Low siring success of females with an acquired male function illustrates the legacy of sexual dimorphism in constraining the breakdown of dioecy

Dioecy has often broken down in flowering plants, yielding functional hermaphroditism. We reasoned that evolutionary transitions from dioecy to functional hermaphroditism must overcome an inertia of sexual dimorphism, because modified males or females will express the opposite sexual function for which their phenotypes have been optimised. We tested this prediction by assessing the siring success of monoecious individuals of the plant Mercurialis annua with an acquired male function but that are phenotypically still female‐like. We found that pollen dispersed by female‐like monoecious individuals was ~ 1/3 poorer at siring outcrossed offspring than pollen from monoecious individuals with an alternative male‐like inflorescence. We conclude that whereas dioecy might evolve from functional hermaphroditism by conferring upon individuals certain benefits of sexual specialisation, reversion from a strategy of separate sexes to one of combined sexes must overcome constraints imposed by the advantages of sexual dimorphism. The breakdown of dioecy must therefore often be limited to situations in which outcrossing cannot be maintained and where selection favours a capacity for inbreeding by functional hermaphrodites.     

Sexual strategy in the freshwater bivalve Galatea paradoxa (Donacidae) from the Volta River estuary,Ghana

The reproductive biology, sexual strategy and sex ratio of the freshwater donacid Galatea paradoxa in the Volta River estuary, Ghana, was studied from March 2008 to February 2010. Histological observations of the gonads showed that G. paradoxa is gonochoristic with dominance by females (80%) and a high incidence of hermaphrodites (9.4%) for individuals ranged between 20 and 82?mm in shell length. There was a single spawning event between July and October. Gametogenesis started in December, progressing steadily to a peak in June–July when spawning began, until November when individuals were spent. The dominance of females coupled with the high incidence of hermaphroditism may be a reflection of populations subjected to increasing fishing pressure and over-exploitation.     

Histological profiling of gonads depicting protandrous hermaphroditism in Eleutheronema tetradactylum

The fourfinger threadfin Eleutheronema tetradactylum is reported as a protandrous hermaphrodite from Australian waters, while being a gonochorist in reports from Singapore and India, with a single report of protandrous hermaphroditism from the latter. Histological analysis of gonads of fish from Indian waters confirms protandrous hermaphroditism in E. tetradactylum. The study was based on 480 fish examined from eight locations along the Indian coast. Mean total length (L_T) of male fish was 240 mm with the transition to female starting from 280 mm L_T. Specimens confirmed as mature females were >380 mm L_T.     

An underground burst of diversity – a new look at the phylogeny and taxonomy of the genus Talpa Linnaeus, 1758 (Mammalia: Talpidae) as revealed by nuclear and mitochondrial genes

Using both nuclear and mitochondrial sequences, we demonstrate high genetic differentiation in the genus Talpa and confirm the existence of cryptic species in the Caucasus and Anatolia, namely, T. talyschensis Vereschagin, 1945, T. ognevi Stroganov, 1948, and Talpa ex gr. levantis. Our data support four clades in the genus Talpa that showed strong geographical associations. The ‘europaea’ group includes six species from the western portion of the genus' range (T. europaea, T. occidentalis, T. romana, T. caeca, T. stankovici, and T. levantis s.l.); another three groups are distributed further east: the ‘caucasica’ group (Caucasus), the ‘davidiana’ group (eastern Anatolia and Elburz) and T. altaica (Siberia). The phylogenetic position of T. davidiana was highlighted for the first time. The order of basal branching remains controversial, which can be attributed to rapid diversification events. The molecular time estimates based on nuclear concatenation estimated the basal divergence of the crown Talpa during the latest Miocene. A putative scenario of Talpa radiation and issues of species delimitation are discussed. © 2015 The Linnean Society of London     

Sex Allocation in Lythrypnus (Gobiidae): Variations on a Hermaphroditic Theme

The gonadal sexual patterns of three Atlantic/Caribbean Lythrypnus (L. nesiotes, L. phorellus, L. spilus) are described, based on histological examination, and compared with previously described sexual patterns of L. dalli and L. zebra, from the eastern Pacific. Overall, there was striking similarity in the sexual patterns of all the species; each exhibited simultaneous hermaphroditism with a high degree of variability in allocation to male tissue among individuals. Among the five species whose sexual patterns are described, at least three distinct sexual patterns were identified. Lythrypnus dalli has an allocation pattern distinct from the other species, with populations consisting of primarily pure females, strongly female-biased hermaphrodites and pure males. Lythrypnus nesiotes was also found to be distinct from the other species, consisting primarily of pure females and strongly female-biased hermaphrodites and male-biased individuals (mean allocation to male tissue = 75%). In contrast to L. dalli, L. nesiotes had no pure males. Indeed, no other species examined included pure males. L. phorellus, L. spilus and L. zebra had much more intermediate allocation patterns than either L. dalli or L. nesiotes. Most individuals were female biased (<10% male tissue), and there were significant numbers of individuals with intermediate allocation (10–90% male). L. zebra differed somewhat from L. phorellus and L. spilus in that more strongly male-biased individuals were common in the sample (i.e., ranging in allocation from 90 to 98% male); this difference was not statistically compelling. This marked variation in allocation among closely related species suggests Lythrypnus is a useful model system to explore factors affecting allocation. For example, models developed for simultaneous hermaphrodites and appropriate for Lythrypnus predict that pure sexed individuals will be more common at higher density (e.g., because mating success is more certain). The observed interspecific variation supports this prediction – both L. dalli and L. nesiotes occur at high density and have the most extreme allocation patterns, whereas L. phorellus, L. spilus, and L. zebra occur at lower density and have more intermediate allocation patterns.     

Early gonadal development and primary males in the protogynous epinepheline, Cephalopholis boenak

Early gonadal development of the protogynous epinepheline, Cephalopholis boenak, was examined histologically in 289 specimens with standard length (L_S) of 42–130 mm, collected from May 2000 to April 2002 in Hong Kong waters, to determine male developmental pathways and establish its sexual pattern. All juvenile gonads developed an ovarian lumen with primary‐growth stage oocytes and scattered spermatogenic tissue prior to sexual differentiation and first sexual maturation. From this bisexual phase containing both female and male tissues, some gonads differentiated as ovaries with further oocyte growth to cortical‐alveolus and vitellogenic stages, the rest differentiated as testes with the proliferation of spermatogenic tissue and the formation of a sperm sinus. All testes retained the lumen and primary‐growth stage oocytes, and sperm sinuses ran within the gonad wall. Unlike most protogynous species, among functional males it was impossible to distinguish those resulting from juveniles through sexual differentiation (i.e. primary male) from those resulting from functional females through sex change (i.e. secondary male) based solely on testicular morphology. A proportion‐spermatogenic‐tissue index (I_ST) was, therefore, developed and determined to be a reliable quantitative indicator for distinguishing differentiating, primary males before a sperm sinus was evident, from differentiating females during sexual differentiation. Since sexually transitional specimens with the concominant appearance of degenerating vitellogenic, or later, stage oocytes and spermatogenic tissue in the gonads were previously noted from Hong Kong, diandric, protogynous hermaphroditism is confirmed in C. boenak. For species, such as this and other epinephelines, in which all males have the same testicular morphology, a complete analysis of a wide range of body sizes from juveniles to adults is necessary for understanding male developmental pathways, and determining sexual pattern.     

Goat RSPO1 over-expression rescues sex-reversal in Rspo1-knockout XX mice but does not perturb testis differentiation in XY or sex-reversed XX mice

RSPO1 is a newly discovered gene involved in sex differentiation. Two goat BAC clones encompassing the RSPO1 gene (gRSPO1) were injected into mouse oocytes and several transgenic lines derived. Both clones induced gRSPO1 over-expression in various tissues, including male and female gonads, with no obvious phenotype and normal sex-ratios. Introgression of the gRSPO1 transgene into a mouse RSPO1 knockout genotype resulted in the rescue of the fertility and the disappearance of the masculinized gonadic features of the females, demonstrating the functionality of the goat protein in a mouse context. On the contrary, over-expression of gRSPO1 within a mSRY or a gSRY-XX genotypes did not interfere with the SRY-induced male phenotype. Laurine Buscara and Fatemeh Montazer-Torbati contributed equally to this work.     

Histology of the reproductive tissue of the clam Ruditapes decussatus from the gulf of Gabès (Tunisia)

Summary

The gametogenic cycle of the clam Ruditapes decussatus was investigated during one year. Samples were collected each month in two different sites located at the gulf of Gabès area. Gametogenic activity was characterized on the basis of histological analysis of the gonads. A classification and a scoring system were then proposed. According to our study, Ruditapes decussatus gametogenesis can be divided in four major steps: (1) a sexual rest (December- February), (2) starting of gametogenesis (February-March), (3) development (March-May), (4) ripe (from April onwards). This study revealed that the gonad of the clam Ruditapes decussatus was surrounded by muscular tissue which could represent the principal storage tissue. Two kinds of hermaphroditism appeared since maturity of gonads: simultaneous one showing gonadal tubules of both males and females in regionally distinct and separate zones and a “superposed” one showing a gonad consisting essentially of mature male gametes and few residual oocytes.     

Environmental condition related reproductive strategies and sex ratio in hydras

Kaliszewicz, A. and Lipińska, A. 2011. Environmental condition related reproductive strategies and sex ratio in hydras. —Acta Zoologica (Stockholm) 00 :1–7. Temperature and food supply appeared to affect sex ratio, sex composition and percentage of sexual individuals in three Hydra species: Hydra vulgaris, Hydra circumcincta and Hydra viridissima. We found three sexes present: females, males and hermaphrodites depending on environmental conditions. Hydra vulgaris appeared to be a species with a temperature‐dependent sex determination (TSD). The males and hermaphrodites were present only under rising temperatures, whereas females were observed exclusively at lowering temperatures. Hydras reproduced asexually at constant room temperature. Unlimited food affected sex ratios and induced the presence of males in H. circumcincta at lowering temperatures. Thus, H. circumcincta may be recognised as another Hydra species in which sex is determined by environmental factors (ESD). Under rising temperatures, the number of hermaphroditic individuals was higher when food supply was unlimited in all three species, indicating that hermaphrodites may need more energy to produce both male and female gonads. Both temperature changes and food supply positively affected asexual reproductive strategies in hydras, especially budding rates. Hydra circumcincta appeared to be less agile than other hydras and able to self‐fertilise. It is likely that self‐fertilisation is an adaptation to the low probability of meeting a mate belonging to the other clone.     

GENETIC DETERMINANTS OF PROTANDRIC SEX IN THE PACIFIC OYSTER,CRASSOSTREA GIGAS THUNBERG

A unique feature of sex in Crassostrea oysters is the coexistence of protandric sex change, dioecy, and hermaphroditism. To determine whether such a system is genetically controlled, we analyzed sex ratios in 86 pair-mated families of the Pacific oyster, Crassostrea gigas Thunberg. The overall female ratios of one-, two-, and three-year-old oysters were 37%, 55%, and 75%, respectively, suggesting that a significant proportion of oysters matured first as males and changed to females in later years. Detailed analysis of sex ratios in factorial and nested crosses revealed significant paternal effects, which corresponded to two types of sires. No major maternal effects on sex were observed. Major genetic control of sex was further indicated by the distribution of family sex ratios in two to four apparently discreet groups. These and other data from the literature are compatible with a single-locus model of primary sex determination with a dominant male allele (M) and a protandric female allele (F), so that MF are true males and FF are protandric females that are capable of sex change. The rate of sex change of FF individuals may be influenced by secondary genes and/or environmental factors. Strong maternal and weak paternal effects on sexual maturation or time of spawning were also suggested.     

Oestrogen blocks the nuclear entry of SOX9 in the developing gonad of a marsupial mammal

Background  

Hormones are critical for early gonadal development in nonmammalian vertebrates, and oestrogen is required for normal ovarian development. In contrast, mammals determine sex by the presence or absence of the SRY gene, and hormones are not thought to play a role in early gonadal development. Despite an XY sex-determining system in marsupial mammals, exposure to oestrogen can override SRY and induce ovarian development of XY gonads if administered early enough. Here we assess the effect of exogenous oestrogen on the molecular pathways of mammalian gonadal development.     

Patterns of gonad structure in hermaphroditic gobies (Teleostei Gobiidae)

Synopsis Hermaphroditism has been reported for a small number of gobiid fishes, but the extent of this sexual pattern within the family is not known. Gonad structure was examined in one or more species from twenty-one gobiid genera. No evidence of hermaphroditism was found in the species selected from 14 genera. Laboratory studies supported the conclusion of gonochorism for the examined species in four of them:Asterropteryx, Bathygobius, Gnatholepis, andPsilogobius. Currently, the absence of precursive testicular tissues associated with the ovary in females, in conjunction with no retained ovarian features in the testes of males, appear to be reliable indicators of a gonochoristic sexual pattern in gobiid fishes. Evidence for hermaphroditism was observed in seven genera:Eviota, Trimma, Fusigobius, Lophogobius, Priolepis, Gobiodon, andParagobiodon. Protogyny was experimentally confirmed inE. epiphanes, and the gonad structure in another nine of ten species ofEviota suggested either protogyny or protogynous tendencies. With the exception ofGobiodon andParagobiodon, which exhibited similar gonadal structure, ovarian and testicular structure varied considerably among the hermaphroditic genera examined, both with regard to the configuration and to the degree of development of ovarian and testicular tissues, or testicular tissue precursors. Findings of this study indicate that hermaphroditic gonad structure will prove to be a useful trait in determining evolutionary relationships within the Gobiidae.     

Flagellar setae of the second antennae in decapod shrimps: sexual dimorphism and possible role in detection of contact sex pheromones

Summary

The antenna 2 (antennal) flagella of decapod shrimps are chemotactile, and their setae are proposed as sensilla involved in recognition of females by males via a contact sex pheromone on the surface of the female. Male recognition of females receptive to mating occurs in many caridean species upon contact of male antennal flagella with the surface of a newly molted parturial female. The hypothesis of sexual dimorphism in the number and kind of setae on the antennal flagella of four caridean and one penaeid shrimp species was tested with setal counts and observations on setal morphology. Unique male antennal setae (“male-specific sensilla”) were not identified in any of the species investigated. However, the abundance of antennal setae was significantly greater in males than in breeding females in the palaemonid carideans Palemonetes pugio and Macrobrachium ohione. In the hippolytid caridean Thor manningi and alpheid caridean Alpheus normanni, no sexual dimorphism in setal abundance was demonstrated. In the penaeoid Rimapenaeus similis, males had a higher abundance of antennal setae than the larger breeding females but so did juvenile females, similar in size to males. The sexual dimorphism in antennal sensilla in the palaemonid species and its absence in A. normanni might be related to their different mating systems, but no such association is suggested for T. manningi and R. similis. Setal morphology suggestive of chemoreceptive function (a terminal pore) was observed in all species.     

Sex determination in humans

In mammals, the Y chromosome induces testis formation and thus male sexual development; in the absence of a Y chromosome, gonads differentiate into ovaries and female development ensues. Molecular genetic studies have identified the Y-located testis determining gene SRY as well as autosomal and X-linked genes necessary for gonadal development. The phenotypes resulting from mutation of these genes, together with their patterns of expression, provide the basis for establishing a hierachy of genes and their interactions in the mammalian sex determination pathway.     

Evolution of sex chromosomes prior to speciation in the dioecious Phoenix species

Understanding the driving forces and molecular processes underlying dioecy and sex chromosome evolution, leading from hermaphroditism to the occurrence of male and female individuals, is of considerable interest in fundamental and applied research. The genus Phoenix, belonging to the Arecaceae family, consists uniquely of dioecious species. Phylogenetic data suggest that the genus Phoenix has diverged from a hermaphroditic ancestor which is also shared with its closest relatives. We have investigated the cessation of recombination in the sex‐determination region within the genus Phoenix as a whole by extending the analysis of P. dactylifera SSR sex‐related loci to eight other species within the genus. Phylogenetic analysis of a date palm sex‐linked PdMYB1 gene in these species has revealed that sex‐linked alleles have not clustered in a species‐dependent way but rather in X and Y‐allele clusters. Our data show that sex chromosomes evolved from a common autosomal origin before the diversification of the extant dioecious species.     

The state of reproductive glands of the pike Esox lucius from water bodies of the Chernobyl trail in 1999–2004

The state of the reproductive system of the pike Esox lucius, descendants of individuals exposed to X-ray irradiation due to the catastrophe at the Chernobyl Nuclear Power Plant in 1986, was studied. The material was collected in water bodies of Ukraine contaminated with radionuclides—Kiev Reservoir, the Teterev River, and Lake Glubokoe in the postcatastrophe period 1999–2004. It was shown that the total number and the extent of disturbances in the pike gonads are positively correlated with the levels of contamination of the water bodies: in the cleanest water body, the Teterev River, the proportion of fish with gonads without considerable deviations was 70%; in the Kiev Reservoir, it was 45%; no individuals with normal gonads were recorded in Lake Glubokoe. The great number of anomalies in sexual cells and gonads has led to a decrease in the reproductive capacity of the pike populations studied. A unique case was revealed of synchronous hermaphroditism in a pike from the Kiev Reservoir that belonged to the fourth generation (F₄) of fish that underwent emergency loads. Among postemergency pike generations, the maximum number of disturbances in sexual cells and gonads was recorded in F_2–4, which is possibly accounted for by the phenomenon of “prolonged mutagenesis.”     

Characterization of the sexual pattern of Parastacus varicosus (Crustacea: Decapoda: Parastacidae)

Abstract. Freshwater crayfishes of the genus Parastacus are intersex, i.e., show characteristics of both sexes in the same individual; also, intersexuality has been documented in hermaphroditic species. The aim of this study was to analyze the gonads of Parastacus varicosus , characterizing its sexual pattern. The animals were collected at Cova do Touro, Gravataí, Rio Grande do Sul, Brazil. Three sexual forms were identified: intersex females, intersex males, and transitional specimens showing an ovotestes gonad that could only be identified by means of histological analysis. All specimens had two pairs of gonopores and gonoducts. The oviduct had a wider diameter in females, whereas in males the vasa deferentia were more developed. Gonads were composed of two parallel structures located in the cephalothorax. Female gonads were classified into three stages. The presence of transitional specimens may indicate that sex change occurs, with protandric hermaphroditism, as observed in other species of Parastacus . Because previous studies had demonstrated that P. varicosus is a gonochoristic species in a population from Uruguay, the present study addressed the possibility that the sexual pattern depends on differences in environmental factors among populations. To improve understanding of evolution of hermaphroditism in these crustaceans, their reproductive dynamics should be studied, including identification of the factors stimulating male and female functions.     

Reproduction and developmental pathways of Red Sea Xeniidae (Octocorallia,Alcyonacea)

This work on Red Sea alcyonaceans describes the reproductive patterns of 21 xeniid species. Gonochorism is the commonest sexual mode but simultaneous hermaphroditism was recorded in 4 species and brooding of planulae was observed in 15 species. The reproductive patterns of Xenia umbellata and Heteroxenia fuscescens were examined. X. umbellata exhibits seasonal spermatogenesis, continuous oogenesis and a 7 month period of planulation each year. H. fuscescens has continuous gametogenesis, and planulation occurs throughout the year, lacking any lunar pattern. The prolonged breeding season of H. fuscescens reflects intrapopulation asynchrony in larval development and subsequent maturation. Xeniids possess structurally similar gonads, but adopt diverse reproductive and developmental pathways.     

The effects of sex ratio and density on the expression of gender in the polychaeteCapitella capitata

Summary Large natural populations of the marine polychaeteCapitella capitata (species type I) contain males, females, and occasionally, hermaphrodites. Environmental conditions control the occurrence of hermaphrodites. At low density or in groups with female-biased sex ratios, males develop into hermaphrodites, and hermaphrodites are common. Crosses suggest that females are heterogametic, and males and hermaphrodites are homogametic. Heterogametic females do not become hermaphrodites.This study shows that in homogametic individuals, environmental conditions determine not only the development of hermaphroditism but also the expression of initial gender. Homogametic individuals can express either male or female gender initially, and homogametic individuals of either gender can develop subsequently into simultaneous hermaphrodites. The choice of initial gender depends on isolation. Most homogametic juveniles become females if reared alone but males if reared with other conspecifics. Homogametic males readily develop into hermaphrodites if females are rare. In contrast, homogametic females rarely become hermaphrodites.     

Morphological study for understanding the sexual pattern in ghost shrimp Callichirus major (Crustacea: Axiidea)

Callichirus major is a species of ghost shrimp that has burrowing habits, building underground galleries in the intertidal region of beaches. This study aimed to analyse some morphological parameters to verify its type of sexual pattern. A sample of 79 individuals, collected at the beach of Corujão, Espírito Santo, Brazil, was used. Data on the external morphology, presence of gonopores, carapace length (CL) and cheliped lengths (Lch) were obtained. In addition, the gonads were processed for histological analysis. The animals were divided into three categories: A, individuals with male characteristics; B, those presenting exclusively female characteristics; C, animals with characteristics of intersex, which presented supernumerary gonopores. Positive allometries were found for Lch versus CL only for category A, characterizing the group's marked heterochely. Histologically, it was possible to observe the presence of oocytes in category A and one case of male tissue in an intersex individual (category C). However, no genital ducts of both sexes were found in that individual. Callichirus major presents controversial sexual characters, which makes it difficult to understand the reproductive parameters and sexual pattern. However, maybe that species can develop a sexual system of hermaphroditism.