Growth rate during early life affects sexual differentiation in roach (<Emphasis Type="Italic">Rutilus rutilus</Emphasis>)

Many environmental factors have been shown to influence sex differentiation in fish, resulting in sex-biased populations, but the effects of growth rate have received limited attention. We conducted a controlled laboratory experiment in which growth rate and population density were manipulated in roach (Rutilus rutilus) during early development, and the subsequent effects on sex ratio determined. Significant differences in growth rate between fish populations were induced through provision of three different ration levels. In the slowest growing population there were fewer females compared within the fastest growing population (19% compared to 36% females), suggesting that in roach it may be more advantageous to become a small male than a small female when growth potential is limited. This may result from the fact that fecundity is limited by body size in female roach and that male roach are able to reproduce at a significantly smaller body size than females. In contrast, where roach were kept at different stocking densities, and there were no differences in growth rate, the subsequent proportion of females did not vary. Our data highlight the importance of controlling for growth rate in research on sexual differentiation in this species, notably when assessing for the effects of endocrine disrupting chemicals and other environmental factors, and have implications for fisheries management and aquaculture. The underlying mechanism for the influence of growth rate on sex differentiation has yet to be determined but is likely to have a strong endocrinological basis.     

Dynamics and histological characteristics of gonadal sex differentiation in pejerrey (Odontesthes bonariensis) at feminizing and masculinizing temperatures

This study evaluated the effects of different temperatures on the histological process of sex differentiation in the pejerrey Odontesthes bonariensis, a fish with marked temperature-dependent sex determination (TSD), at feminizing, neutral, and masculinizing temperatures. Fish reared at three temperatures (17 degrees C, 24 degrees C, and 29 degrees C) from hatching were sampled weekly until 11 weeks and their gonads were examined by histology. The percentages of females at 17 degrees C, 24 degrees C, and 29 degrees C were 100%, 73%, and 0%, respectively. Sex differentiation occurred earlier and at a smaller body size at higher temperatures in both sexes. The first signs of ovarian differentiation were observed at 4 and 7 weeks at 24 degrees C and 17 degrees C, respectively, and those of testicular differentiation at 4 and 7 weeks at 29 degrees C and 24 degrees C, respectively. Body or gonadal growth rates before sex differentiation were not proportional to temperature and showed no sexual dimorphism at 24 degrees C, where both sexes were present. Thus, differential growth rate is probably not a factor in TSD or histological sex differentiation in pejerrey. Blood vessels were formed before sex differentiation in both sexes and at all temperatures, and may be important for sex differentiation. No signs of intersexuality were found in any of the groups, and this characterizes pejerrey as the differentiated type of gonochorist even at feminizing and masculinizing temperatures. Ovaries were formed by the same histological processes at feminizing (17 degrees C) and neutral (24 degrees C) temperatures and without any pathological features such as germ cell degeneration. The process of testicular formation was generally similar at 24 degrees C and 29 degrees C, but some fish at 29 degrees C had widespread germ cell degeneration before sex differentiation. This suggests that pathological processes leading to germ cell death, such as heat-induced dysfunction of the supporting somatic cells, could be involved in masculinization of the genetic females at high temperatures.     

Sex determination in flatfishes: Mechanisms and environmental influences

Flounder of the genus Paralichthys exhibit a unique mode of sex determination where both low and high temperatures induce male-skewed sex ratios, while intermediate temperatures produce a 1:1 sex ratio. Male differentiation is thus easily induced in genetic females creating a combination of genetic (GSD) and environmental sex determination (ESD). Since male flounder become reproductively fit at substantially smaller body sizes than females, temperature or other environmental variables that elicit lower growth rates may also influence sex differentiation toward male development. This review covers our current knowledge of sex determination and differentiation in flatfishes including possible adaptive significance of ESD and involvement of factors such as aromatase (cyp19).     

Sex differences in growth rates of early life stage Japanese eels Anguilla japonica under experimental conditions

To assess the relationship between growth rate of body mass and sex in the Japanese eel Anguilla japonica in the early life stage; the growth rates of males and females were compared under experimental conditions. The mean growth rate of females was significantly slower than that of males. To assess the relative priority of growth rate and sex, growth was delayed by restricted feeding, resulting in a significantly higher proportion of females in the delayed than in the normal growth group. These findings indicate that the mean growth rate of A. japonica is slower in females than in males in the early life stage around sex determination and differentiation under experimental rearing conditions. Moreover, growth rate probably has priority over sex determination, with slow growth rate increasing the probability of being female.     

Growth and sex effects on the expression of syndecan-4 and glypican-1 in turkey myogenic satellite cell populations

The adult skeletal muscle stem cells, satellite cells, are responsible for skeletal muscle growth and regeneration. Satellite cells represent a heterogeneous cell population that differentially express cell surface markers. The membrane-associated heparan sulfate proteoglycans, syndecan-4, and glypican-1, are differentially expressed by satellite cells during the proliferation and differentiation stages of satellite cells. However, how the population of syndecan-4- or glypican-1-positive satellite cells changes during proliferation and differentiation, and how sex and muscle growth potential affect the expression of these genes is unknown. Differences in the amount of satellite cells positive for syndecan-4 or glypican-1 would affect the process of proliferation and differentiation which would impact both muscle mass accretion and the regeneration of muscle. In the current study, the percentage of satellite cells positive for syndecan-4 or glypican-1 from male and female turkeys from a Randombred Control Line 2 and a line (F) selected for increased 16-week body weight were measured during proliferation and differentiation. Growth selection altered the population of syndecan-4- and glypican-1-positive satellite cells and there were sex differences in the percentage of syndecan-4- and glypican-1-positive satellite cells. This study provides new information on dynamic changes in syndecan-4- and glypican-1-positive satellite cells showing that they are differentially expressed during myogenesis and growth selection and sex affects their expression.     

Sex-lethal in neurons controls female body growth in Drosophila

Sexual size dimorphism (SSD), a sex difference in body size, is widespread throughout the animal kingdom, raising the question of how sex influences existing growth regulatory pathways to bring about SSD. In insects, somatic sexual differentiation has long been considered to be controlled strictly cell-autonomously. Here, we discuss our surprising finding that in Drosophila larvae, the sex determination gene Sex-lethal (Sxl) functions in neurons to non-autonomously specify SSD. We found that Sxl is required in specific neuronal subsets to upregulate female body growth, including in the neurosecretory insulin producing cells, even though insulin-like peptides themselves appear not to be involved. SSD regulation by neuronal Sxl is also independent of its known splicing targets, transformer and msl-2, suggesting that it involves a new molecular mechanism. Interestingly, SSD control by neuronal Sxl is selective for larval, not imaginal tissue types, and operates in addition to cell-autonomous effects of Sxl and Tra, which are present in both larval and imaginal tissues. Overall, our findings add to a small but growing number of studies reporting non-autonomous, likely hormonal, control of sex differences in Drosophila, and suggest that the principles of sexual differentiation in insects and mammals may be more similar than previously thought.     

Zebrafish monosex population reveals female dominance in sex determination and earliest events of gonad differentiation

The zebrafish is a popular model for genetic analysis and its sex differentiation has been the focus of attention for breeding purposes. Despite numerous efforts, very little is known about the mechanism of zebrafish sex determination. The lack of discernible sex chromosomes and the difficulty of distinguishing the sex of juvenile fish are two major obstacles that hamper the progress in such studies. To alleviate these problems, we have developed a scheme involving methyltestosterone treatment followed by natural mating to generate fish with predictable sex trait. Female F1 fish that gave rise to all-female offspring were generated. This predictable sex trait enables characterization of gonadal development in juvenile fish by histological examination and gene expression analysis. We found the first sign of zebrafish sex differentiation to be ovarian gonocyte proliferation and differentiation at 10 to 12 days post-fertilization (dpf). Somatic genes were expressed indifferently at 10 to 17 dpf, and then became sexually dimorphic at three weeks. This result indicates clear distinction of male and female gonads derived independently from primordial gonads. We classified the earliest stages of zebrafish sex determination into the initial preparation followed by female germ cell growth, oocyte differentiation, and somatic differentiation. Our genetic selection scheme matches the prediction that female-dominant genetic factors are required to determine zebrafish sex.     

甲壳动物性别分化的遗传决定及外界因素的影响(英文)

本文综述了甲壳动物的性别决定机理及外界因素对性别分化的影响。绝大多数甲壳动物没有明显的性染色体 ,促雄腺被认为是甲壳动物性别分化的最主要的决定因子 ,其作用已得到了广泛的证明。由于甲壳动物幼体在早期发育过程中具有向两性发育的潜能 ,促雄腺可以决定个体未来发育的性别 ,并且通过人为摘除或移植促雄腺的方法可以使性别已经分化的个体发生性逆转 ,从而改变幼体的性别。虽然甲壳动物的性别是由遗传决定的 ,但外界的因素比如寄生、光周期、温度或激素可以改变其性比 ,其中以寄生的影响研究比较多 ,并认为是影响某些甲壳动物性别分化的主要外界因子。由于大多数养殖的甲壳动物雌雄性之间有体重和体长的差异 ,在水产养殖中可以利用这些特征进行全雌或全雄种苗的生产 ,以提高产量和效益。     

Sex dimorphism in digital formulae of children

This paper presents results of a study designed to: 1) test for a sex difference in the relative lengths of the finger bones, including the second-to-fourth digit ratio (2D:4D), using left-hand radiographs taken in young children, 2) test whether sex differences can be explained by sex differences in fetal growth, and 3) test the serial stability of sex differences in relative digit lengths, including 2D:4D. Results are presented from 1,060 subjects of the California Child Health and Development Studies. One serial replication at about 9 years old is available from 271 subjects. Results indicate that relative digit lengths are sex-dimorphic in children (Manning et al. [1998] Hum. Reprod. 13:3000-3004, [2004] Early Hum. Dev. 80:161-168). Sex differences in digit length ratios are more pronounced within sibships, where shared family factors are controlled, and are not strongly associated with gross measures of fetal growth, like birth length or weight. Thus, sex differences in the fetal growth of the body are not implicated in sex differences in digital formulae, leaving open the possibility of more direct hormonal and/or genetic causation. However, 2D:4D declined between ages 6-8 in a longitudinal sample, and was a less consistent sex-dimorphic marker than 3D:4D across ethnic groups, suggesting that 3D:4D may be a better marker of perinatal sex differentiation. Prior conflicting findings about 2D:4D may be partly explained by variations in age and ethnicity of populations studied.     

Ontogeny and the evolution of adult body size dimorphism in apes

This analysis investigates the ontogeny of body size dimorphism in apes. The processes that lead to adult body size dimorphism are illustrated and described. Potential covariation between ontogenetic processes and socioecological variables is evaluated. Mixed-longitudinal growth data from 395 captive individuals (representing Hylobates lar [gibbon], Hylobates syndactylus [siamang], Pongo pygmaeus [orangutan], Gorilla gorilla [gorilla], Pan paniscus [pygmy chimpanzee], and Pan troglodytes [“common” chimpanzee]) form the basis of this study. Results illustrate heterogeneity in the growth processes that produce ape dimorphism. Hylobatids show no sexual differentiation in body weight growth. Adult body size dimorphism in Pongo can be largely attributed to indeterminate male growth. Dimorphism in African apes is produced by two different ontogenetic processes. Both pygmy chimpanzees (Pan paniscus) and gorillas (Gorilla gorilla) become dimorphic primarily through bimaturism (sex differences in duration of growth). In contrast, sex differences in rate of growth account for the majority of dimorphism in common chimpanzees (Pan troglodytes). Diversity in the ontogenetic pathways that produce adult body size dimorphism may be related to multiple evolutionary causes of dimorphism. The lack of sex differences in hylobatid growth is consistent with a monogamous social organization. Adult dimorphism in Pongo can be attributed to sexual selection for indeterminate male growth. Interpretation of dimorphism in African apes is complicated because factors that influence female ontogeny have a substantial effect on the resultant adult dimorphism. Sexual selection for prolonged male growth in gorillas may also increase bimaturism relative to common chimpanzees. Variation in female growth is hypothesized to covary with foraging adaptations and with differences in female competition that result from these foraging adaptations. Variation in male growth probably corresponds to variation in level of sexual selection. © 1995 Wiley-Liss, Inc.     

Sexual differentiation in Salaria (=Blennius) pavo

The sexual differentiation of Salaria (= Blennius ) pavo is described from the stage of hatching to a body length of 35 mm. At hatching, the primordial germ cells (PGCs) can be recognized clearly. At a standard body length of 5 mm, they begin to protrude into the peritoneal cavity and at 14 mm they transform to oogonia. At 17 mm length, the first oocytes can be observed. In males at a standard length of 16–17 mm, the first signs of a differentiation into a testis can be recognized. Shortly after the differentiation of the male sex, the division of the male gonad into a testis and a testicular gland can be seen. The fine structural characteristics of the PGCs and of differentiation stages are presented.     

FGF and PI3 kinase signaling pathways antagonistically modulate sex muscle differentiation in C. elegans

Myogenesis in vertebrate myocytes is promoted by activation of the phosphatidyl-inositol 3'-kinase (PI3 kinase) pathway and inhibited by fibroblast growth factor (FGF) signaling. We show that hyperactivation of the Caenorhabditis elegans FGF receptor, EGL-15, similarly inhibits the differentiation of the hermaphrodite sex muscles. Activation of the PI3 kinase signaling pathway can partially suppress this differentiation defect, mimicking the antagonistic relationship between these two pathways known to influence vertebrate myogenesis. When ectopically expressed in body wall muscle precursor cells, hyperactivated EGL-15 can also interfere with the proper development of the body wall musculature. Hyperactivation of EGL-15 has also revealed additional effects on a number of fundamental processes within the postembryonic muscle lineage, such as cell division polarity. These studies provide important in vivo insights into the contribution of FGF signaling events to myogenesis.     

Body mass and sex‐biased parasitism in wood mice Apodemus sylvaticus

Male sex‐biased parasitism (SBP) occurs across a range of mammalian taxa and two contrasting sets of hypotheses have been suggested for its establishment. The first invokes body size per se and suggests that larger individuals are either a larger target for parasites, trade off growth at the expense of immunity or cope better with parasitism than smaller individuals. The second suggests a sex‐specific handicap whereby males have reduced immunocompetence compared to females due to the immunodepressive effects of testosterone. The current study investigated whether sex‐biased parasitism is driven by host ‘body size’ or ‘sex’ using a rodent–tick (Apodemus sylvaticus–Ixodes ricinus) system. Moreover, the presence or absence of large mammals at study sites were used to control the presence of immature ticks infesting wood mice, allowing the impacts of parasitism on host body mass and female reproduction to be assessed. As expected, male mice had greater tick loads than females and analyses suggested this sex‐bias was driven by body mass as opposed to sex. It is therefore likely that larger individuals are a larger target for parasites, trade off growth at the expense of immunity or adapt behavioural responses to parasitism based on their body size. Parasite load had no effect on host body mass or female reproductive output suggesting individuals may alter behaviour or life history strategies to compensate for costs incurred through parasitism. Overall, this study lends support to the ‘body size’ hypothesis for the formation of sex‐biased parasitism.     

Growth and reproductive development in the male tree shrew (Tupaia belangeri) from birth to sexual maturity

The growth and reproductive development of the male tree shrew were studied from birth to sexual maturity. An infantile phase from birth to Day 30 was characterized by the rapid involution of the testis and adrenal gland from a fetal condition followed by a nadir in testosterone levels and slow growth and differentiation of the testis and accessory sex organs. The initiation of puberty occurred collaterally with the emergence of the young from the nest and was marked by a sharp rise in testosterone levels from Days 30 to 35 to maximum levels at Days 40-55. Peak testosterone levels were temporally correlated with the onset of maximum growth and differentiation of the testis and accessory sex organs, descent of the testis, development of the scrotum, and a pronounced peak in the weight-velocity curve. The rapid growth of males at puberty contributed to a moderate degree of sexual dimorphism in this species. Puberty was attained at about Day 90 with the completion of spermatogenesis and the functional differentiation of the accessory sex organs. The postnatal development of the tree shrew conforms with the general primate pattern. The precise endocrine correlates established during puberty make Tupaia belangeri a useful small animal model for the study of puberty in primates.     

Sex‐specific growth in chicks of the sexually dimorphic Black‐tailed Godwit

Sexual size dimorphism (SSD) is common in birds and has been linked to various selective forces. Nevertheless, the question of how and when the sexes start to differentiate from each other is poorly studied. This is a critical knowledge gap, as sex differences in growth may cause different responses to similar ecological conditions. In this study, we describe the sex‐specific growth – based on body mass and five morphometric measurements – of 56 captive Black‐tailed Godwit Limosa limosa limosa chicks raised under ad libitum food conditions, and conclude that all six growth curves are sex‐specific. Females are the larger sex in terms of body mass and skeletal body size. To test whether sex‐specific growth leads to sex‐specific susceptibility to environmental conditions, we compared the age‐specific sizes of male and female chicks in the wild with those of Black‐tailed Godwits reared in captivity. We then tested for a relationship between residual growth and relative hatching date, age, sex and habitat type in which the wild chicks were born. Early‐hatched chicks were relatively bigger and in better condition than late‐hatched chicks, but body condition and size were not affected by natal habitat type. Female chicks deviated more negatively from the sex‐specific growth curves than male chicks for body mass and total‐head length. This suggests that the growth of the larger females is more susceptible to limiting environmental conditions. On average, the deviations of wild chicks from the predicted growth curves were negative for all measurements, which suggests that conditions are limiting in the current agricultural landscape. We argue that in estimating growth curves for sexually dimorphic species, it is critical first to make accurate sex and age determinations.     

Social environment and sex differentiation in the false clown anemonefish, Amphiprion ocellaris

Plasticity in sex differentiation is known to be common in teleost fishes. Anemonefishes are protandrous; females are the largest and dominant members of social groups, displaying frequent aggressive behavior towards other members of groups. The second-ranked individuals become males and others remain as non-reproductive individuals. Here we examine the influence of social interaction in-group on sex differentiation in the false clown anemonefish, Amphiprion ocellaris, under laboratory conditions. Three juvenile anemonefish were kept in a tank for 180 days and their behaviors observed once a month. The social rank of individuals was distinguishable by their interactions in a group, with rank order clearly correlated with aggressive and appeasing behaviors. The dominant individuals occupied the shelter in the tank from the start to completion of the observation period. The body mass of dominant individuals increased compared to group-housed control fish, while third-ranked individuals showed growth suppression. The ratio of testicular tissue in gonads increased in dominant and second-ranked individuals but decreased in the third-ranked individuals. Differences in the plasma concentrations of estradiol, testosterone, and cortisol were not significant, but the concentration of 11-ketotestosterone was significantly higher in dominant individuals. These results suggest that, in false clown anemonefish, reproductive suppression of lower-ranked individuals becomes apparent in the first stage of group formation, and sex differentiation of upper-ranked individuals is gradually determined by long-term social interactions.     

Histological study of the development and sex differentiation of the gonad in the European eel

The histological structure of the gonads was studied in yellow eels sampled from a coastal lagoon and from stocks reared in an aquaculture plant showing different sex ratios. Gonad development related to body size rather than to age and underwent an intermediate stage characterized by a structure of an early testis but containing oogonia and oocytes. This gonad was called the Syrski organ and the stage juvenile ambisexual. Ovaries were found in eels from 22–30 cm in length, possibly derived from undifferentiated gonads or from Syrski organs. Fully differentiated testes were found in eels >35 cm, derived from Syrski organs. These observations support the results of previous research. From elvers and in eels up to 15–16 cm in length, growth of the gonadal primordium is due to primordial germ cell migration. In eels > 15 cm multiplication of primordial cells begins. Oogonial clones were found in eels > 18 cm in length, whilespermatogonium B clones were observed in eels >30 cm in length. The dynamics of sex differentiation was different among stocks with different ultimate sex ratios: ovaries were found in shorter eels in stocks with a prevalence of females, in longer eels in stocks with a prevalence of males. This result supports the hypothesis of a metagametic (environmental) sex determination. The somatic cells in contact with germ cells and those in the interstitium appeared early during gonad development and preceded germ cell differentiation. This suggests that somatic cells are the targets of the environmental factors influencing sex differentiation.