the quantity of biologically fixed N that is incorporated into vegetative tissue before being mobilized to pods. For the three varieties of cowpea included in the experiment a maximum of 50% of the N, biologically fixed two days earlier, was contained in the pods. The remaining N was distributed throughout the vegetative portion of the plant with at least 30% in stems and leaves which indicates that much of the newly fixed N must cycle through a N pool in these tissues before reaching the pods.     

Reproductive biology of two biotypes of Edovum puttleri, a parasitoid of Colorado potato beetle eggs

Emergence of adults of Edovum puttleri Grissell (Hymenoptera: Eulophidae) has a diel periodicity: most emergence occurred within 2 h of lights on (16L:8D, 24±1°C). Males emerged 2.0±3.0 h prior to females, and soon thereafter mated with emerging females. Courtship behavior did not differ between the Colombian and Mexican biotypes and was similar to that of members of the eulophid subfamily Tetrastichinae. The preoviposition period and oviposition period did not differ between biotypes. Males and females of both biotypes were long-lived, with the longevity of females of the Mexican biotype being significantly longer than that of Colombian females. males of both biotypes lived equally long. Mexican females produced more offspring than Colombian females, although this difference was not significant. After approximately 3 weeks, production of female offspring declined in both biotypes. A single mating was sufficient for females to produce a full complement of female offspring. Head capsule width of females correlated positively with fecundity.

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Résumé L'émergence des adultes de E. puttleri Grissell (Hymen.: Eulophidae) présente une périodicité quotidienne, la plupart de émergences ayant lieu dans les deux premières heures de la photophase (16L/8D, 24°±1°C). Les mâles apparaissent 2±3 h avant les femelles, et s'accouplant immédiatement avec les femelles émergentes. Les biotypes mexicains et columbiens ont le même comportement de cour, qui correspond à celui décrit pour les Tetrastichini et pour les Entedontinae. Il en est de même pour la durée, d'une part des périodes précédent la ponte (3,7 ±2,8 j), et d'autre part de la ponte elle-même (25,1 ±4,5 j). Les adults des deux biotypes vivent longtemps, mais les femelles mexicaines vivent plus longtemps (57,4±18 j) que les colombiennes (46,2±14,8 j); les longévités des mâles sont identiques (45,7 ±18 j). Les femelles mexicaines produisent apparemment plus d'oeufs (221,2±29,5) que les colombiennes (166,6±58,1), mais ces différences ne sont pas significatives. Après trois semaines environ la production de descendants régresse chez les deux types. Un seul accouplement suffit à produire le contingent total de filles. La largeur de la capsule céphalique des femelles est liée positivement à leur fécondité.

     

Developmental pattern formation of somatic embryos induced in cell suspension cultures of cowpea [Vigna unguiculata (L.) Walp]

The sequence of events in the functional body pattern formation during the somatic embryo development in cowpea suspensions is described under three heads. Early stages of somatic embryogenesis were characterized by both periclinal and anticlinal cell divisions. Differentiation of the protoderm cell layer by periclinal divisions marked the commencement of somatic embryogenesis. The most critical events appear to be the formation of apical meristems, establishment of apical-basal patterns of symmetry, and cellular organization in oblong-stage somatic embryo for the transition to torpedo and cotyledonary-stage somatic embryos. Two different stages of mature embryos showing distinct morphology, classified based on the number of cotyledons and their ability to convert into plantlets, were visualized. Repeated mitotic divisions of the sub-epidermal cell layers marked the induction of proembryogenic mass (PEM) in the embryogenic calli. The first division plane was periclinally-oriented, the second anticlinally-oriented, and the subsequent division planes appeared in any direction, leading to clusters of proembryogenic clumps. Differentiation of the protoderm layer marks the beginning of the structural differentiation in globular stage. Incipient procambium formation is the first sign of somatic embryo transition. Axial elongation of inner isodiametric cells of the globular somatic embryo followed by the change in the growth axis of the procambium is an important event in oblong-stage somatic embryo. Vacuolation in the ground meristem of torpedo-stage embryo begins the process of histodifferentiation. Three major embryonic tissue systems; shoot apical meristem, root apical meristem, and the differentiation of procambial strands, are visible in torpedo-stage somatic embryo. Monocotyledonary-stage somatic embryo induced both the shoot apical meristem and two leaf primordia compared to the ansiocotyledonary somatic embryo.     

Exploring the foraging environment of a natural enemy of Callosobruchus maculatus: Spatial egg distribution in stored cowpea

Knowledge of the spatial distribution of stored product insects may reduce the dependency on chemicals for control of these insects. Biological control, for instance, could be improved based on such knowledge. In this paper we describe the three-dimensional spatial oviposition pattern of Callosobruchus maculatus in stored cowpea. Individual C. maculatus females oviposited in clusters of 70±15 (SD) eggs. These clusters were variable in shape. In any cluster 90 to 95% of the eggs fitted into a volume of 19.1±3.5 cm³. The egg density was highest (0.6 eggs bean^–1) at the center of a cluster and decreased towards the periphery. A statistically significant relationship existed between the number of eggs n in a cluster and the cluster volume, V(cm³): V=11.5+0.11n. We also investigated the spatial egg distribution of beetles which emerged from egg clusters such as those produced by individual females. Their oviposition was not confined to one specific area but was scattered throughout the bean mass. A point pattern analysis showed that the density of the `parent' cluster had no effect on the spatial egg pattern. These results give insight into the foraging environment which the egg parasitoid Uscana lariophaga, a promising candidate for biological control of C. maculatus, is facing. We argue that the probability p of encountering at least one other bean with eggs after a parasitization is a function of the number n of beans that are visited: p=1–0.42 (0.37)^(n–1).     

Performance of bean bruchids Callosobruchus maculatus and Zabrotes subfasciatus (Coleoptera: Bruchidae) reared on resistant (IT81D-1045) and susceptible (Epace 10) Vigna unguiculata seeds: relationship with trypsin inhibitor and vicilin excretion

Callosobruchus maculatus (Cm) and Zabrotes subfasciatus (Zs) were reared on resistant (IT81D-1045) and on susceptible (Epace 10) cowpea seeds. The emergence of adult insects, total developmental period (TDP) and excretion of trypsin inhibitor and vicilin were determined for both bruchid populations. Parameter evaluation showed that the Zs populations emerged from both seeds had no significant differences in emergence and TDP. The Cm population raised from resistant seeds had lower emergence (5.6+/-1.3%) and delayed TDP (46+/-1.25 days) than those emerged from susceptible seeds. The excretion of defense proteins showed that Zs reared in resistant seeds excreted 1.7 times more trypsin inhibitor, but this did not affect emergence or TDP. Furthermore, Cm population emerged from resistant seeds excreted 7 times higher vicilin and 0.4 times less trypsin inhibitor than that emerged from susceptible seeds. These results indicate that vicilins from resistant seeds are involved to significantly longer TDP (46 days) and also drastic reduction of insect emergence ( approximately 5%) of C. maculatus.     

Effects of water deficit on N2(C2H2) fixation in cowpea and groundnut

The effect of water deficit on nodulation, N₂ fixation, photosynthesis, and total soluble sugars and leghemoglobin in nodules was investigated in cowpea and groundnut. Nitrogenase activity completely ceased in cowpea with a decrease in leaf water potential ( _leaf) from –0.4 MPa to –0.9 MPa, while in groundnut it continued down to –1.7 MPa. With increasing water stress, the acetylene reduction activity (ARA) declined very sharply in cowpea, but ARA gradually decreased in groundnut. Even with mild water stress ( _leaf of 0.2 MPa), nodule fresh weight declined 50% in cowpea partly due to a severe nodule shedding whereas nodule fresh weight declined in groundnut only when _leaf decreased by 1.0 MPa. No nodule shedding was noticed even at a higher stress level in groundnut. Photosynthesis and stomatal conductance were also more stable in groundnut than in cowpea under water stress. There was a sharp increase in total soluble sugars and leghemoglobin in the nodules of groundut with water stress, but no definite trend could be found in cowpea.     

Flight initiation in the egg parasitoid Trichogramma minutum: Effects of ambient temperature, mates, food, and host eggs

Emergence, preening, and flight initiation were studied in laboratory-reared Trichogramma minutum Riley (Hymenoptera: Trichogrammatidae). Male parasitoids emerged first and flew before females. When both sexes were present in flight cylinders, female parasitoids flew before males. Flight propensity in males was negatively related to the number of emerging females, while flight propensity in females was independent of the number of males present. Ambient temperature significantly affected the propensity and timing of flight; between 70–80% of the parasitoids flew at 25 and 30 °C while less than 4%, mostly males, flew at 20 °C. No flights were observed at 15 °C. The presence of fresh host eggs caused a reduction in the proportion of female parasitoids that flew and a delay in the time to flight for the females that did fly. The presence of food increased the flight propensity of female parasitoids, but did not affect the timing of flight. The relationship between flight behaviour and the efficiency of mass-reared Trichogramma is discussed in terms of its importance for inundative release programmes.     

The genetic basis of egg lay response to conditioned medium in the flour beetle,Tribolium castaneum

Summary The rate of egg laying in Tribolium castaneum is affected by the quality of the environment and can thus serve not only as a component of fitness of the individual, but also as an adaptation to the long-term survival of the population. In an attempt to determine whether it is an independent genetic character, selection for high and low rates of response was carried jut in beetles from three wild populations. When tests were done on virgin females, and responsiveness, corrected for scale effects, was used as the criterion for selection, separation between the lines was achieved in one generation. It is suggested that in natural populations of Tribolium, the responsiveness of the rate of egg laying to the quality of the environment is controlled by a small number of genes.     

Characterization of two cDNAs for novel drought-inducible genes in the highly drought-tolerant cowpea

Ten cDNAs for drought-inducible genes were isolated using differential screening of a cDNA library prepared from 10-hr dehydrated cowpea plants,Vigna unguiculata (S. Iuchi, K. Yamaguchi-Shinozaki, T. Urao, T. Terao, K. Shinozaki; Plant Cell Physiology, 1996 in press). Two of the cDNA clones, designated CPRD12 and CPRD46, were sequenced and characterized. The CPRD12 and CPRD46 cDNAs encode putative proteins related to nonmetallo-short-chain alcohol dehydrogenase (CPRD12) and chloroplastic lipoxygenase (CPRD46). Northern blot analysis revealed that these genes are induced by high-salinity stress and exogenous abscisic acid, but not by cold stress. The CPRD46 gene is also responsive to heat stress and methyl jasmonate and salicylic acid. Genomic Southern blot analysis suggested that CPRD12 constitutes a small gene family, but that CPRD46 is a single copy gene. We discuss the possible functions of these two CPRD gene products under drought stress.     

The effect of host size on quality attributes of the egg parasitoid,Trichogramma pretiosum

In a study of the quality ofTrichogramma pretiosum Riley (Hymenoptera: Trichogrammidae), we compared female wasps emerging from natural hosts, parasitized in the laboratory or the field with those emerging from factitious hosts used for commercial mass production. Females from the natural hosts were larger, more fecund, and longer lived than those from the factitious hosts. Compared to small females, large female wasps are substantially more fecund when honey (carbohydrate) is available but marginally more fecund when honey is unavailable. The size of a femaleT. pretiosum depends on two factors: the size of the host egg from which it emerges even when the wasp was gregarious, and the number of conspecifics that emerge with it. The similarities in the size distribution of female wasps emerging from natural hosts, in conjunction with the mechanism by whichTrichogramma measure host size and allocate eggs accordingly, suggests the hypothesis that size related components of fitness in femaleT. pretiosum are under strong selection in the field.