The Emerging Phenotype: Ecological Genetics of Color, Form and Sex in the Common Frog

One of the main aims of evolutionary biology is to explain why organisms vary phenotypically as they do. Proximately, this variation arises from genetic differences and from environmental influences, the latter of which is referred to as phenotypic plasticity. Phenotypic plasticity is thus a central concept in evolutionary biology, and understanding its relative importance in causing the phenotypic variation and differentiation is important, for instance in anticipating the consequences of human induced environmental changes. The aim of this thesis was to study geographic variation and local adaptation, as well as sex ratios and environmental sex reversal, in the common frog (Rana temporaria). These themes cover three different aspects of phenotypic plasticity, which emerges as the central concept for the thesis. The first two chapters address geographic variation and local adaptation in two potentially thermally adaptive traits, namely the degree of melanism and the relative leg length. The results show that although there is an increasing latitudinal trend in the degree of melanism in wild populations across Scandinavian Peninsula, this cline has no direct genetic basis and is thus environmentally induced. The second chapter demonstrates that although there is no linear, latitudinally ordered phenotypic trend in relative leg length that would be expected under Allen s rule an ecogeographical rule linking extremity length to climatic conditions there seems to be such a trend at the genetic level, hidden under environmental effects. The first two chapters thus view phenotypic plasticity through its ecological role and evolution, and demonstrate that it can both give rise to phenotypic variation and hide evolutionary patterns in studies that focus solely on phenotypes. The last three chapters relate to phenotypic plasticity through its ecological and evolutionary role in sex determination, and consequent effects on population sex ratio, genetic recombination and the evolution of sex chromosomes. The results show that while sex ratios are strongly female biased and there is evidence of environmental sex reversals, these reversals are unlikely to have caused the sex ratio skew, at least directly. The results demonstrate that environmental sex reversal can have an effect on the evolution of sex chromosomes, as the recombination patterns between them seem to be controlled by phenotypic, rather than genetic, sex. This potentially allows Y chromosomes to recombine, lending support for the recent hypothesis suggesting that sex-reversal may play an important role on the rejuvenation of Y chromosomes.

Consanguinity, twinning and secondary sex ratio in the population of Karnataka, South India

Consanguineous marriages are strongly favoured in the state of Karnataka. Of 65492 marriages studied 33·07% were consanguineous, equivalent to a coefficient of inbreeding (F) of 0·0298. The twinning rate was low, 6·9 per thousand, whereas the secondary sex ratio, 0·5221, was higher than in comparable major human populations. Consanguinity exerted no significant effect on either parameter. The results also indicate that consanguinity is not associated with excess antenatal losses and suggest the possibility of enhanced selection against mutations at X chromosome loci.

Proposed role of w chromosome inactivation and the absence of dosage compensation in avian sex determination

Three features of avian sex chromosomes - female heterogamety (ZZ male, ZW female), the apparently inactive state of the W chromosome, and dose-dependent expression of Z-linked genes - are examined in regard to their possible relation to sex determination. It is proposed that the W chromosome is facultatively heterochromatic and that the Z and W chromosomes carry one or more homologous sex-determination genes. The absence of dosage compensation in ZZ embryos, and W inactivation in ZW embryos, would then bring about a 2n(ZZ)-n(ZW) inequality in the effective copy number of such genes. The absence of dosage compensation of Z-linked genes in ZZ embryos is viewed as a means by which two copies of Z-W homologous sex determination genes are kept active to meet the requirements of testis determination. W inactivation may promote ovarian development by reducing the effective copy number of these genes from 2n to n. If there is a W-specific gene for femaleness, spread of heterochromatization to this gene in cells forming the right gonadal primordium may explain the latter's normally undifferentiated state; reversal of heterochromatization may similarly explain the development of the right gonad into a testis following left ovariectomy.

The X Chromosome of Hemipteran Insects: Conservation, Dosage Compensation and Sex-Biased Expression

Insects of the order Hemiptera (true bugs) use a wide range of mechanisms of sex determination, including genetic sex determination, paternal genome elimination, and haplodiploidy. Genetic sex determination, the prevalent mode, is generally controlled by a pair of XY sex chromosomes or by an XX/XO system, but different configurations that include additional sex chromosomes are also present. Although this diversity of sex determining systems has been extensively studied at the cytogenetic level, only the X chromosome of the model pea aphid Acyrthosiphon pisum has been analyzed at the genomic level, and little is known about X chromosome biology in the rest of the order. In this study, we take advantage of published DNA- and RNA-seq data from three additional Hemiptera species to perform a comparative analysis of the gene content and expression of the X chromosome throughout this clade. We find that, despite showing evidence of dosage compensation, the X chromosomes of these species show female-biased expression, and a deficit of male-biased genes, in direct contrast to the pea aphid X. We further detect an excess of shared gene content between these very distant species, suggesting that despite the diversity of sex determining systems, the same chromosomal element is used as the X throughout a large portion of the order.

Effects of temperature on the biology of the northern shrimp, Pandalus borealis, in the Gulf of Maine

Length-frequency data collected from inshore and offshore locations in the Gulf of Maine in 1966-1968 indicated that ovigerous female northern shrimp (Pandalus borealis) first appeared offshore in August and September and migrated inshore in the fall and winter. Once eggs hatched, surviving females returned offshore. Juveniles and males migrated offshore during their first two years of life. Sex transition occurred in both inshore and oll'shore waters, but most males changed sex offshore during their third and fourth years. Most shrimp changed sex and matured as females for the first time in their fourth year. Smaller females and females exposed to colder bottom temperatures spawned first. The incidence of egg parasitism peaked in January and was higher for shrimp exposed to warmer bottom temperatures. Accelerated growth at higher temperatures appeared to result in earlier or more rapid sex transition. Males and non-ovigerous females were observed to make diurnal vertical migrations, but were not found in near- surface waters where the temperature exceeded 6°C. Ovigerous females fed more heavily on benthic molluscs in inshore waters in the winter, presumably because the egg masses they were carrying prevented them from migrating vertically at night. Northern shrimp were more abundant in the southwestern region of the Gulf of Maine where bottom temperatures remain low throughout the year. Bottom trawl catch rates were highest in Jeffreys Basin where bottom temperatures were lower than at any other sampling location. Catch rates throughout the study area were inversely related to bottom temperature and reached a maximum at 3°C. An increase of 40% in fecundity between 1973 and 1979 was associated with a decline of 2-3°C in April-July offshore bottom temperatures. Furthermore, a decrease in mean fecundity per 25 mm female between 1965 and 1970 was linearly related to reduced landings between 1969 and 1974. It is hypothesized that temperature-induced changes in fecundity and, possibly, in the extent of egg mortality due to parasitism, may provide a mechanism which could partially account for changes in the size of the Gulf of Maine northern shrimp population during the last thirty years. (PDF file contains 28 pages.)

Sex-ratio and fecundity of catfish, Clarias gariepinus in Opa reservoir, Obafemi Awolowo University, Ile-Ife

The sex-ratio of Clarias gariepinus in Opa Reservoir was 2:1 (male/female). The fecundity of C. gariepinus in Opa reservoir ranged between 1,567 and 650,625 egg. The fish species had extended spawning period which probably spreads the risk of predation on the eggs. The population of the fish species could be improved by stocking with the female breeders

Biologie de Penaeus duorarum notialis en Côte d'Ivoire. 4 - Rélations entre la répartition et les conditions du milieu. Étude des variations du sex-ratio

The analysis of the geographic and bathymetric distribution of Penaeus duorarum and, particularly P. d. notialis off Côte d'Ivoire and in its whole distribution area leads to the definition of the adult ecological requirements (temperature, salinity, grain size and sediment composition, organic matter) and the importance of the thermocline in the bathmetric distribution. The population structure study shows: (1) variations of size with depth, (2) variations of sex ratio, with size, depth and seasons.

Length and sex effects on the spatial structure of catches of Pacific halibut (Hippoglossus stenolepis) on longline gear

Field experiments were conducted to test the hypotheses that Pacific halibut (Hippoglossus stenolepis) display small-scale spatial structure within longline catches, relative to other species and empty hooks, or within-species based on sex or length. Sequential hook-by-hook inventories, along with length and sex data, were taken at thirty-one survey stations. Two-dimensional spatial statistics were used to test for 1) aggregation, defined as the clustering of individuals within a given demographic of size or sex over small intervals of distance; and 2) segregation, defined as the sequential occurrence of individuals within a given demographic of size or sex, uninterrupted by other observations, irrespective of the distance between individuals. Statistically significant structure was detected within catches that is more commonly associated with fish length than sex. Significant spatial structuring occurred at 60% of all stations tested. Significant aggregation of halibut of legal length for commercial retention (≥82 cm) was detected at 44% of stations and aggregation of sublegal-size halibut was detected at 11%. Maleand female-based aggregations were observed at 22% and 11% of stations, respectively. Significant segregation of females was observed at 20% of stations, male segregation occurred at 8% of stations, and segregation by size at 16% of stations. Understanding small-scale spatial structure within longline catches may help us interpret changes in survey and commercial catch data. If structure is generated by behavior, then observed size-at-age or relative sex-ratios may be biased relative to underlying distributions. Although physical processes such as gape limitation should remain stable over the time, dynamic processes may be spatially and temporally variabl

Estimates of body sizes at maturation and at sex change, and the spawning seasonality and sex ratio of the endemic Hawaiian grouper (Hyporthodus quernus, F. Epinephelidae)

A case study of the reproductive biology of the endemic Hawaiian grouper or hapu’upu’u (Hyporthodus quernus) is presented as a model for comprehensive future studies of economically important epinephelid groupers. Specimens were collected throughout multiple years (1978–81, 1992–93, and 2005–08) from most reefs and banks of the Northwestern Hawaiian Islands. The absence of small males, presence of atretic oocytes and brown bodies in testes of mature males, and both developed ovarian and testicular tissues in the gonads of five transitional fish provided evidence of protogynous hermaphroditism. No small mature males were collected, indicating that Hawaiian grouper are monandrous (all males are sex-changed females). Complementary microscopic criteria also were used to assign reproductive stage and estimate median body sizes (L50) at female sexual maturity and at adult sex change from female to male. The L50 at maturation and at sex change was 580 ±8 (95% confidence interval [CI]) mm total length (TL) and 895 ±20 mm TL, respectively. The adult sex ratio was strongly female biased (6:1). Spawning seasonality was described by using gonadosomatic indices. Females began ripening in the fall and remained ripe through April. A February–June main spawning period that followed peak ripening was deduced from the proportion of females whose ovaries contained hydrated oocytes, postovulatory follicles, or both. Testes weights were not affected by season; average testes weight was only about 0.2% of body weight—an order of magnitude smaller than that for ovaries that peaked at 1–3% of body weight. The species’ reproductive life history is discussed in relation to its management.

Accuracy of sex determination for northeastern Pacific Ocean thornyheads (Sebastolobus altivelis and S. alascanus)

Determining the sex of thornyheads (Sebastolobus alascanus and S. altivelis) can be difficult under field conditions. We assessed our ability to correctly assign sex in the field by comparing results from field observations to results obtained in the laboratory through both macroscopic and microscopic examination of gonads. Sex of longspine thornyheads was more difficult to determine than that of shortspine thornyheads and correct determination of sex was signif icantly related to size. By restricting the minimum size of thornyheads to 18 cm for macroscopic determination of sex we reduced the number of fish with misidentified sex by approximately 65%.

Reproductive biology of blue marlin (Makaira nigricans) in the western Pacific Ocean

The reproductive biology of blue marlin (Makaira nigricans) was assessed from 1001 fish (ranging from 121 to 275 cm in eye-to-fork length; EFL) caught by Taiwanese offshore longliners in the western Pacific Ocean from September 2000 to December 2001 and from 843 gonad samples from these fish, The overall sex ratio of the catch was approximately 1:1 dur ing the sampling period, but blue marlin are sexually dimorphic; females are larger than males. Reproductive activity (assessed by histology), a gonadosomatic index, and the distribution of oocyte diameters, indicated that spawning occurred predominantly from May to September. The estimated sizes-at-maturity (EFL50) were 179.76 ±1.01 cm (mean ±standard error) for females and 130 ±1 cm EFL for males. Blue marlin are multiple spawners and oocytes develop asynchronously. The proportion of mature females with ovaries containing postovulatory follicles (0.41) and hydrated oocytes (0.34) indicated that the blue marlin spawned once every 2–3 days on average. Batch fecundity (BF) for 26 females with the most advanced oocytes (≥1000 μm), but without postovulatory follicles, ranged from 2.11 to 13.50 million eggs (6.94 ± 0.54 million eggs). The relationships between batch fecundity (BF, in millions of eggs) and EFL and round weight (RW, kg) were BF = 3.29 × 10 –12 EFL5.31 (r2 = 0.70) and BF = 1.59 × 10–3 RW 1.73 (r2= 0.67), respectively. The parameters estimated in this study are key information for stock assessments of blue marlin in the western Pacific Ocean and will contribute to the conservation and sustainable yield of

Analysis of sex-specific spawning biomass per recruit of the sailfish (Istiophorus platypterus) in the waters off eastern Taiwan

Analyses of sex-specific yield per recruit and spawning stock biomass per recruit were conducted to evaluate the current status of the sailfish (Istiophorus platypterus) fishery in the waters off eastern Taiwan. Natural mortality rates estimated from Pauly’s empirical equation were 0.26/yr for females and 0.27/yr for males. The current fishing mortality rates were estimated as 0.24/yr and 0.43/yr for females and males, respectively, which are much lower than the estimated F0 .1 (0.62/yr and 0.79/yr for females and males, respectively) and FSSB40 (0.46/yr for females) which are commonly used as target reference points in fisheries management. The effects of the fishing mortality, natural mortality, and age at first capture on the estimates of biological reference points were evaluated by using the Monte Carlo simulation. The results indicate that failure to consider the uncertainty in parameters such as natural mortality or age at first capture may lead to the improper estimation of biological reference points. This study indicates the possibility of current fishing mortality exceeding the target biological reference points may be negligible for sailfish in the waters off eastern Taiwan. However, in view of the recent rapid increase in fishing effort, it is evident that the stock status and development of the fishery need to be closely monitore

The western blue groper (Achoerodus gouldii), a protogynous hermaphroditic labrid with exceptional longevity, late maturity, slow growth, and both late maturation and sex change

The western blue groper (Achoerodus gouldii) is shown to be a temperate protogynous hermaphrodite, which spawns between early winter and mid-spring. Because A. gouldii changes body color at about the time of sex change, its color can be used as a proxy for sex for estimating the size and age at sex change and for estimating growth when it is not possible to use gonads for determining the sex of this fish. The following characteristics make A. gouldii highly susceptible to overfishing: 1) exceptional longevity, with a maximum age (70 years) that is by far the greatest yet estimated for a labrid; 2) slow growth for the first 15 years and little subsequent growth by females; and 3) late maturation at a large total length (TL50 = 653 mm) and old age (~17 years) and 4) late sex change at an even greater total length (TL50 = 821 mm) and age (~35 years). The TL50 at maturity and particularly at sex change exceeded the minimum legal total length (500 mm) of A. gouldii and the lengths of many recreationally and commercially caught fish. Many of these characteristics are found in certain deep-water fishes that are likewise considered susceptible to overfishing. Indeed, although fishing effort for A. gouldii in Western Australia is not particularly high, per-recruit analyses indicate that this species is already close to or fully exploited.

Distribution of red deepsea crab (Chaceon quinquedens) by size and sex in the Gulf of Mexico

The red deepsea crab (Chaceon quinquedens (Smith, 1879)) has supported a commercial fishery off the coast of New England since the 1970s (Wigley et al., 1975) and has had annual harvests from 400 metric tons (t) (1996) to 4000 t (2001) (NEFMC, 2002). In 2002, a fishery management plan for the northeast fishery on the Atlantic coast was implemented and total allowable catch was reduced to approximately 2500 t (NEFMC, 2002). Although there are management plans for the golden crab (C. fenneri) and the red deep sea crab for Atlantic coast regions, there is no fishery management plan for red deepsea crabs in the Gulf of Mexico. Successful management for sustainable harvests should be based on a knowledge of the life history of the species, but C. quinquedens has been a difficult species for which to obtain life history and abundance information because of its deep distribution.