Temperature-dependent sex determination of Ascension Island green turtles

We examined the role played by temperature in the duration of incubation and sex ratio of green turtle hatchlings at Ascension Island, one of the most important green turtle rookeries in the Atlantic. Temperature at control sites at nest depth and in 39 green turtle nests was measured using small temperature recording devices. The sex ratio of hatchlings was ascertained in a sub-sample of monitored nests allowing the description of the relationship between intranest temperature and hatchling sex ratio, demonstrating a pivotal incubation temperature of 28.8°C. The seasonal profile in sex ratio of hatchlings produced on all nesting beaches at Ascension Island was estimated, showing that a female-biased sex ratio would be expected with a female:male ratio of the order of 3:1. The use of nest temperature, air temperature, sand temperature at control sites, and incubation duration as proxies to estimate hatchling sex ratio are discussed.

Breeding periodicity for male sea turtles, operational sex ratios, and implications in the face of climate change

Species that have temperature-dependent sex determination (TSD) often produce highly skewed offspring sex ratios contrary to long-standing theoretical predictions. This ecological enigma has provoked concern that climate change may induce the production of single-sex generations and hence lead to population extirpation. All species of sea turtles exhibit TSD, many are already endangered, and most already produce sex ratios skewed to the sex produced at warmer temperatures (females). We tracked male loggerhead turtles (Caretta caretta) from Zakynthos, Greece, throughout the entire interval between successive breeding seasons and identified individuals on their breeding grounds, using photoidentification, to determine breeding periodicity and operational sex ratios. Males returned to breed at least twice as frequently as females. We estimated that the hatchling sex ratio of 70:30 female to male for this rookery will translate into an overall operational sex ratio (OSR) (i.e., ratio of total number of males vs females breeding each year) of close to 50:50 female to male. We followed three male turtles for between 10 and 12 months during which time they all traveled back to the breeding grounds. Flipper tagging revealed the proportion of females returning to nest after intervals of 1, 2, 3, and 4 years were 0.21, 0.38, 0.29, and 0.12, respectively (mean interval 2.3 years). A further nine male turtles were tracked for short periods to determine their departure date from the breeding grounds. These departure dates were combined with a photoidentification data set of 165 individuals identified on in-water transect surveys at the start of the breeding season to develop a statistical model of the population dynamics. This model produced a maximum likelihood estimate that males visit the breeding site 2.6 times more often than females (95%CI 2.1, 3.1), which was consistent with the data from satellite tracking and flipper tagging. Increased frequency of male breeding will help ameliorate female-biased hatchling sex ratios. Combined with the ability of males to fertilize the eggs of many females and for females to store sperm to fertilize many clutches, our results imply that effects of climate change on the viability of sea turtle populations are likely to be less acute than previously suspected.

Retracted: Experimental evidence that maternal corticosterone controls adaptive offspring sex ratios

1. Sex allocation theory has received considerable attention, yet the mechanism(s) by which mothers skew offspring sex ratios remain unknown. In birds, females are the heterogametic sex, which potentially gives them control of whether gametes will be male or female. How females might control the sex of the gamete is unclear, but one possibility is that variation in steroid hormones may mediate this process. 2. We experimentally altered circulating levels of corticosterone in female Gouldian finches (Erythrura gouldiae), a species that demonstrates both extreme stress responses and extreme offspring sex ratio biases when breeding with a low-quality (genetically incompatible) partner. 3. During egg production, individual females received both corticosterone and metyrapone (a corticosterone-synthesis inhibitor) implants, in random order, to induce both high and low levels of circulating stress hormones (within physiological limits). 4. We found that females with elevated corticosterone levels produced male-biased sex ratios, but when the same females were treated with metyrapone they produced female-biased offspring sex ratios. 5. These stress responses are adaptive because females constrained to breeding with low-quality males can substantially increase their fitness by overproducing sons. Changes in maternal corticosterone levels during stressful situations, such as the quality of a breeding partner, may provide an endocrine mechanism that can be exploited for strategic sex allocation.

Sex as a determinant of relapse incidence and progressive course of multiple sclerosis

The aim of this work was to evaluate sex differences in the incidence of multiple sclerosis relapses; assess the relationship between sex and primary progressive disease course; and compare effects of age and disease duration on relapse incidence. Annualized relapse rates were calculated using the MSBase registry. Patients with incomplete data or <1 year of follow-up were excluded. Patients with primary progressive multiple sclerosis were only included in the sex ratio analysis. Relapse incidences over 40 years of multiple sclerosis or 70 years of age were compared between females and males with Andersen-Gill and Tweedie models. Female-to-male ratios stratified by annual relapse count were evaluated across disease duration and patient age and compared between relapse-onset and primary progressive multiple sclerosis. The study cohort consisted of 11 570 eligible patients with relapse-onset and 881 patients with primary progressive multiple sclerosis. Among the relapse-onset patients (82 552 patient-years), 48 362 relapses were recorded. Relapse frequency was 17.7% higher in females compared with males. Within the initial 5 years, the female-to-male ratio increased from 2.3:1 to 3.3:1 in patients with 0 versus ≥4 relapses per year, respectively. The magnitude of this sex effect increased at longer disease duration and older age (P < 10−12). However, the female-to-male ratio in patients with relapse-onset multiple sclerosis and zero relapses in any given year was double that of the patients with primary progressive multiple sclerosis. Patient age was a more important determinant of decline in relapse incidence than disease duration (P < 10−12). Females are predisposed to higher relapse activity than males. However, this difference does not explain the markedly lower female-to-male sex ratio in primary progressive multiple sclerosis. Decline in relapse activity over time is more closely related to patient age than disease duration.

Reproductive and behavioral aspects of red-winged tinamous (Rhynchotus rufescens) in groups with different sex ratios

The aim of this research study was to evaluate the reproductive performance of tinamous submitted to five different male:female ratios. The study was carried out with 72 birds in a randomized experimental design with 4 replications. Tinamous were housed in cages, using the ratios of one (1:1), two (2:1), three (3:1) and four (4:1) females per male, and also one male was housed with three females individually (3R:1), in a rotational system. Reproductive records of the breeding season from September 2004 to March 2005 were used. The reproductive traits studied were: number of eggs laid, fertility, and percentage of eggs damaged and cracked by pecking. Nonparametric analyses of these traits were performed using Kruskal-Wallis test. Two replications of treatments 1:1 and 4:1, and one of treatment 2:1 were video-taped for three days, 12 hours/day. The videotapes were sampled according to the scan method to fit an ethogram. Birds were also watched for one hour per day to study dominance and agonistic behavior. None of the reproductive traits was affected by mating sex ratio (p<0.05). Female dominance could be related to displacement behavior (r=1.00), and male sitting in immobility plus sitting in activity behaviors were related to lower number of damaged eggs (r=-0.90). Social dominance was indirectly determined by displacement behavior in the study situation. A large number of damaged eggs occurred in all treatments, thereby not allowing a clear conclusion on the best male:female ratio.

Size and sex compositions, length-weight relationship, and occurrence of the Brazilian sharpnose shark, Rhizoprionodon lalandii, caught by artisanal fishery from southeastern Brazil

Although the Brazilian sharpnose shark, Rhizoprionodon lalandii (Muller and Henle, 1839), is an inshore species widely distributed in the Western Atlantic from Panama to Uruguay, there is little available information on its biology. During a long-term study of small coastal sharks caught by gill net fisheries in southeastern Brazil (PROJETO CACAO), 3643 specimens of R. lalandii were examined, comprising 61.3% of the total sharks,and including all sizes classes, from 30 to 78,5 cm TL., and weights from 100 to 2950 g. The length-weight relationships were not significantly different between sexes, Overall sex ratio favoured the males slightly at the rate of 1.3: 1. Sex ratios, however, did differ significantly between season and size classes. This species occurred in this area all year long. Three seasonal size-class Occurrence patterns were recognized: (1) between October and March, the juveniles were more frequents (2) from April to July, adults were most common, and (3) from August to September, neonates were most numerically abundant. Such patterns we to associated with reproductive tactics that may reduce intra-specific and inter-specific competition with hammerhead shark neonates (Sphyrna lewini). probably result in reduced natural mortality of the offspring during their first few months. (C) 2005 Elsevier B.V. All rights reserved.

Sex allocation in the polydomous leaf-cutting ant Acromyrmex balzani

Logistic regression analysis was used to analyse sex allocation in a population of the leaf-cutting ant Acromyrmex balzani occurring in a pasture in southern Brazil. The field sample consisted of 151 fungus-garden chambers (18 queenright and 133 queenless), belonging to 50 nests with three vertically stacked chambers per nest on average. Taking nest chamber as the unit of analysis, seven predictor variables were considered: sampling date, chamber depth, chamber volume, weight of fungus garden, presence of a queen, number of large workers, and number of small to medium workers. The population-level numerical proportion of females was 0.548 and the inferred proportional energetic investment in females 0.672. The former was not significantly different from 0.5 (P=0.168), but the latter was (P=0.0003). The proportional investment in females per fungus garden increased with the number of large workers present (P=0.0002) and decreased with the dry weight of the fungus garden (P=0.012). This implies that resource acquisition through foraging is likely to be a major proximate determinant of sex allocation. The negative correlation between female bias and fungus garden weight might be due to developing adult females requiring more food than males, but this hypothesis could not be confirmed by direct statistical evidence.

Chromosome mapping of repetitive sequences in Anostomidae species: Implications for genomic and sex chromosome evolution

Background: Members of the Anostomidae family provide an interesting model system for the study of the influence of repetitive elements on genome composition, mainly because they possess numerous heterochromatic segments and a peculiar system of female heterogamety that is restricted to a few species of the Leporinus genus. The aim of this study was to isolate and identify important new repetitive DNA elements in Anostomidae through restriction enzyme digestion, followed by cloning, characterisation and chromosome mapping of this fragment. To identify repetitive elements in other Leporinus species and expand on studies of repetitive elements in Anostomidae, hybridisation experiments were also performed using previously described probes of LeSpeI repetitive elements. Results: The 628-base pair (bp) LeSpeII fragment was hybridised to metaphase cells of L. elongatus individuals as well as those of L. macrocephalus, L. obtusidens, L. striatus, L. lacustris, L. friderici, Schizodon borellii and S. isognathus. In L. elongatus, both male and female cells contained small clusters of LeSpeII repetitive elements dispersed on all of the chromosomes, with enrichment near most of the terminal portions of the chromosomes. In the female sex chromosomes of L. elongatus (Z2,Z2/W1W 2), however, this repeated element was absent. In the remaining species, a dispersed pattern of hybridisation was observed on all chromosomes irrespective of whether or not they were sex chromosomes. The repetitive element LeSpeI produced positive hybridisations signals only in L. elongatus, L. macrocephalus and L. obtusidens, i.e., species with differentiated sex chromosomes. In the remaining species, the LeSpeI element did not produce hybridisation signals. Conclusions: Results are discussed in terms of the effects of repetitive sequences on the differentiation of the Anostomidae genome, especially with respect to sex chromosome evolution. LeSpeII showed hybridisation patterns typical of Long Interspersed Elements (LINEs). The differential distribution of this element may be linked to sex chromosome differentiation in L. elongatus species. The relationship between sex chromosome specificity and the LeSpeI element is confirmed in the species L. elongatus, L. macrocephalus and L. obtusidens. © 2012 da Silva et al.; licensee BioMed Central Ltd.

No evidence of predation causing female-biased sex ratios in marine pelagic copepods

Although sex ratios close to unity are expected in dioecious species, biased sex ratios are common in nature. It is essential to understand causes of skewed sex ratios in situ, as they can lead to mate limitation and have implications for the success of natural populations. Female-skewed sex ratios are commonly observed in copepods in situ. Here we discuss the challenges of copepod sex ratio research and provide a critical review of factors determining copepod sex ratios, focusing on 2 main objectives. The first is a critique of the male predation theory, which is currently the main process thought to be responsible for female-skewed sex ratios. It assumes that males have higher mortality because of increased vulnerability to predation during their search for mates. We show that there is little support for the male predation theory, that sex ratios skewed toward females occur in the absence of predation, that sex ratios are not related to predation pressure, and that where sex-skewed predation does occur, it is biased toward females. Our second objective is to suggest alternative hypotheses regarding the determination of sex ratios. We demonstrate that environmental factors, environmental sex determination and sex change have strong effects on copepod sex ratios, and suggest that differential physiological longevity of males and females may be more important in determining sex ratios than previously thought. We suggest that copepod sex ratios are the result of a mixture of factors.

Female-biased sex ratios in marine pelagic copepods: Response to Hirst et al. (2013)

Hirst et al. (2013; Mar Ecol Prog Ser 489:297-298) suggest that Gusmão et al. (2013; Mar Ecol Prog Ser 482:279-298) misinterpreted the findings of Hirst et al. (2010; Limnol Oceanogr 55:2193-2206). They restate that the major factors determining sex ratio in pelagic copepods act upon the adult stage, but they place less emphasis on the idea that predation on male copepods is a likely determinant, and highlight the role of physiological longevity. Here we reconsider the data and confirm our position that at present there is limited evidence to support the theory of male-skewed predation. However, we agree that sex determination is governed by a combination of factors, with the relative emphasis being the main point of contention between the 2 parties.

Sex determination and temperature-induced sex differentiation in three natural populations of Nile tilapia (Oreochromis niloticus) adapted to extreme temperature conditions

As a species of major interest for aquaculture, the sex determination system (SDS) of Nile tilapia, Oreochromis niloticus, has been widely investigated. In this species, sex determination is considered to be governed by the interactions between a complex system of genetic sex determination factors (GSD) and the influence of temperature (TSD) during a critical period. Previous studies were exclusively carried out on domestic stocks with the genetic and maintenance limitations associated. Given the wide distribution and adaptation potential of the Nile tilapia, we investigated under controlled conditions the sex determination system of natural populations adapted to three extreme thermal regimes: stable extreme environments in Ethiopia, either cold temperatures in a highland lake (Lake Koka), or warm temperatures in hydrothermal springs (Lake Metahara), and an environment with large seasonal variations in Ghana (Kpandu, Lake Volta). The sex ratio analysis was conducted on progenies reared under constant basal (27 degrees C) or high (36 degrees C) temperatures during the 30 days following yolk-sac resorption. Sex ratios of the progenies reared at standard temperature suggest that the three populations share a similar complex GSD system based on a predominant male heterogametic factor with additional influences of polymorphism at this locus and/or action of minor factors. The three populations presented a clear thermosensitivity of sex differentiation, with large variations in the intensity of response depending on the parents. This confirms the presence of genotype-environment interactions in TSD of Nile tilapia. Furthermore the existence of naturally sex-reversed individuals is strongly suggested in two populations (Kpandu and Koka). However, it was not possible here to infer if the sex-inversion resulted from minor genetic factors and/or environmental influences. The present study demonstrated for the first time the conservation of a complex SDS combining polymorphic GSD and TSD components in natural populations of Nile tilapia. We discuss the evolutionary implications of our findings and highlight the importance of field investigations of sex determination. (c) 2007 Elsevier B.V. All rights reserved.

Invasion and fixation of sex-reversal genes

We simulated a meta-population with random dispersal among demes but local mating within demes to investigate conditions under which a dominant female-determining gene W, with no individual selection advantage, can invade and become fixed in females, changing the population from male to female heterogamety. Starting with one mutant W in a single deme, the interaction of sex ratio selection and random genetic drift causes W to be fixed among females more often than a comparable neutral mutation with no influence on sex determination, even when YY males have slightly reduced viability. Meta-population structure and interdeme selection can also favour the fixation of W. The reverse transition from female to male heterogamety can also occur with higher probability than for a comparable neutral mutation. These results help to explain the involvement of sex-determining genes in the evolution of sex chromosomes and in sexual selection and speciation.

Sex identification and PIT-tagging: tools and prospects for studying intersexual differences in freshwater fishes

This study evaluated a technique to allow the long-term monitoring of individual fishes of known sex in the wild using sex confirmation in close proximity to the reproductive period combined with individual tagging. Hundreds of partially migratory roach Rutilus rutilus were tagged with passive integrated transponders (PIT) following sex determination in spring and various performance measures were compared with fish tagged outside the reproductive period in autumn. Short-term survival was > 95% for R. rutilus sexed and tagged under natural field conditions. Total length (LT) did not affect the probability of survival within the size range tagged (119–280mm), nor were there differences in timing of migration the following season between individuals sexed and tagged in spring and individuals tagged in autumn (i.e. outside the reproductive period). Also, a similar per cent of R. rutilus sexed and tagged in spring and tagged in autumn migrated the following season (34·5 and 34·7%). Moreover, long-term recapture data revealed no significant differences in body condition between R. rutilus individuals sexed and tagged in spring, individuals tagged in autumn and unmanipulated individuals. The observed sex ratio of recaptured fish did not differ from the expected values of equal recapture rates between males and females. Hence, there is no observable evidence for an adverse effect of tagging close to the reproductive period and therefore this method is suitable for studying intersexual differences and other phenotypic traits temporarily expressed during reproduction at the individual level in fishes.