Interference of asexual and sexual reproduction in the green hydra

The green hydra, Hydra viridissima, has three sexes: hermaphrodite, male, and female. I investigated the reproductive strategies of the green hydra and the relationship between asexual budding and sexual reproduction. The proportion of mature individuals in the asexually reproducing population increased with increasing temperature. Sexual reproduction did not interrupt asexual budding in hermaphrodites or males; sexual reproduction did interrupt asexual budding in females. Sexual reproduction also resulted in exponential population growth during the reproductive season. The number of asexual buds on each parental individual was positively correlated with the parental individual size in asexual individuals and in males. The same positive correlation was found between the number of testicles and the size of males. These correlations reflect a common tendency in asexual and sexual reproduction: larger parental individuals have a greater number of propagules or gametes. No correlation was found between size and buds or size and gonads in hermaphrodites; hermaphrodites had at most one asexual bud and were significantly larger than males, females, and asexual individuals. The larger size of hermaphrodites supports the hypothesis that producing both female and male gonads is more energetically costly than producing only one type of gamete (gonochorism).     

Inherited XX sex reversal in the cocker spaniel dog

Summary Nine XX true hermaphrodites and two XX males were discovered in a family of American cocker spaniels. The true hermaphrodites were partially-masculinized females with ovotestes; the XX males had malformed male external genitalia and cryptorchid aspermatogenic testes. Wolffian and Mullerian duct derivatives were present in both true hermaphrodites and XX males. All four sires of sex-reversed dogs were normal XY males; five of the dams were anatomically normal females and one was an XX true hermaphrodite. A second true hermaphrodite reproduced as a female, producing anatomically normal offspring.All matings that produced sex-reversed offspring were consanguineous. Matings of the parents of sex-reversed cocker spaniels to normal beagles with no family history of intersexuality produced only normal offspring. Examination of G-banded karyotypes of the affected animals, their parents, and siblings, revealed no structural anomalies of the chromosomes that were consistently associated with sex-reversal.In assays for serologically-detectable H-Y antigen, the group of XX true hermaphrodites and the group of XX males had mean levels of the antigen not significantly different from that in normal male controls. Female parents of sex-reversed dogs and some of their female siblings were typed H-Y antigen positive, but the mean level of the antigen in this group was less than that of normal male controls.It is proposed that XX sex reversal in cocker spaniels is due to a mutant gene which when homozygous in females, results in a level of H-Y antigen similar to that found in normal males and the gonads develop as ovotestes or testes. When the gene is heterozygous in females, the level of serologically-detectable H-Y antigen is lowr than that found in normal males and the gonads develop as normal ovaries. The persistence of Mullerian structures in the presence of testicular tissue suggests that Mullerian inhibiting substance is deficient or ineffective in its action in this condition.Supported by NIH Postdoctoral Fellowship IF32 HL05515, University of Pennsylvania Genetics Center Grant, No. GM 20138, and NIH grants AI-19456, HD 17049, and HD 14357; and a grant from the Mrs. Cheever Porter Foundation.     

XX sex reversal in the American cocker spaniel dog: phenotypic expression and inheritance

Summary This study was conducted to define the range of phenotypic expression and mode of inheritance of XX sex reversal in the cocker spaniel dog. Breeding experiments produced F1, F1BC, and F2 generations in which 29 XX true hermaphrodites and 3 XX males were defined by chromosome constitution, serial histologic sections of the gonads, and examination of the internal and external genitalia. In XX true hermaphrodites, the most common combination of gonads was bilateral ovotestes, followed by ovotestis and ovary, then ovotestis and testis. The amount of testicular tissue in the two gonads was closely correlated within each true hermaphrodite. The distribution of testicular tissue within ovotestes of true hermaphrodites was consistent with the hypothesis that testicular differentiation is initiated in the center of the gonad and spreads outward. XX males had bilateral aspermatogenic testes and the internal ducts and external genitalia were more masculinized than in true hermaphrodites. Results of breeding experiments are consistent with autosomal recessive inheritance, the affected phenotype being expressed only in dogs with an XX chromosome constitution. The phenotypic expression and mode of inheritance of this disorder is compared to XX sex reversal in humans and other animals.     

Sexuality and hermaphroditism in fishes. I. Synchronous functional hermaphroditism in the serranid fish Serranus scriba L

Anatomical and histological examination of Serranus scriba L. showed the existence of primary females (67%), hermaphrodites (31%) and primary males (2%). Synchronous functional hermaphroditism is described on the basis of an anatomical and histological study of the gonads. Although they function simultaneously, the testicular and ovarian parts of hermaphrodite gonads have completely separate ducts. Females and hermaphrodites have the same annual reproduction cycle. In hermaphrodites, the testicular part matures one month sooner than the ovarian part. Cross fertilization between primary females and hermaphrodite individuals and between two different hermaphrodites probably occurs, while self-fertilization is less likely. The testicular tissues of primary males are of the acinar type and those of hermaphrodites are of the radial type. It is possible that primary males do not take part in reproduction. Serranus scriba in Egyptian Mediterranean waters is a longperiod spawner, which spawns from June to the end of October, i.e. it is a summer-autumn spawner.     

Disorders of sexual development and associated changes in the pituitary-gonadal axis in dogs

Normal sexual differentiation depends on completion of chromosomal sex determination, gonadal differentiation, and development of the phenotypic sex. An irregularity in any of these three steps can lead to a disorder in sexual development (DSD). We examined nine dogs with DSD by abdominal ultrasonography, laparotomy, histologic examination of the gonads, and reproductive tract, cytogenetic analysis, and mRNA expression of the SRY gene. We also determined the plasma concentrations of luteinizing hormone (LH), estradiol-17β, and testosterone before and after administration of gonadotropin-releasing hormone (GnRH) and compared these results with those obtained in anestrous bitches and male control dogs. The gonads of three dogs with DSD contained both testicular and ovarian tissue, while in the other six only testicular tissue was found. Each of the dogs had a uterus. Based on gynecologic examination, cytogenetic analysis, and the histology of the gonads, seven of the nine dogs appeared to be XX sex reversals. Three of these were XX true hermaphrodites and four were XX males; the other two dogs had incomplete XY gonadal dysgenesis. All seven XX sex-reversed dogs were found to be negative for the SRY gene by polymerase chain reaction. The basal plasma luteinizing hormone (LH) concentration was significantly higher in dogs with DSD than in anestrous bitches but not significantly different from that in male dogs. The basal plasma LH concentration increased significantly after GnRH administration in all dogs with DSD. The basal plasma estradiol concentration was significantly higher in dogs with DSD than in anestrous bitches but not significantly different from that in male dogs. The basal plasma testosterone concentration was lower in dogs with DSD than in male dogs. In all dogs with DSD both the basal and GnRH-induced plasma testosterone concentrations were above the upper limit of their respective ranges in the anestrous bitches. In conclusion, the secretion of LH and estradiol in these dogs with DSD, all of which had testicular tissue in their gonads, was similar to that in male control dogs. These results indicate that the basal and/or GnRH-stimulated plasma testosterone concentration might be used to detect the presence of testicular tissue in dogs with DSD.     

Variation in reproductive success and gonadal allocation in the simultaneous hermaphrodite,Serranus fasciatus

Summary This paper examines the correlates of individual size, reproductive success, gonadal allocation, and growth in a hermaphroditic reef fish. Individuals in S. fasciatus mature as simultaneous hermaphrodites; large individuals subsequently lose female function and become functional males. Daily female reproductive success was highly correlated with both hermaphrodite size and amount of female gonadal tissue. Three separate comparisons gave a positive correlation between male reproductive success and male gonadal allocation: (1) Males had higher levels of male gonadal allocation and male reproductive success than hermaphrodites. (2) The percent of gonad allocated to male tissue in hermaphrodites was higher in the year they had higher male mating success. (3) Male gonadal tissue of hermaphrodites was positively correlated with male reproductive success in the year that male reproductive success by hermaphrodites was higher and more variable. There was no evidence for a trade-off between male function, female function, and growth among hermaphrodites. Many of these patterns have also been observed in plants, but the selective pressures leading to these patterns in S. fasciatus and plants are probably quite different.     

Physiological and reproductive differences between hermaphrodites and males in the androdioecious plant Fraxinus ornus

Androdioecy, the co-occurrence of males and hermaphrodites in a breeding population, is a rare reproductive system in the nature. This rarity may be the result of the large fitness gain required for male plants to be maintained by selection. Physiological, vegetative and reproductive characters of males and hermaphrodites of the androdioecious species Fraxinus ornus (Oleaceae) were compared, supporting the hypothesis that males compensate the fitness advantage of hermaphrodites with greater reproductive -but not vegetative- output, with concomitant differences in physiological capacities between the genders. Photosynthetic rate was similar between both genders, but hermaphrodites had lower water potential and carbon isotope discrimination than males. Photosynthesis rates decreased with decreasing water potentials more steeply in males than in hermaphrodites, indicating that hermaphrodites were more drought tolerant than males. Vegetative characters such as current year shoot growth or tree size did not differ between genders. Males produced 1.6 times more inflorescences than hermaphrodites. This difference was consistent across years and populations.     

Sex allocation in a simultaneously hermaphroditic marine shrimp

Two fundamental questions dealing with simultaneous hermaphrodites are how resources are optimally allocated to the male and female function and what conditions determine shifts in optimal sex allocation with age or size. In this study, I explored multiple factors that theoretically affect fitness gain curves (that depict the relationship between sex-specific investment and fitness gains) to predict and test the overall and size-dependent sex allocation in a simultaneously hermaphroditic brooding shrimp with an early male phase. In Lysmata wurdemanni, sperm competition is absent as hermaphrodites reproducing in the female role invariably mated only once with a single other shrimp. Shrimps acting as females preferred small over large shrimps as male mating partners, male mating ability was greater for small compared to large hermaphrodites, and adolescent males were predominant in the population during the breeding season. In addition, brooding constraints were not severe and varied linearly with body size whereas the ability to acquire resources increased markedly with body size. Using sex allocation theory as a framework, the findings above permitted to infer the shape of the male and female fitness gain curves for the hermaphrodites. The absence of sperm competition and the almost unconstrained brooding capacity imply that both curves saturate, however the male curve levels off much more quickly than the female curve with increasing level of investment. In turn, the predominance of adolescent males in the population implies that the absolute gain of the female curve is greater than that of the male curve. Last, the size-dependent female preference and male mating ability of hermaphrodites determines that the absolute gain of the male curve is greater for small than for large hermaphrodites. Taking into consideration the inferred shape of the fitness gain curves, two predictions with respect to the optimal sex allocation were formulated. First, overall sex allocation should be female biased; it permits hermaphrodites to profit from the female function that provides a greater fitness return than the male function. Second, sex allocation should be size-dependent with smaller hermaphrodites allocating more than proportionally resources to male reproduction than larger ones. This size-dependent sex allocation permits hermaphrodites to profit from male mating opportunities that are the greatest at small body sizes. Size-dependent sex allocation is also expected because the male fitness gain curve decelerates more quickly than the female gain curve and experiments indicated that resources are greater for large than small hermaphrodites. These two predictions were tested when determining the sex allocation of hermaphrodites by dissecting their gonad and quantifying ovaries versus testes mass. Supporting the predictions above, hermaphrodites allocated, on average, 118 times more to the female than to the male gonad and the proportion of resources devoted to male function was higher in small than in large hermaphrodites. A trade-off between male and female allocation is assumed by theory but no negative correlation between male and female reproductive investment was observed. In L. wurdemanni, the relationship between sex-specific investment and fitness changes during ontogeny in a way that is consistent with an adjustment of sex allocation to improve size-specific reproductive success.     

The evolution of males: support for predictions from sex allocation theory using mating arrays of sagittaria latifolia (alismataceae)

Investment in male function should often yield diminishing fitness returns, subjecting the evolution of male phenotypes to substantial constraints. In plants, the subdivision of male function via the gradual presentation of pollen might minimize these constraints by preventing the saturation of receptive stigmas. Here, we report on an investigation of (1) patterns of investment in male function by plants in hermaphroditic (monoecious) and dioecious populations of Sagittaria latifolia, and (2) patterns of siring success by males versus hermaphrodites in experimental mating arrays. We show that in natural populations, males from dioecious populations had greater investment in male function than hermaphrodites in monoecious populations. However, as a proportion of total flower production, males presented substantially fewer flowers at once than hermaphrodites. In comparison with hermaphrodites, therefore, males prolonged the period over which they presented pollen. In mating arrays comprised of females, males, and hermaphrodites, siring success by males increased linearly with flower production. This finding is consistent with the existence of a linear gain curve for male function in S. latifolia and supports the idea that the gradual deployment of male function enables plants to avoid diminishing returns on the investment in male function.     

Non‐functional hermaphroditism in three species of Clupeiformes from Tokyo Bay,Japan

Intersexual gonads were detected in three clupeiform species, Sardinops melanostictus, Sardinella zunasi and Englaulis japonicus, collected in Tokyo Bay off Yokohama City, Japan, of which appearance rates were 18.2, 34.6 and 33.9 %, respectively. While spermatogenesis proceeded normally in these individuals, previtellogenic stage oocytes were detected outside of the testes in the hermaphrodites, appearing as a single cell or small clusters. The results suggest that these species are non‐functional hermaphrodites. As intersexual gonads have been reported in other clupeiform species collected before significant pollution and/or at sites far from urban influences, such non‐functional hermaphrodites may be common occurrence in the order. However, high rates of hermaphrodite appearance also suggest the possibility that the development of these oocytes were induced by environmental estrogens.     

Sexual selection and simultaneous hermaphroditism among the Unionidae (Bivalvia: Mollusca)

Simultaneous hermaphroditism is an infrequent mode of reproduction among bivalves of the family Unionidae: only five of the 220 North American species are simultaneous hermaphrodites. However, hermaphroditic individuals of otherwise predominantly dioecious species have been encountered in 30 of I01 species examined. These hermaphroditic individuals as well as simultaneous hermaphrodites can exhibit considerable variability in the ratio of spermatogenic: oogenic tissue within the gonad, and the purposes of this paper are to determine the underlying causes of both this variability and the occurrence of occasional hermaphroditic individuals among dioecious species. Results indicate that the ratio of male: female gonodal tissue of a simultaneous hermaphrodite is bimodally distributed, and several hypotheses to account for this observation are presented. It is proposed that populations occurring in different habitats and under conditions of different individual density are subject to fundamentally different sexual selection pressures acting on ratios of allocation to male and female gametes. Occasional hermaphroditism among otherwise predominantly dioecious species was in this study associated with infection of the gonads by digenean trematodes. A model of sexual determination among unionids presented in this paper proposes that sex is determined by genetically controlled hormone levels: occasional hermaphrodites result from alterations in these hormone levels caused by developmental errors and trematodal infections which mimic the results of such errors. Predictions of this model are consistent with observed levels of variability in male: females gonadal tissue among occasional hermaphrodites.     

Male–Male Competition in a Mixed‐Mating Fish

Mating systems that comprise a mixture of pure males and self‐fertilising hermaphrodites remain an evolutionary enigma. In particular, our understanding of the sexual selection pressures associated with such mating systems is nascent. Males can only reproduce by fertilising hermaphrodites’ eggs, but hermaphrodites can also fertilise their own eggs and gain a genetic advantage by doing so. Consequently, there should be intense competition among males to access hermaphrodites. Here, we test the importance of male size, colour and heterozygosity in predicting the outcome of male–male competition using the mangrove rivulus, which has a male‐hermaphrodite mixed‐mating system. We pitted males against one another in dyadic laboratory trials to develop a dominance score for each male. We then correlated these scores with male length, several components of male colour, and heterozygosity. Male size was the only significant correlate of dominance: larger males dominated smaller males, implying selection for large male size. However, male mangrove rivulus are similar in size to hermaphrodites, indicating that directional selection for large body size in males is no greater than it is in hermaphrodites. Across all trials, colour was unrelated to dominance, but contests between similarly sized males were usually won by more colourful individuals. As mangrove rivulus are dichromatic, we suspect that male colour may prove to be more important in mate choice than we found it to be in intrasexual competition. Heterozygosity did not explain dominance directly, but correlated strongly with male size, implying an indirect role in intrasexual competition.     

Biology of the amphioxus,Branchiostoma belcheri in the Ariake Sea,Japan II. Reproduction

We investigated the reproduction of the amphioxus Branchiostoma belcheri in the southern Ariake Sea, Japan, from 1999 to 2002. Gonads were very small or almost absent from September to December and began to develop in January. Changes in the gonad length index indicated that spawning began in mid June, after the gonads had attained maximum size. Although most gametes were extruded during the first spawning, some remained in the gonads. Shrunken gonads were much smaller but contained gametes, and the second spawning occurred around 10 July. It was unclear whether all adults spawned twice, but a large proportion of individuals did. After the first spawning, both males and females lost 30% of their body weight. Most one-year-old individuals did not spawn, and the minimum size at maturity was ca. 20 mm. We found no termination of reproduction among very large individuals. The average number of gonads was significantly larger on the right side of the body (26.2 in males and 26.1 in females) than on the left side (24.0 in males and 23.5 in females). We found two hermaphrodites in a total of 11,184 specimens examined. Each had four or three ovaries among a total of 45 or 54 gonads.     

Increased sperm allocation delays body growth in a protandrous simultaneous hermaphrodite

Sex allocation theory for simultaneous hermaphrodites predicts that hermaphrodites have a fixed amount of reproductive resources to allocate to both sex functions and that they trade off their allocation to both sexes depending on environmental conditions. Ophryotrocha diadema is a simultaneous hermaphroditic polychaete worm which has a protandrous phase prior to the hermaphroditic phase. The ability of adolescent males to compete with mature hermaphrodites for egg fertilization and the costs of an increase in male expenditure during the protandrous phase were tested in experiments where adolescent males had the possibility to fertilize hermaphrodites' eggs. Results document that adolescent males were able to compete with hermaphrodites for egg fertilization and caused the hermaphrodites fitness losses of 31.4%. Adolescent males which fertilized hermaphrodites' eggs had a slower growth rate than males which had no possibility to fertilize eggs. This result indicates that an increased male expenditure is a cost for adolescent males and that, during the protandrous phase, resources are allocated to reproduction at the expense of somatic development.  © 2003 The Linnean Society of London . Biological Journal of the Linnean Society , 2003, 78 , 149–154.     

Is there sperm storage in the clam shrimp Eulimnadia texana?

Abstract. Androdioecy is a rare form of mating system in which species comprise males and hermaphrodites. One recently described case of androdioecy is the freshwater crustacean Eulimnadia texana. A mathematical model of the mating system of this shrimp suggests that males and hermaphrodites should only coexist under limited circumstances. One possible factor not considered in this model would extend the conditions for coexistence: the possibility of sperm storage in the hermaphrodites. Here we use genetically marked matings between males and hermaphrodites to determine if hermaphrodites can store male sperms. Eggs were collected from hermaphrodites both in the presence of a male and after the male was removed. A total of 30 of these matings had successful hatches, but only 14 of these 30 could be used to test for sperm storage. In these 14 cases, an average of 35% of the eggs were outcrossed when males were present, but only 0.4% were outcrossed after males were removed. Thus, sperm storage by hermaphrodites was an insignificant factor in the production of offspring. These data suggest that sperm storage cannot help explain the coexistence of males and hermaphrodites in natural populations of this crustacean.     

High male reproductive success of hermaphrodites in the androdioecious Phillyrea angustifolia

Androdioecy, the coexistence of males and hermaphrodites within a population, is a rare breeding system, often considered as unlikely to evolve because of restrictive conditions for its maintenance. Phillyrea angustifolia, a wind-pollinated shrub, is one of the handful species reported to be androdioecious. Our previous studies have shown that natural populations of this species in southern France exhibit higher male frequencies (approximately 50%) than predicted on theoretical grounds. Thus, the male functionality of hermaphrodites is still debated. To assess the functional breeding system of this species in the wild, a paternity analysis was performed with two highly polymorphic microsatellite loci on 729 seeds collected on 10 maternal shrubs in a natural population of 24 mature individuals of P. angustifolia. A large proportion of seeds were found to have been sired by pollen from outside the population. Analysis of seeds sired by individuals within the study population revealed a high male fertility of hermaphrodites resulting in a low male advantage in fertility for male plants. Intermate distances were found to have a strong impact on male reproductive success, whereas sexual morph had no effect, with males and hermaphrodites performing equivalently. This study is the first to unequivocally document the occurrence of a male function of hermaphrodites in a natural population of an androdioecious species.     

AN ECOLOGICAL GENETIC STUDY OF GYNODIOECY IN LIMNANTHES DOUGLASII (LIMNANTHACEAE)

Gynodioecy in two populations of Limnanthes douglasii var. rosea was studied for its maintenance requirements, role in population structure and its influence on the levels of outbreeding. The mode of inheritance of male sterility appeared to be nucleo-cytoplasmic, although the precise number of nuclear genes and dominance relationships could not be ascertained. Measurements on the plants sampled from natural stands and controlled experiments showed that male-steriles and their progeny had greater biomass and more flowers per plant than hermaphrodites, though these results varied with the environmental conditions. The hermaphrodites of gynodioecious populations had higher rates of selfing than the hermaphrodite individuals in populations lacking male sterility. Estimates of the inbreeding depression and homozygosity levels were also higher in gynodioecious populations. Variation in these parameters of breeding system and relative heterozygosis among populations may explain why male sterility has a restricted distribution, as theoretical models also predict rather specific conditions for this stable polymorphism. These data suggest that the advantage of male sterility is associated with lowered inbreeding depression. However, the potential ecological resource reallocation to the female function needs to be investigated. The fitness differences observed in this study between male-steriles and hermaphrodites appear inadequate to maintain the nucleo-cytoplasmic male sterility but could account for the observed frequencies (10–20%) of male-steriles in nature if the genetic system has evolved from a cytoplasmic system.     

Males outcompete hermaphrodites for seed siring success in controlled crosses in the polygamous Fraxinus excelsior (Oleaceae)

Polygamy (including trioecy and subdioecy), the co-occurrence of males, hermaphrodites, and females in natural populations, is a rare and poorly studied breeding system expressed in Fraxinus excelsior L. (Oleaceae), a wind-pollinated tree. Here we investigate siring ability of pollen from male vs. hermaphrodite individuals to better understand this sex polymorphism. We conducted single-donor and two-donor pollination experiments and compared both fruit set and seed siring success, assessed with polymorphic microsatellite markers, of male and hermaphrodite individuals. Single pollen donor crosses allowed us to verify the male function of hermaphrodites. However, pollen from hermaphrodites was much less proficient than male pollen, with males siring 10 times as many fruits in single donor pollination treatments. This result was strengthened by the surprisingly low reproductive success of hermaphrodites in pollen competition conditions: of the 110 seedlings analyzed three were selfed and only one was sired by the hermaphrodite donor. The remaining 106 were sired by the male pollen donor. These results raise the question of the maintenance of male fertility in hermaphrodites in Fraxinus excelsior. Male function of hermaphrodites in this species now needs to be assessed under field conditions.