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1.
Specimens, representative of each of the major taxa of mosquitoes, were fixed in copula and the external genitalia examined by scanning electron microscopy. The periphery of the basin-like everted aedeagus of Aedus aegypti precisely matches that of the everted atrial membrane of the female. These structures are appressed during coitus and sealed by pressure of the paraprocts, aedeagal pouch and proctiger. When everted, the aedeagus of male Culex pipiens reveals a ridged dome that surrounds the genital opening. This dome seals itself laterally into a gutter formed by pad-like extensions of the female's genital lips and is sealed dorsally by pressure of the aedeagal apodeme. The aedeagus of another culicine species, Wyeomyia smithii, bears the gonopore at the apex of a spined tube. This tube is inserted between the female's genital lips and is sealed within the genital atrium. The aedeagus of the toxorhynchitine species Toxorhynchitis brevipalpus is immobile and is inserted deep within the genital atrium of the female where it is sealed by pressure of the atrial walls. Males of each of these mosquitoes deliver a mixture of semen and sperm to the copulatory bursa of the female. After withdrawal of the aedeagus, sperm is transferred to the spermathecae. In contrast, sperm of Anopheles quadrimaculatus are delivered directly to the spermathecal duct. The tube-like aedeagus is positioned by its leaflets during sperm transfer and is driven deep into the atrium, where a mixture of semen and sperm is ejaculated. The significance of mechanical barriers to mating between species is discussed.  相似文献   

2.
Theory predicts that costly secondary sexual traits will evolve heightened condition dependence, and many studies have reported strong condition dependence of signal and weapon traits in a variety of species. However, although genital structures often play key roles in intersexual interactions and appear to be subject to sexual or sexually antagonistic selection, few studies have examined the condition dependence of genital structures, especially in both sexes simultaneously. We investigated the responses of male and female genital structures to manipulation of larval diet quality (new versus once‐used mung beans) in the bruchid seed beetle Callosobruchus maculatus. We quantified effects on mean relative size and static allometry of the male aedeagus, aedeagal spines, flap and paramere and the female reproductive tract and bursal spines. None of the male traits showed a significant effect of diet quality. In females, we found that longer bursal spines (relative to body size) were expressed on low‐quality diet. Although the function of bursal spines is poorly understood, we suggest that greater bursal spine length in low‐condition females may represent a sexually antagonistic adaptation. Overall, we found no evidence that genital traits in C. maculatus are expressed to a greater extent when nutrients are more abundant. This suggests that, even though some genital traits appear to function as secondary sexual traits, genital traits do not exhibit heightened condition dependence in this species. We discuss possible reasons for this finding.  相似文献   

3.
Koen Martens 《Hydrobiologia》1991,218(2):127-131
The male of Sclerocypris tuberculata (Methuen), thus far unknown, is here described. Relying on the morphology of the copulatory appendages and of the prehensile palps, it appears that this taxon belongs to a separate species group, together with S. zelaznyi and perhaps also S. sarsi. There are some interesting sexual dimorphic characters in the valve morphology: males have shorter valves with a dorsal margin which runs nearly parallel to the ventral one (more elongated valves with sloping dorsal margin in females) and there is lobe-like projection of the valve margin on the ventro-caudal corner of the LV in females which is lacking in the male. Furthermore, the female genital region has a very aberrant morphology, and all specimens from the present collection possess the tuberculated and noded valves.  相似文献   

4.
It is widely admitted that sexual selection is the responsible force behind genital traits. However, the particular mechanisms of genital evolution are still debated. Recently, studies of genital static allometry in insects have been used to elucidate such mechanisms. Insect genital traits are often reported to show negative allometry (i.e., a slope < 1), which has generated a number of ideas on how genital traits are selected. However, many studies that have inferred selection mechanisms have omitted consideration of the function of genital traits, used unreliable indicators of body size, and only rarely included female genitalia in their analysis. We investigated whether negative allometry operates for genitalia in two damselfly species (Protoneura cara and Ischnura denticollis). Damselflies are suitable for genital allometry tests as their genital function and body size indicators (wing length and head width) are relatively well known and established. First, we show that the aedeagus is used to physically remove sperm from both sperm storage organs (bursa and spermatheca) and that wing length and head width correlate positively with other morphological traits for the two study species. Second, we estimated genital allometry by measuring aedeagal length, vaginal length, bursal volume, and spermathecal volume. Our results indicate no consistent allometric pattern. Allometry for aedeagal length and vaginal width was not the same. Thus, there was no support for a negative allometric relationship. We urge researchers investigating allometry to look directly at how genitalia function rather than inferring function from allometric relationships only.  相似文献   

5.
Summary Diplo-X flies homozygous for the transform-er-2 ts (tra-2 ts) mutation develop into females at 16° C, while they develop into males at 29° C (Belote and Baker 1982). By means of this conditional mutation, we have carried out a detailed analysis of the development of the genital disc. Temperature shifts between 16 and 29° C, in both directions, and temperature pulses at 29° C, have been applied during the larval growth of tra-2 ts homozygous diplo-X flies, and the external derivatives of the genital disc have been analysed. Genital discs shifted from 16 to 29° C rapidly lose their capacity to differentiate female genital structures, while they become able to differentiate male genital structures whose inventory is more complete the earlier in larval development the temperature shift is carried out; moreover, duplicated male genital structures were observed. In the shift from 29 to 16° C, the genital disc loses its capacity to differentiate male genital structures, while it becomes able to differentiate female genital structures. The inventory of male structures is smaller, and the inventory of the female structures is more complete, the earlier in larval development the temperature is shifted. No duplicated female or male genital structures were observed in the downshift experiment. With respect to the analia, the shift from 16 to 29° C resulted in the quick formation of pure male anal plates, while in the opposite shift the formation of pure female anal plates occurred gradually. Moreover, the time course for the dorsal and ventral anal plates to show normal female phenotype was different: when the dorsal anal plates were completely normal, it was still possible to find incomplete ventral anal plates. In the pulse experiment at 29° C, the genital disc is able to differentiate both female and male genital structures, although the inventory of the latter ones was not complete. In addition, the capacity of the genital disc to differentiate male genital structures depended on the duration of the temperature pulse. The anal plates were always female, although they showed a reduction in their size, the ventral female anal plate being more affected than the dorsal one. No male anal plates were observed. The results have revealed that the genital disc follows a sequence in its capacity to differentiate female or male adult structures. We suggest that this sequence reflects the sequence of determination events occurring in the genital disc during its larval growth. In addition, results shown here provide evidence for the existence in the female genital primordium of a set of cells capable of giving rise either to female genital structures (ventral vaginal plates) or to male genital structures (hypandrium and penis apparatus). We also present evidence supporting the previous idea of two primordia for the anal plates.  相似文献   

6.
Mutations of the transformer-2 (tra-2) locus of Drosophila melanogaster cause chromosomally female (XX) animals to develop as males, but have no effect on the development of chromosomally male (XY) animals. In the female genital disc, such mutations cause repression of growth and inhibition of differentiation in the female genital primordium, while allowing growth and differentiation of the otherwise repressed male genital primordium. We used a temperature-sensitive mutation of this locus (tra-2ts1) to switch development from one sexual pathway to the other. Following development at the male-determining temperature (29°C), subsequent culture of the XX;tra-2ts1 genital disc in vivo at the female-determining temperature (16°C) allowed the previously repressed female genital primordium to develop and form female genital structures, whereas the formation of male genital elements was grossly disturbed. Conversely, following development at the female-determining temperature, subsequent culture in vivo at the male-determining temperature allowed the formerly repressed male genital primordium to grow and produce male genital structures, and repressed the formation of female elements from the already fully developed female genital primordium. The experiments indicate that the tra-2 product has to operate during the culture period in order to maintain the female state of sex determination, i.e., to promote the development of female structures, as well as to repress that of male structures. The experimental treatments, as well as the results of temperature shifts on developing larvae, resulted in sexual transformation of the anal plates, and clarified the sexual homologies of these structures. In both genitalia and analia, a switch from the female to the male developmental pathway was accomplished more rapidly and effectively than the reverse change.  相似文献   

7.
Each muscle of the third metasomal segment of the male of Nomia melanderi Ckll. is described in detail. The points of attachment of each muscle are compared with their homologs in other pregenital segments and with their homologs in the female. The function desgnated for each muscle describes its action alone or in conjunction with other muscle(s). New names are given to genital muscles by referring in the name to their points of attachment. Each intratergal muscle has homologous points of attachment in the pregenital segments of both sexes. The median tergo-dorsoplical muscle of the seventh segment and the oblique tergo-dorsoplical muscle of the eighth segment have changed their points of attachment. The intrasternal muscles are modified to suit the needs of courtship and mating, thus they are different from their homologs in the female. The spiracular muscles are well developed in all segments except the eighth, where the sterno-spiracular muscle is absent. The extrinsic genital muscles are derived from the intrasternal muscles of the eighth and ninth segments. The parameral and volsellar muscles are reduced in number. The aedeagal muscles, except the aedeago-phallic, have retained similar points of attachment to those found in primitive Hymenoptera. The topography of the metasomal nervous system is reported in detail by following each nerve and nervule to its termination. The study shows that (at least in Nomia) the criterion of nerve-concentration should not be used alone to indicate evolutionary levels. To accommodate the morphological changes in the terminal segments the Anterior and Posterior Lateral Nerves have migrated to new locations. The pattern of nerve topography (even at the nervule level) is homologous both in the different pregenital segments and between the sexes. The fact that homology does not exist between the external genitalia of the male and the modified ovipositor of the female supports the thesis that the male genital capsule is of phallic rather than prephallic origin. A pair of intersegmental membrane glands located between the seventh and eighth sterna is described. These glands may be the source of a pheromon responsible for gregariousness among “sleeping” males.  相似文献   

8.
Although the great genital diversity of the barklouse genus Trichadenotecnum has been described in previous studies, the specific function of the genital structures during the copulation process has received less investigative attention. We reconstructed 3D-models of each structure of the male and female genitalia of Trichadenotecnum incognitum in copula and those of uncopulated male and female of Trichadenotecnum pseudomedium. By comparing the changes in male and female genital structures and related muscles in copulated and uncopulated states, the function of each genital structure can be described. During the copulation, we found that the female subgenital plate was hooked into the male body by the distal process on the male paraproct and was fixed by the male epiproct, hypandrium and phallosome. In addition, sexual coevolution was suggested by tightly contacting structures, that is, thorny male hypandrium and thickened membrane around the female spermapore plate. These results not only give us a new understanding copulation process of Trichadenotecnum, but also explain the reasons why genital structures are so divers in the genus.  相似文献   

9.
The autosomal mutations transformer (tra) and transformer-2 (tra-2) of Drosophila convert chromosomal females (X/X) into phenotypical males. Our analysis aims at an understanding of the role which the transformer genes play in the development of the sexually dimorphic genital disc. In each Drosophila embryo, this disc starts development with a male and a female genital primordium, and an anal primordium. Our experiments involved the production of cell clones that were made homozygous for tra or tra-2 at different times of development. Homozygous clones were obtained by inducing mitotic recombination in three types of females heterozygous for tra or tra-2. The cells of the homozygous tra/tra or tra-2/tra-2 clones responded by changing from the female into the male pathway. Male genital structures developed if the clones were induced not later than 81 hr into development. In the analia, male clones appeared up to 120 hr. Our results show that the action of the wild-type alleles of tra+ and tra-2+ is required until late in larval development to repress the male genital primordium and to support development of the female primordium, as well as to maintain the anal primordium in the female pathway. Our data also suggest that the embryonic genital disc consists of two compartments, one containing the precursors for penis and analia, the other those of the male and female genitalia.  相似文献   

10.
Sasabe M  Takami Y  Sota T 《Molecular ecology》2010,19(23):5231-5239
Animals with internal fertilization often exhibit marked diversification in genital morphology among closely related species. However, our knowledge of the genetic architecture underlying genital evolution is still limited. We constructed genetic linkage maps and analysed quantitative trait loci (QTL) for F2 hybrids of two closely related species of the carabid beetles Carabus (Ohomopterus) iwawakianus and C. (O.) maiyasanus, which show matching male and female genital shapes within species, but marked differences in genital morphologies between species. The linkage maps comprised both amplified fragment length polymorphism and microsatellite markers. Composite interval mapping to detect QTL for three traits of male copulatory piece (length, width, weight) and two traits for female vaginal appendix (length, width) resulted in the detection of one to five significant QTL for each trait. The QTL explained large proportions of phenotypic variance. Thus, the interspecific difference in the genital morphologies appeared to be determined by relatively small numbers of genes with large genetic effects. QTL of different traits for the same or different sexes co‐occurred on five of eight linkage groups with significant QTL; in particular, three QTL for different male and female genital traits occurred almost at the same position. Each of the male genital traits showed uniform signs of additive genetic effects, suggesting that directional selection has led to species‐specific morphologies. However, the signs of additive genetic effects in each female genital trait were not uniform, suggesting that coevolution between sexes is not necessarily concerted. This result requires further assessment because the sample size of F2 females was small.  相似文献   

11.
Summary The genital imaginal disc ofDrosophila differentiates the terminalia, i.e. the genitalia and analia, of both sexes. It represents a composite anlage, containing a female genital primordium, a male genital primordium and an anal primordium. In normal males and females, only one of the two genital primordia differentiates; the other is developmentally repressed. Therefore, cell-lineage relationships between the male and female genital primordia can only be studied in sexual mosaics which differentiate female and male cells. We producedMinute (M)non-Minute(M+) gynandromorphs and selected those with sexually mosaic terminalia for a cell-lineage analysis. In these mosaics, either the male (XO) or female (XX) cells wereM + and thus had a growth advantage. The differential growth rates served as a tool to detect clonal restrictions. In control gynandromorphs (M +M +), the amount of female genitalia differentiated was largely independent of the amount of male genitalia present. In contrast, male and female anal structures, as a rule, added up to one full set. The same was true for the experimentalMM + gynandromorphs, but the contribution ofXX andXO cells to mosaic terminalia changed drastically due toM + cells competing successfully against the more slowly growingM cells. Specific subsamples ofMM + gynandromorphs showed thatM cells in a non-mosaic primordium are shielded from cell competition taking place in the neighbouring mosaic primordium. We conclude that the three primordia of the genital disc represent developmental compartments. In the genital primordia, even developmentally repressedM + cells compete successfully against developmentally activeM cells.  相似文献   

12.
The coremata and male genital structures together with the associated muscles of Zygaena trifolii are described and illustrated in detail. Of the female genitalia only the external parts are dealt with. The closure of the male coremata pouch was found to be effected by a special muscle. Mating pairs were quickly fixed while in copula and dissected afterwards in order to examine the mechanism of copulation. It turns out that the female is held by the male at three points: first, the gonopods (= valvae) grip the female abdominal tip laterally. Secondly, the praephallus together with the everted endotheca (= vesica) are tightly locked inside the female ductus bursae. Thirdly, female postvaginal structures become evaginated and are then held between the spined anellus and the eighth abdominal tergum of the male.  相似文献   

13.
The rapid divergence of genital morphology is well studied in the context of sexual selection and speciation; however, little is known about the developmental mechanisms underlying divergence in genitalia. Ground beetles in the subgenus Ohomopterus genus Carabus have species‐specific genitalia that show coevolutionary divergence between the sexes. In this study, using X‐ray microcomputed tomography, we examined the morphogenesis of male and female genitalia in two closely related Ohomopterus species with divergent genital morphologies. The morphogenetic processes generating the male and female genitalia at the pupal stage were qualitatively similar in the two species. The male aedeagus and internal sac and female bursa copulatrix were partially formed at pupation and developed gradually thereafter. The species‐specific genital parts, male copulatory piece, and female vaginal appendix differed in the timing and rate of development. The relatively long copulatory piece of Carabus maiyasanus began to develop earlier, but subsequent rates of growth were similar in the two species. The timing of the formation of the vaginal appendix and initial growth rates were similar, but subsequent rapid growth led to a longer vaginal appendix in C. maiyasanus. Thus, substantial interspecific differences in the size of genital parts were mediated by different underlying developmental mechanisms between the sexes (i.e., a shift in the developmental schedule in males and a change in growth rate in females). These results revealed the spatio–temporal dynamics of species‐specific genital structure development, providing a novel platform for evo–devo studies of the diversification of genital morphologies.  相似文献   

14.
The contemporary explanation for the rapid evolutionary diversification of animal genitalia is that such traits evolve by post‐copulatory sexual selection. Here, we test the hypothesis that the male genital spines of Drosophila ananassae play an adaptive role in post‐copulatory sexual selection. Whereas previous work on two Drosophila species shows that these spines function in precopulatory sexual selection to initiate genital coupling and promote male competitive copulation success, further research is needed to evaluate the potential for Drosophila genital spines to have a post‐copulatory function. Using a precision micron‐scale laser surgery technique, we test the effect of spine length reduction on copulation duration, male competitive fertilization success, female fecundity and female remating behaviour. We find no evidence that male genital spines in this species have a post‐copulatory adaptive function. Instead, females mated to males with surgically reduced/blunted genital spines exhibited comparatively greater short‐term fecundity relative to those mated by control males, indicating that the natural (i.e. unaltered) form of the trait may be harmful to females. In the absence of an effect of genital spine reduction on measured components of post‐copulatory fitness, the harm seems to be a pleiotropic side effect rather than adaptive. Results are discussed in the context of sexual conflict mediating the evolution of male genital spines in this species and likely other Drosophila.  相似文献   

15.
Static allometry of sexually selected traits has been the subject of intense research recently. However, some pitfalls for this kind of research are: (a) the functions of sexual traits are largely unknown; (b) more than one body size indicator must be measured; and, (c) allometry must be examined under different environmental circumstances to see whether allometric values change. Using Hetaerina americana damselflies, we investigated the type of allometry exhibited by a wing red spot and aedeagal width. These traits are positively selected during pre-copulatory male-male contests and post-copulatory female stimulation, respectively. As body size indicators, we used wing length and head width. It has been documented that expression of both sexual traits varies throughout the year. Thus, allometry was examined in different times of the year. We also investigated the allometry of aedeagal width and vaginal width at the zone where female stimulation takes place. We found no clear pattern of any allometric relationship for male and female traits and for both body size indicators at all times sampled. Our results contrast with patterns of negative allometry exhibited by genital traits in other animals.  相似文献   

16.
Males of several insect species inflict wounds on female genitalia during copulation. Such copulatory wounding also occurs in the fruit fly Drosophila melanogaster Meigen, 1830, one of the most important model organisms. Using a flash fixation technique with mating pairs of D. melanogaster, I examined the use and functions of the male phallic organ within the female reproductive tract. Paired components of the phallic organ (gonopods and two pairs of branches of the basal processes of the aedeagus) opened sequentially, from outer to inner components, during copulation. The dorsal branches of the aedeagal basal processes pierced the intima of the female reproductive tract at the lateral shallow folds. Consequently, mated females usually had a pair of melanized patches from repaired copulatory wounds. The sites that were stabbed by the dorsal branches were also clutched on the outside of the female oviscape (ovipositor) by the posterior process, which is a component of the periphallic organ. These structures likely function together as a mate-holding device. Male ejaculate labeled with rhodamine-B fluorescent dye entered the copulatory wounds in D. eugracilis Bock and Wheeler (Univ Texas Publ 7213:1–102, 1972), a related species, but not in D. melanogaster. Thus, copulatory wounds may function as an entrance for male seminal chemicals into the female circulatory system in D. eugracilis, but might not in D. melanogaster.  相似文献   

17.
The aedeagal gland of male Tenebrio molitor consists of numerous acini containing several secretory units (organules) of three epithelial cells in series. The distal cortical cell and intermediate cell are secretory cells. Secretory products are passed into microvilli-lined extracellular reservoirs. From these storage areas products flow through minute canaliculi and into the efferent ductule. Canaliculi, cuticular trabeculae, and fibrillar material are characteristic features of the efferent ductules within the extracellular reservoirs of secretory cells. After passing from the secretory cells, the efferent ductule penetrates the basal ductule cell. The thin epicuticle that comprises the wall of the ductule is confluent with the epicuticle of the cuticular sheath forming the wall of the genital pocket. Secretory products flow from the cortical cell ductule into the intermediate cell and eventually empty into the genital pocket. A chemical reaction apparently takes place in the intermediate cell ductule, resulting in a frothy secretion product. When released from the ductule, this frothy product forms a foam-like layer that coats the inner wall of the genital pocket. Ultrastructural and probable functional aspects of this gland are described and discussed.  相似文献   

18.
One of the most sweeping of all patterns in morphological evolution is that animal genitalia tend to diverge more rapidly than do other structures. Abundant indirect evidence supports the cryptic female choice (CFC) explanation of this pattern, which supposes that male genitalia often function to court females during copulation; but direct experimental demonstrations of a stimulatory function have been lacking. In this study, we altered the form of two male genital structures that squeeze the female’s abdomen rhythmically in Glossina pallidipes flies. As predicted by theory, this induced CFC against the male: ovulation and sperm storage decreased, while female remating increased. Further experiments showed that these effects were due to changes in tactile stimuli received by the female from the male’s altered genitalia, and were not due to other possible changes in the males due to alteration of their genital form. Stimulation from male genital structures also induces females to permit copulation to occur. Together with previous studies of tsetse reproductive physiology, these data constitute the most complete experimental confirmation that sexual selection (probably by CFC) acts on the stimulatory properties of male genitalia.  相似文献   

19.
The rapid evolution of male genital morphology is a characteristic feature of several animal groups. Such rapid divergence makes this trait a useful key for species identification. The aedeagus, the intromittent organ of male genitalia, is considered the main diagnostic trait in the Drosophila repleta group. In this study we analysed phenotypic plasticity and genetic variations associated with aedeagus size and shape in the cactophilic sibling species Drosophila gouveai Tidon‐Sklorz and Sene, 2001 and Drosophila antonietae Tidon‐Sklorz and Sene, 2001. Phenotypic plasticity in aedeagus morphology was evaluated in terms of the response to rearing media prepared with each species’ natural host plant, Pilosocereus machrisii Dawson, 1957 and Cereus hildmannianus Schum, 1890 respectively. Our results show that aedeagal shape differed significantly between species and that both shape and size presented host‐related phenotypic plasticity in both species. Flies reared on P. machrisii had, on average, larger aedeagi than those grown in C. hildmannianus. The general shape of aedeagus also differed significantly between flies that emerged in different host cactus. Patterns of variation in aedeagus morphology are discussed in the light of the current knowledge of evolutionary relationships and host plant use, in the D. buzzatii cluster, an assemblage of species in active cladogenesis.  相似文献   

20.
Typically, sexually selected traits show positive allometry and high coefficients of variation (CV). To date, many studies on the allometry of genitalia have focused on insects. In addition, studies have largely ignored the potential for sexual selection on female genitalia, despite male and female structures presumably co-evolving. Insects tend to show negative allometry in both male and female genitalia, while in contrast, the few studies carried out in mammals (males only) show positive allometry. Reasons for these differences between the taxa still remain unclear. However, in mammals, three main mechanisms have been proposed for genital evolution, namely, sperm competition, female cryptic choice and sexual conflict. In the first such study that we are aware of, we examined intra-specific genital allometry in both males and females of a mammal, the subterranean solitary Cape dune mole-rat, Bathyergus suillus. We found positive allometry occurring in male genitalia, which is consistent with previous vertebrate studies. Similarly, we found that female genitalia also exhibited positive allometry further supporting the notion of co-evolution of male and female genitalia. Although it is difficult to distinguish between the forces or mechanisms determining this directional selection, we suggest that several reproductive advantages are incurred as a result of positive allometric relationship of the genitalia in B. suillus and such advantages are also likely in other subterranean mammals. Our study further highlights the differences in genital allometry across taxa.  相似文献   

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