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William A Overholt - One of the best experts on this subject based on the ideXlab platform.
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release and establishment of Cotesia flavipes cameron hymenoptera braconidae an exotic parasitoid of chilo partellus swinhoe lepidoptera crambidae in east and southern africa
Annales De La Societe Entomologique De France, 2006Co-Authors: C. O. Omwega, William A Overholt, Eric Muchugu, Fritz SchulthessAbstract:Abstract Cotesia flavipes Cameron (Hymenoptera: Braconidae) was first imported into Kenya in 1991 from Pakistan for control of Chilo partellus (Swinhoe) (Lepidoptera: Crambidae). First releases were made at the Kenya coast in 1993 and a few recoveries of the parasitoid were made the following year. Additional foreign exploration for C. flavipes was conducted in central India in 1996 and 1998, which resulted in additional importation of the parasitoid for subsequent releases in eastern and southern Africa. Region-wide releases commenced with releases in Mozambique in 1996, Somalia in 1997 and Uganda in 1998. By 2005, many releases had been made in 10 countries in East and Southern Africa with establishment being reported in all of them except Eritrea but including Ethiopia where releases had never been made. This paper describes the progress made in the release and establishment of C. flavipes in East and Southern Africa and quantifies the rate of spread from the initial release sites in Kenya.
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responses of the stem borer larval endoparasitoid Cotesia flavipes hymenoptera braconidae to plant derived synomones laboratory and field cage experiments
Biocontrol Science and Technology, 2005Co-Authors: Esther N Ngumbi, William A Overholt, Adele J Ngisong, E N M Njagi, Rita Torto, Lester J Wadhams, Michael A Birkett, J A Pickett, Baldwyn TortoAbstract:Abstract Laboratory and field cage experiments investigated the response of females of the stem borer larval endoparasitoid Cotesia flavipes to two synthetic synomone components, the terpenoid (E)-β-farnesene and the green leaf volatile, (Z)-3-hexenyl acetate, both compounds identified previously in headspace volatiles of maize plants damaged by stem borer (Chilo partellus). In dose response tests performed in a Y-tube olfactometer, parasitoids were significantly more attracted to the arms bearing 10 or 15 µg of (Z)-3-hexenyl acetate and (E)-β-farnesene than to the control arm. (E)-β-Farnesene was as attractive as the essential oil from the plant Hemizygia petiolata (Lamiaceae) rich in the same compound (80% relative amount). The plant essential oil elicited responses from females of the parasitoid comparable to those elicited by two positive controls, stem borer larval frass and adult parasitoid diet (20% honey solution), tested in the laboratory assays. In field cage trapping experiments, captures in tr...
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Cotesia flavipes cameron hymenoptera braconidae does not exhibit complementary sex determination ii evidence from laboratory experiments
Applied Entomology and Zoology, 2004Co-Authors: Emmanuel Iyamulemye Niyibigira, William A Overholt, Richard StouthamerAbstract:Sex determination in the order Hymenoptera is based on haplodiploid arrhenotoky; in which males develop from unfertilized eggs and are haploid, whereas females develop from fertilized eggs and are diploid. However, some hymenopteran species produce diploid males through a mechanism known as single-locus complementary sex determination (sl-CSD). In these species, heterozygous individuals at a single sex locus develop into females, whereas hemizygotes (haploids) and homozygotes (diploids) develop into males. Inbreeding leads to homozygosity and consequent production of diploid males. We investigated the presence of sl-CSD in the braconid Cotesia flavipes using a series of inbreeding crosses among five isofemale lines. Sex ratio (proportion of females) did not differ among the within-line crosses, between-line crosses and crosses carried out between isofemale lines and a mixed (outbred) colony. Brood size of within-line and between-line crosses did not differ. Culturing populations for 25 generations did not result in changes to more male-biased sex ratios. We conclude that CSD does not exist in C. flavipes. The implications of absence of CSD in C. flavipes are discussed in context of mass rearing for classical biological control programmes.
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comparative studies on the influence of relative humidity and temperature on life table parameters of two populations of Cotesia flavipes hymenoptera braconidae
Biocontrol Science and Technology, 2004Co-Authors: Emana Getu, William A Overholt, Eunicf KairuAbstract:Cotesia flavipes Cameron (Hymenoptera: Braconidae) is a gregarious larval koinobiont parasitoid of gramineous stemborers from the Indo-Australian region. More recently, it has been introduced into several countries in East and southern Africa for regulation of Chilo partellus (Swinhoe) (Lepidoptera: Crambidae). Establishment has varied from country to country and within country, suggesting that abiotic factors, such as temperature and relative humidity, may influence parasitoid performance. In this work, the effect of temperature and relative humidity on life table parameters of two populations of C. flavipes were measured. The results indicated that the factors and their interactions significantly affected the population growth of C. flavipes. The intrinsic rate of increase of the North Pakistan population of Cotesia flavipes was higher than that of the Indian population at all humidities at 28°C, but there were no differences at other temperatures or humidities.
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Cotesia flavipes Cameron and Cotesia sesamiae (Cameron) (Hymenoptera : Braconidae) do not exhibit complementary sex determination : Evidence from field populations
Applied Entomology and Zoology, 2004Co-Authors: Emmanuel Iyamulemye Niyibigira, William A Overholt, Richard StouthamerAbstract:Diploid males are expected to occur in populations of some Hymenoptera because of the single-locus complementary sex determination (sl-CSD) mechanism. If sl-CSD is found in a species that produces gregarious broods and sib mates at a high frequency, a fraction of these sib-matings (matched mating) produces diploid male offspring. We developed models to predict the matched mating frequency in populations with different frequencies of sib mating and egg fertilization. The predictions of these models are used to determine if we can use the distributions of brood sex ratio and brood size to determine if a species has sl-CSD. The models show that sl-CSD can be detected from these brood sex ratios if the diploid male offspring survives. We applied our models to Cotesia sesamiae and its exotic congener Cotesia flavipes (Hymenoptera: Braconidae), to determine if sl-CSD occurs in these species. Parasitoids were reared from stemborer larvae sampled from maize fields in Kenya between 1992 and 1999. We found no evidence for the presence of sl-CSD with survival of diploid males in both braconid species, but we cannot exclude the possibility that sl-CSD with diploid male mortality takes place in these species.
C. O. Omwega - One of the best experts on this subject based on the ideXlab platform.
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release and establishment of Cotesia flavipes cameron hymenoptera braconidae an exotic parasitoid of chilo partellus swinhoe lepidoptera crambidae in east and southern africa
Annales De La Societe Entomologique De France, 2006Co-Authors: C. O. Omwega, William A Overholt, Eric Muchugu, Fritz SchulthessAbstract:Abstract Cotesia flavipes Cameron (Hymenoptera: Braconidae) was first imported into Kenya in 1991 from Pakistan for control of Chilo partellus (Swinhoe) (Lepidoptera: Crambidae). First releases were made at the Kenya coast in 1993 and a few recoveries of the parasitoid were made the following year. Additional foreign exploration for C. flavipes was conducted in central India in 1996 and 1998, which resulted in additional importation of the parasitoid for subsequent releases in eastern and southern Africa. Region-wide releases commenced with releases in Mozambique in 1996, Somalia in 1997 and Uganda in 1998. By 2005, many releases had been made in 10 countries in East and Southern Africa with establishment being reported in all of them except Eritrea but including Ethiopia where releases had never been made. This paper describes the progress made in the release and establishment of C. flavipes in East and Southern Africa and quantifies the rate of spread from the initial release sites in Kenya.
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performance of Cotesia flavipes hymenoptera braconidae in parasitizing chilo partellus lepidoptera crambidae as affected by temperature and host stage
Biological Control, 2004Co-Authors: Nanqing Jiang, Mamoudou Setamou, Adele J Ngisong, C. O. OmwegaAbstract:Abstract The braconid larval parasitoid, Cotesia flavipes , is used as a biological control agent against the crambid Chilo partellus , a serious pest of cereal crops in eastern and southern Africa. We examined the survival, development parameters, and body growth patterns of the host and its parasitoid at different temperatures (22, 26, and 30 °C) using third and fourth instars of Ch. partellus . For non-parasitized hosts, larval mortality tended to be highest at lowest temperature and for parasitized at third host instars only at highest temperature. Development time of Co. flavipes immatures significantly decreased with host instar and with temperature. Sex ratio of Co. flavipes varied from male- to female-biased with increase in temperature. The increase in body weight of parasitized fourth instar Ch. partellus was higher than in non-parasitized larvae at all temperatures. Parasitism by Co. flavipes had no effect on the food uptake by Ch. partellus , but significantly less food was consumed by both parasitized and non-parasitized larvae at 26 °C. The results of this study were discussed in light of the performance of Co. flavipes under different climatic conditions.
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Near-infrared Spectroscopic Method to Identify Cotesia flavipes and Cotesia sesamiae (Hymenoptera: Braconidae)
Annals of The Entomological Society of America, 2003Co-Authors: T J Cole, C. O. Omwega, William A Overholt, Floyd E. Dowell, Sonny B RamaswamyAbstract:Parasitoids of the Cotesia flavipes complex (C. flavipes and Cotesia sesamiae) are natural enemies of stemboring lepidopteran pests in sub-Saharan Africa. The two species are difÞcult to differentiate using morphological markers, and a quick, reliable test was sought for their correct identiÞcation. After numerous, unsuccessful attempts at developing species-speciÞc monoclonal antibodies that could differentiate between the two species, we were successful in using near-infrared spectroscopy to distinguish the cocoons of the two species with an accuracy of better than 85%. Calibrations were established using partial least squares analysis, enabling identiÞcation of cocoons of known species, cocoons from an alternative host, as well as in blind tests. This technology would greatly expedite identiÞcation of Þeld-caught insects used to determine ecological parameters and parasitization rates of an individual species.
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Evidence of the establishment of Cotesia flavipes (Hymenoptera: Braconidae), a parasitoid of cereal stemborers, and its host range expansion in Ethiopia.
Bulletin of Entomological Research, 2003Co-Authors: Emana Getu, Eunicf Kairu, William A Overholt, C. O. OmwegaAbstract:Three lepidopteran cereal stemborers, Chilo partellus (Swinhoe) (Crambidae), Busseola fusca Fuller, and Sesamia calamistis Hampson (Noctuidae) were collected from maize and sorghum in Ethiopia. The noctuid stemborers are indigenous to Africa while C. partellus is an introduced species from Asia. In 1999, the Asian stemborer parasitoid, Cotesia flavipes Cameron (Braconidae) was found to be widespread in Ethiopia, even though it had never been released in the country. In addition to attacking Chilo partellus, Cotesia flavipes was reared from B. fusca and S. calamistis. The origin of C. flavipes in Ethiopia may have been Somalia where it was released in 1997 near the border with eastern Ethiopia. Percent parasitism of borers by C. flavipes was higher in eastern Ethiopia than other surveyed regions, and parasitism was higher in 2000 than 1999. Parasitism was higher when cereals were intercropped with other plants and when wild grass hosts of stemborers were present.
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Cereal stemborer species complex and establishment of Cotesia flavipes Cameron in Eastern Uganda.
International Journal of Tropical Insect Science, 2001Co-Authors: T. Matama-kauma, J. A. Ogwang, C. O. Omwega, Samuel Kyamanywa, H R WillsonAbstract:Studies were conducted in two districts of eastern Uganda from 1997 to 1999 to introduce and monitor the establishment of an exotic parasitoid, Cotesia flavipes Cameron (Hymenoptera: Braconidae), for the control of the stemborer Chilo Partellus (Swinhoe) (Lepidoptera: Pyralidae), and also to determine the stemborer species complex in maize and sorghum. The study confirmed the presence of four important stemborers, two pyralids Ch. partellus and Eldana saccharina Walker and two noctuids, Busseola fusca Fuller and Sesamia calamistis Hampson. Chilo Partellus was dominant, constituting 53-88% of stemborers fourd followed by B. fusca at 8–37 %. The most abundant local parasitoid was the larval parasitoid Cotesia sesamiae (Cameron) (Hymenoptera: Braconidae). The pupal parasitoids Pediobius furvus Gahan (Hymenoptera: Eulophidae) and Dentichasmias busseolae Heinrich (Hymenoptera: Ichneumonidae) were also recorded. Parasitism of Co. sesamiae on Ch. partellus ranged between 0 and 13.1 %. The introduced Co. flavipes was recovered from all sites in four consecutive seasons (between 1998 and 1999) causing parasitism of between 4 and 32.9 % on Ch. partellus. Cotesia flavipes was also recovered from the indigenous stemborers B. fusca and S. calamistis. This study indicates that Co. flavipes has established in eastern Uganda.
Elisa Aparecida Gregório - One of the best experts on this subject based on the ideXlab platform.
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mitochondria in the midgut epithelial cells of sugarcane borer parasitized by Cotesia flavipes cameron 1891
Brazilian Journal of Biology, 2010Co-Authors: Daniela O Pinheiro, M D Silva, Elisa Aparecida GregórioAbstract:The sugarcane borer Diatraea saccharalis (Lepidoptera: Crambidae) has been controlled by Cotesia flavipes (Hymenoptera: Braconidae); however, very little is known about the effect of the parasitism in the host organs, including the midgut. This work aims to verify mitochondrial alteration in the different midgut epithelial cells of D. saccharalis parasitized by C. flavipes. Midgut fragments (anterior and posterior region) of both non-parasitized and parasitized larvae were processed for transmission electron microscopy. The mitochondria of midgut epithelial cell in the parasitized larvae exhibit morphological alteration, represented by matrix rarefaction and vacuolisation. These mitochondrial alterations are more pronounced in the anterior midgut region during the parasitism process, mainly in the columnar cell.
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spherites in the midgut epithelial cells of the sugarcane borer parasitized by Cotesia flavipes
Biocell, 2008Co-Authors: Daniela O Pinheiro, Helio Conte, Elisa Aparecida GregórioAbstract:Diatraea saccharalis, the main pest of sugarcane, has been controlled by Cotesia flavipes. Very little is known about the effect of parasitism on the host organs, including the midgut. The Lepidoptera midgut epithelium is composed of columnar, goblet, regenerative, and endocrine cells. Spherites have been described in columnar and regenerative cells of several Lepidoptera species, and presented a lot of functional meaning. We identified spherites in the midgut epithelial cells of non-parasitized D. saccharalis larvae analyzed the effect of parasitism on spherite morphology and distribution along the length of the midgut. Midgut fragments of both non-parasitized and parasitized larvae were processed for transmission electron microscopy. All the midgut epithelial cells showed spherites, but they were not preferentially located in a particular part of the cells. Parasitized larvae had more spherites, mainly in the columnar cells, than non-parasitized larvae. This observation was associated with an ionic imbalance within the insect host. Spherites were more abundant in the anterior midgut region than in other regions, which suggests that this region is involved in ion transport by intracellular and/or paracellular route. The morphological variability of spherites in the cells of parasitized larvae was related to the developmental stages of these structures.
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morphometry of the midgut epithelium of diatraea saccharalis fabricius 1794 lepidoptera parasitized by Cotesia flavipes cameron 1891 hymenoptera
Journal of Invertebrate Pathology, 2006Co-Authors: Daniela O Pinheiro, Reinaldo Jose Da Silva, Elisa Aparecida GregórioAbstract:The morphometric study of the midgut in Diatraea saccharalis (Lepidoptera) larvae parasitized by the Cotesia flavipes (Hymenoptera) showed that there was significant increase in the columnar, goblet and regenerative cells and their nuclei; the midgut lumen diameter and the epithelial height were also increased in the parasitized larvae. The multivariate analysis showed that parasitism affected the columnar cell only in the posterior region, and the goblet cells along the midgut length (anterior and posterior regions).
R.p.j. Potting - One of the best experts on this subject based on the ideXlab platform.
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geographic variation in host selection behaviour and reproductive success in the stemborer parasitoid Cotesia flavipes hymenoptera braconidae
Bulletin of Entomological Research, 1997Co-Authors: R.p.j. Potting, L. E. M. Vet, William A OverholtAbstract:Local parasitoid populations may be adapted to their sympatric major plant host complex. Parasitoid strains may thus differ in their propensity to search for a particular micro-habitat or host or they may differ in their physiological compatibility with particular plant or host species. Cotesia flavipes Cameron, a larval parasitoid used worldwide in biological control against tropical stemborers, has a wide host range in diverse habitats. The existence of plant and/or host specific strains in C. flavipes has been postulated. To provide insight into the existence of strains in C. flavipes, we compared the plant/host complex selection behaviour, and physiological compatibility with different stemborers, of six different geographic strains of C. flavipes that differed in the plant/host complex they were obtained from. The results of the host selection experiments indicate that there is no intraspecific variation in host selection behaviour among C. flavipes strains. However, our comparative experiments show variation in reproductive success among strains. The most significant result was that the strain with the longest period of co-existence with the new host Diatraea saccharalis Fabricius had the highest reproductive success on this host species. We argue that the reported existence of C. flavipes strains is based not on differences in host selection behaviour, but on differences in physiological compatibility between local parasitoid and host population.
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absence of odour learning in the stemborer parasitoid Cotesia flavipes
Animal Behaviour, 1997Co-Authors: R.p.j. Potting, H OttenAbstract:Abstract Parasitic wasps are commonly found to learn olfactory and visual cues that are associated with successful host location. For many parasitoids the cues that are associated with hosts vary in space and time, and are therefore unpredictable. An ability to learn allows the wasp to concentrate on those cues that will lead it to new hosts most effectively in a particular area. In contrast, parasitoids that forage in a predictable homogeneous environment and/or make only a few foraging decisions do not need to learn and should rely on innate responses to specific cues. The role of learning in host foraging was studied in Cotesia flavipes (Hymenoptera: Braconidae), a parasitoid of stemborer larvae with an ecology where learning is expected to be of low adaptive value. There was no evidence that C. flavipes uses odour learning in host-micro-habitat location. There was no significant effect of the development and emergence environment on the response level or preference towards the odour of infested plants. Neither was there evidence that experience with a particular plant–host complex during foraging influences subsequent foraging decisions in C. flavipes females. The absence of learning in C. flavipes which seems an exception among the parasitoids studied, is discussed in relation to its ecology.
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evolutionary and applied aspects of the behavioural ecology of the stemborer parasitoid Cotesia flavipes
International Journal of Tropical Insect Science, 1997Co-Authors: R.p.j. PottingAbstract:Cotesia flavipes (Hymenoptera: Braconidae) is a braconid larval parasitoid that has been used in the biological control of tropical stemborers in gramineous plants. The current research status on the behavioural ecology of this parasitoid is reviewed. The stimuli involved in the location of the host-microhabitat and the behaviour on infested plants and inside stemborer tunnels are discussed. Several aspects of the host foraging behaviour on C. flavipes are discussed in relation to the ecology of the parasitoid, such as the role of learning in foraging and clutch size allocation decisions. Variation in behaviour can be an obstacle in the effective use of natural enemies in biological control. Behavioural variation may exist because individuals differ genetically in their propensity to find or accept different hosts. Individuals may also differ because they have experienced different environments (i.e. learning). The second part of this paper focuses on the intraspecific variability in C. flavipes behaviour and discusses to what extent the reported plant and host specificity in C. flavipes has a genetic basis or is due to phenotypic plasticity through learning.
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Fitness consequences of superparasitism and mechanism of host discrimination in the stemborer parasitoid Cotesia flavipes
Entomologia Experimentalis et Applicata, 1997Co-Authors: R.p.j. Potting, H. M. Snellen, L. E. M. VetAbstract:The fitness consequences of superparasitism and the mechanism of host discrimination in Cotesia flavipes, a larval parasitoid of concealed stemborer larvae was investigated. Naive females readily superparasitized and treated the already parasitized host as an unparasitized host by allocating the same amount of eggs as in an unparasitized host. However, there was no significant increase in the number of emerging parasitoids from superparasitized hosts due to substantial mortality of parasitoid offspring in superparasitized hosts. Furthermore, the developmental time of the parasitoids in a superparasitized host was significantly longer than in a singly parasitized host and the emerging progeny were significantly smaller (body length and head width). Naive females entered a tunnel in which the host was parasitized 4 h previously and accepted it for oviposition. Experienced females (oviposition experience in unparasitized host) refused to enter a tunnel with a host parasitized by herself or by another female. In experiments where the tunnel and/or host was manipulated it was demonstrated that the female leaves a mark in the tunnel when she parasitizes a host. The role of patch marking in C. flavipes is discussed in relation to the ecology of the parasitoid.
Daniela O Pinheiro - One of the best experts on this subject based on the ideXlab platform.
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external morphology of Cotesia flavipes hymenoptera braconidae during larval development
Zoologia, 2010Co-Authors: Daniela O Pinheiro, Guilherme Duarte Rossi, Fernando Luis CônsoliAbstract:Cotesia flavipes (Cameron, 1891) (Hymenoptera) is a gregarious endoparasitoid used in applied biological control against Diatraea saccharalis (Fabricius, 1794) (Lepidoptera). In this study, we characterize the larval external morphology and the number of instars of C. flavipes. Parasitized larvae of D. saccharalis were sampled from the 1st to the 10th day after parasitism and dissected in an anticoagulant buffer for collection of C. flavipes immatures. Immatures were processed for scanning electron microscopy. Larvae of C. flavipes were prepared in NaOH solution and slide mounted to allow for mandible size measurements. Analysis of measurements of the parasitoid larval mandible size indicated that C. flavipes has three instars. Newly hatched larvae are caudate-mandibulate, assuming a hymenopteriform shape later in their development. The anal vesicle began to expand in the first instar and, once expanded, remained unchanged up to the beginning of the third instar. At the third instar, the anal vesicle decreased in volume. Herein we report the development and possible functions of the larval external structures modified during the development of C. flavipes, as for example their role in aiding newly-eclosed larvae to avoid the host immune response and to move within the host. To summarize the morphological changes during parasitoid growth, we should mention that the modifications in the anal vesicle were correlated with the feeding activity, and the maintenance of the anal vesicle indicates that this structure remained functional, probably playing a role in nutrient absorption and host regulation. On the other hand, the mandibles of early stage larvae are probably used to assist the parasitoid larvae during eclosion.
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mitochondria in the midgut epithelial cells of sugarcane borer parasitized by Cotesia flavipes cameron 1891
Brazilian Journal of Biology, 2010Co-Authors: Daniela O Pinheiro, M D Silva, Elisa Aparecida GregórioAbstract:The sugarcane borer Diatraea saccharalis (Lepidoptera: Crambidae) has been controlled by Cotesia flavipes (Hymenoptera: Braconidae); however, very little is known about the effect of the parasitism in the host organs, including the midgut. This work aims to verify mitochondrial alteration in the different midgut epithelial cells of D. saccharalis parasitized by C. flavipes. Midgut fragments (anterior and posterior region) of both non-parasitized and parasitized larvae were processed for transmission electron microscopy. The mitochondria of midgut epithelial cell in the parasitized larvae exhibit morphological alteration, represented by matrix rarefaction and vacuolisation. These mitochondrial alterations are more pronounced in the anterior midgut region during the parasitism process, mainly in the columnar cell.
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spherites in the midgut epithelial cells of the sugarcane borer parasitized by Cotesia flavipes
Biocell, 2008Co-Authors: Daniela O Pinheiro, Helio Conte, Elisa Aparecida GregórioAbstract:Diatraea saccharalis, the main pest of sugarcane, has been controlled by Cotesia flavipes. Very little is known about the effect of parasitism on the host organs, including the midgut. The Lepidoptera midgut epithelium is composed of columnar, goblet, regenerative, and endocrine cells. Spherites have been described in columnar and regenerative cells of several Lepidoptera species, and presented a lot of functional meaning. We identified spherites in the midgut epithelial cells of non-parasitized D. saccharalis larvae analyzed the effect of parasitism on spherite morphology and distribution along the length of the midgut. Midgut fragments of both non-parasitized and parasitized larvae were processed for transmission electron microscopy. All the midgut epithelial cells showed spherites, but they were not preferentially located in a particular part of the cells. Parasitized larvae had more spherites, mainly in the columnar cells, than non-parasitized larvae. This observation was associated with an ionic imbalance within the insect host. Spherites were more abundant in the anterior midgut region than in other regions, which suggests that this region is involved in ion transport by intracellular and/or paracellular route. The morphological variability of spherites in the cells of parasitized larvae was related to the developmental stages of these structures.
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morphometry of the midgut epithelium of diatraea saccharalis fabricius 1794 lepidoptera parasitized by Cotesia flavipes cameron 1891 hymenoptera
Journal of Invertebrate Pathology, 2006Co-Authors: Daniela O Pinheiro, Reinaldo Jose Da Silva, Elisa Aparecida GregórioAbstract:The morphometric study of the midgut in Diatraea saccharalis (Lepidoptera) larvae parasitized by the Cotesia flavipes (Hymenoptera) showed that there was significant increase in the columnar, goblet and regenerative cells and their nuclei; the midgut lumen diameter and the epithelial height were also increased in the parasitized larvae. The multivariate analysis showed that parasitism affected the columnar cell only in the posterior region, and the goblet cells along the midgut length (anterior and posterior regions).
