Seasonal variations in the physiological status and energy content of somatic and reproductive tissues of chub

The seasonal fluctuations in condition, nutrition and somatic energy content, and gonad development cycle were investigated in chub Leuciscus pyrenaicus from the headwaters of the Guadalete River, a freshwater ecosystem characterized by strong seasonal fluctuations in discharge and water level, temperature and food supply. The relationship between somatic stage and gonad cycle was also investigated and discussed. Condition, nutrition and somatic energy cycles could be divided into two distinct periods: from April to January with summer decreases and autumn increments, which is common for juveniles and mature fish; and from January to March, when juveniles and mature fish displayed different temporal variations related to the reproductive cycle. Gonad development took place from the end of the winter into the summer, the testes developing before the ovaries. Spawning started in late spring (May) and continued into summer (June and July), with fish quiescent by autumn (September). The results suggest that, for L. pyrenaicus , both environmental factors (e.g. food supply, water temperature) and reproduction needs affect the condition, nutrition and somatic energy storage of fish, which have been used as indicators of the physiological status of the population.     

The seasonal cycle of condition in the pollan, Coregonus autumnalis pollan Thompson, of Lough Neagh, Northern Ireland

The length–weight relationship for Lough Neagh pollan was log W= 3·2 log L –2·24. Covariance analysis indicated that there were no differences between the sexes, with maturity, or between years. While a seasonal cycle in relative condition was in phase with gonad development during late summer and autumn, changes in somatic condition presented a more complex pattern. Somatic condition increased in spring, as did the quantity and diversity of food consumed. Despite good conditions for feeding and growth, there was an autumn fall in somatic condition, caused by the diversion of energy to gonad growth.     

Sex differences in the energetic content of reproductive tissues in the Blackeye Goby,Rhinogobiops nicholsii

Differential energetic investment in reproduction between the sexes has been a driving a force of life history theory and sexual selection. However, reproductive costs between the sexes have often been based on morphology, such as gonad mass and gonadosomatic indices (GSI), and few have directly measured the energy content of gonadal tissues in relation to GSI. Using the blackeye goby, Rhinogobiops nicholsii, we measured the energetic content of whole gonadal tissues, specifically testes, ovaries and associated reproductive tissues using oxygen bomb calorimetry. The energy content per gram unit of gonadal tissues was generally predictive of GSI, indicating that GSI is a reasonable measure of energetic costs. Interestingly, although females had greater gonadal mass, GSI and energy content per gram than males, the sex difference in energy content per mass unit was only 13 %, suggesting that gross indices such as gonadal mass or GSI may overestimate energetic costs where instead the cost difference in a unit gram of gonadal tissues between the sexes is smaller than often predicted. This study also demonstrates that although the cost of ovaries is greater than testes, males’ investment in reproductive tissue can be considerable, which is consistent with the often inflated reproductive success for males in haremic mating systems.     

Seasonal changes in endogenous ABA and IAA and the influence of applied ABA and auxin in relation to shoot growth and abscission in Valencia Orange (Citrus sinensis (L.) Osbeck)

Seasonal changes in endogenous IAA and ABA were measured by gas chromatography/mass-spectrometry. Highest concentrations of ABA occurred in leaves. There was a major ABA peak in early spring (up to 1360 ng g^–1 dw). Levels were low in summer (90 ng g^–1 dw). There was a minor ABA peak in autumn. Endogenous IAA in leaves was highest in winter/spring (up to 76 ng g^–1 dw). Applied ABA promoted abscission of leaves and shoots while applied NAA delayed abscission. The main peak in leaf-ABA content was followed by extensive shoot abscission. The involvement of ABA and IAA in regulation of flush growth was not clear.This paper is dedicated to Michael G. Mullins, who died on 13 November 1990, for his outstanding contribution to horticulture.     

Reproductive energy expenditure and changes in body morphology for a population of Chinook salmon Oncorhynchus tshawytscha with a long distance migration

Energetic demands of a long freshwater migration, extended holding period, gamete development and spawning were evaluated for a population of stream‐type Chinook salmon Oncorhynchus tshawytscha. Female and male somatic mass decreased by 24 and 21%, respectively, during migration and by an additional 18 and 12% during holding. Between freshwater entry and death after spawning, females allocated 14% of initial somatic energy towards gonad development and 78% for metabolism (46, 25 and 7% during migration, holding and spawning, respectively). Males used only 2% of initial somatic energy for gonad development and 80% on metabolic costs, as well as an increase in snout length (41, 28 and 11% during migration, holding and spawning, respectively). Individually marked O. tshawytscha took between 27 and 53 days to migrate 920 km. Those with slower travel times through the dammed section of the migration corridor arrived at spawning grounds with less muscle energy than faster migrants. Although energy depletion did not appear to be the proximate cause of death in most pre‐spawn mortalities, average final post‐spawning somatic energy densities were low at 3·6 kJ g^?1 in females and 4·1 kJ g^?1 in males, consistent with the concept of a minimum energy threshold required to sustain life in semelparous salmonids.     

Seasonal variation of energy metabolism in ghost crab Ocypode quadrata at Siriú Beach (Brazil)

The aim of this study was to analyze the seasonal variations of carbohydrate and lipid metabolism of the ghost crab Ocypode quadrata (Fabricius, 1787) on a sandy beach in the southern region of Brazil. Crabs and hemolymph samples were collected monthly in the field. Hepatopancreas, gills, gonads and claw muscles were used for glycogen determination. In males, blood glucose levels increased in the summer and in the winter. The glycogen values increased significantly in the hepatopancreas in the winter, but remained constant in the muscle, gonads and gills. In females, hemolymph glucose levels, glycogen values in the hepatopancreas and in the gills remained constant throughout the year; however, muscular glycogen increased in the spring and gonad glycogen decreased in the summer. Hemolymph triglyceride levels of males and females and total cholesterol of males decreased significantly in the spring. In females, a significant increase of total cholesterol levels was found in the winter. The findings suggest that in O. quadrata lipids seem to be an important reserve of energy used during reproduction, both in males and females, while glycogen may be used during periods of intense activity or fasting.     

Seasonal variation of energy metabolism in ghost crab Ocypode quadrata at Siriú Beach (Brazil)

The aim of this study was to analyze the seasonal variations of carbohydrate and lipid metabolism of the ghost crab Ocypode quadrata (Fabricius, 1787) on a sandy beach in the southern region of Brazil. Crabs and hemolymph samples were collected monthly in the field. Hepatopancreas, gills, gonads and claw muscles were used for glycogen determination. In males, blood glucose levels increased in the summer and in the winter. The glycogen values increased significantly in the hepatopancreas in the winter, but remained constant in the muscle, gonads and gills. In females, hemolymph glucose levels, glycogen values in the hepatopancreas and in the gills remained constant throughout the year; however, muscular glycogen increased in the spring and gonad glycogen decreased in the summer. Hemolymph triglyceride levels of males and females and total cholesterol of males decreased significantly in the spring. In females, a significant increase of total cholesterol levels was found in the winter. The findings suggest that in O. quadrata lipids seem to be an important reserve of energy used during reproduction, both in males and females, while glycogen may be used during periods of intense activity or fasting.     

Gonad development and filleting yield of common carp Cyprinus carpio L. reared in ponds in Eastern France

The processing of 232 common carp Cyprinus carpio made it possible to study the correlation between gonad maturation and filleting yield. The study was undertaken between April and August 1992 and was carried out on 4-year-old carp cultured in extensive ponds. Morphological parameters were recorded and relations between total weight and length, biomass of fillet, and biomass of gonad were determined. The histology of male and female gonads was examined in relation to the gonadosomatic index (GSI: up to 16% for females and less than 11% for males) and condition indices (total K or somatic K1). Filleting yields (32–36% of the total weight) decreased for females just prior to spawning and corresponded to a decline of K1. A rematuration phenomenon was observed, although females weighing less than 1200g were still immature. We suggest avoiding the filleting of large females before spawning and suggest estimating the date of appearance of this stage by referring to the positive correlation established in this paper between the number of degree-days and total fish weight.     

Testosterone and estradiol potentiate the serum gonadotropin response to gonadotropin-releasing hormone in goldfish

The effects of gonadal steroids on gonadosomatic index (GSI; gonad wt/total body wt x 100), pituitary gonadotropin (GTH) content, and serum GTH response to [D-Ala6,Pro9-Net]-luteinizing hormone-releasing hormone (LHRH-A) were investigated throughout the seasonal reproductive cycle of the goldfish. Gonad-intact female fish were implanted i.p. for 5 days with silastic pellets containing no steroid (blank), testosterone (T; 100 micrograms/g), or estradiol (E2; 100 micrograms/g). The serum GTH response at 6 h following i.p. injection of saline or 0.1 microgram/g LHRH-A was assessed. In blank-implanted, saline-injected animals, seasonal variations in GSI, pituitary GTH content, and serum GTH levels were evident; maximal and minimal levels were noted in the spring and summer months, respectively. In blank-implanted fish, LHRH-A effectively stimulated GTH release in females undergoing gonadal recrudescence (late autumn and winter) and in sexually mature (spring) females, but not in sexually regressed (summer and early autumn) females. Implantation of T or E2 raised serum steroid levels to those found during ovulation in goldfish. Steroid treatments did not affect unstimulated serum GTH levels at any time of the year. Testosterone effectively potentiated the serum GTH response to LHRH-A during the entire reproductive cycle, whereas the positive effects of E2 were evident in sexually regressed and post-spawning females only. Both T and E2 potentiated the GTH response to LHRH-A in male fish. To examine the involvement of T aromatization in mediating its actions on induced GTH secretion, male and female fish were implanted with T or the nonaromatizable androgens 5 alpha-dihydroxytestosterone (DHT; 100 micrograms/g) and 11-keto-testosterone (11-KT; 250 micrograms/animal). Testosterone potentiated the GTH response to LHRH-A in both males and females whereas DHT and 11-KT were without effect. Furthermore, the positive action of T on induced GTH secretion was blocked by 2-day pretreatment with the aromatase inhibitor 1,4,6-androstatrien-3,17-dione (100 or 300 micrograms/g). Multiple i.p. injections of hCG (0.2 microgram/g every 3 days for 39 days), probably through stimulation of endogenous T secretion, resulted in potentiation of the GTH response to LHRH-A in mature male goldfish. These results clearly demonstrate that T, through aromatization to E2, can increase pituitary responsiveness to exogenous LHRH-A in gonad-intact male and female goldfish.     

Production of Excirolana armata (Dana, 1853) (Isopoda, Cirolanidae) on an exposed sandy beach in southeastern Brazil

The somatic and gonad productions of the cirolanid isopod Excirolana armata were analyzed by taking monthly samples from December 2003 to November 2005 on Una beach, S?o Paulo state (24°S), southeastern Brazil. Sampling was performed along three fixed transects established from the base of the foredunes to the waterline. Weight-specific growth rate was used to estimate the E. armata somatic production for 2004 and 2005, separately. The gonad production was estimated based on the monthly reproductive potential (mean number of eggs/embryos per female × monthly abundance of ovigerous females with near-release broods) for 2004. The annual somatic production of E. armata population varied from 15.57 to 17.25?g AFDW m^?1?year^?1 and the somatic production/biomass ratio (P _s/B) from 3.55 to 3.14?year^?1 for 2004 and 2005, respectively. The P _s/B ratios were higher for males (4.02 and 3.19?year^?1 for 2004 and 2005) than for females (3.10?year^?1 for both years). The annual gonad production (P _g?=?1.07?g AFDW m^?1?year^?1) contributed about 15 and 6% to the total production (P _s?+?P _g) of females and the population, respectively. The proportion of gonad to somatic production of females (P _g/P _s) increased with individual size (ca 90% in the 7.5?mm size class), and the annual weight-specific gonad production (P _g/B ratio) was estimated to 0.24?year^?1. The high P _s/B ratios estimated for E. armata derive from the fast growth of individuals and show the importance of this population to the energy flow on Una beach ecosystem. However, the low percentage of juveniles verified in this population and in other studies of populations of the genus Excirolana is discussed as an important source of underestimation of P _s/B ratio.     

Gonadal maturation,fecundity, spawning,and timing of reproduction in the mud snail,Cerithidea cingulata,a pest in milkfish ponds in the Philippines

Summary

Gonadal maturation, spawning, fecundity and timing of reproduction of the snail Cerithidea cingulata in a brackish water pond in Molo, Iloilo, Philippines, are described. Snails 4–41 mm in shell length were sampled monthly from May 1997 to May 1998; 25% were <25 mm, 67% were 20–30 mm, and 8% were >30 mm. The sexes are separate and could first be distinguished at 15 mm. Males are aphallic, have narrower shells than females of the same length, and have bright yellow-orange testes overlying the digestive gland deep inside the shell. Females have more robust shells, an ovipositor at the right side of the foot, and yellow-green ovaries overlying the digestive gland. The sex ratio was one male to two females in the pond population studied. Gonadal maturation was monitored by means of gonadosomatic index (GSI, gonad weight as a percent of visceral weight); maturation stages were based on the gonad appearance (immature, developing, mature) and histology (immature, developing, mature, redeveloping). GSI increased with snail size, and reached 16% in a 33-mm female. The smallest mature males and females were 18–19 mm, and most snails >20 mm were mature, spawning, or redeveloping. Histological sections showed all stages of gametogenesis in mature male snails. The oocyte size-frequency distributions in mature females showed mostly mature oocytes and secondary oocytes, but also oogonia and primary oocytes. GSI and the frequency of snails at different maturation stages varied over the year. Both GSI and the frequency of mature snails were highest during the summer months, April to August. Nevertheless, mature snails occurred throughout the whole year, as did mating and egg-laying. Fecundity (= number of oocytes >70 pμ) increased with size in mature females 2041 mm; an average 25-mm female produced about 1,500 oocytes and larger females produced a maximum of about 2,500 oocytes. Eggs strings laid on the pond bottom were 45–75 mm long; an average 64-mm string contained 2,000 eggs 210+20 pm in diameter. The density of eggs strings was highest (80–120/m²) during March-September. Eggs hatched after 6–7 d into planktonic veligers, which in turn settle on the pond bottom 11–12 d later as juveniles. Juveniles 2–6-mm long were most abundant in the pond during August-October.     

The reproductive biology of an endemic cyprinid,Zacco pachycephalus,in Taiwan

Synopsis Reproductive parameters including body weight-length relationship, condition factor, gonadosomatic index (GSI) and fecundity of an endemic cyprinid,Zacco pachycephalus, were studied from October 1991 to January 1993. Although the condition factor was similar between both sexes, GSI values of female (3.72 to 14.24%) were significantly larger than that of males (0.21 to 2.23%) during the major breeding season. The number of yolky eggs per fish ranged from 459 to 2059. Ovum size distribution, annual GSI value, and recruitment of juvenile fish indicated thatZ. pachycephalus conducted two major spawning activities in spring (February to April) and summer (June to August). The larger GSI value and number of yolky eggs in spring compared to those in summer implies thatZ. pachycephalus has evolved a reproductive style by investing more energy in spring to avoid the damage of summer freshet on its recruitment success. The mating behavior ofZ. pachycephalus is similar to that of otherZacco fishes.     

Sex steroids,gonadal histology and biological indices of fall and spring Caspian lamprey (<Emphasis Type="Italic">Caspiomyzon wagneri</Emphasis>) spawning migrants in the Shirud River,Southern Caspian Sea

This study investigates sex steroids, gonadal histology and some biological indices of fall and spring spawning migrants of Caspian lamprey Caspiomyzon wagneri (Kessler, 1870). Blood and gonad samples were collected from 15 migrants during fall and spring. Serum sex steroid levels including testosterone (T), 17β-estradiol (E₂) and progesterone (P) were determined with ELISA and gonadal development was studied using conventional histological methods. Our results showed serum E₂ level in females were similar in the two seasons but in males, fall migrants had higher serum E₂ concentrations. No differences were found in T levels in fall and spring migrants. Serum P concentration in fall migrant males were significantly higher than spring migrant males while spring migrant females had higher serum P levels compare to fall migrant females. Gonads in both fall and spring migrants were in the final stage of maturity. Fall and spring migrant males had similar HSI and GSI; fall migratory female had significantly higher HSI and GSI. Sex ratios were close to 1:1 in both seasons. There was a significant relationship between fecundity and length in both seasons. Comparison of fecundity and egg diameter between fall and spring seasons showed no significant difference. These results indicate that both fall and spring migrants Caspian lamprey were ready for spawning. Further studies are needed to clarify when spawning occurs in Caspian lamprey.     

Allocation of resources in an European cyprinid during maturation period as measured by nucleic acids

Changes in the nucleic acid levels of different tissues during a regular gonadal maturation were used to investigate the sex-related size differences in Iberian barbel, Barbus sclateri. Gonadal DNA concentration was associated with gonadal development in both sexes. There was a decline in DNA when ovaries were ripe. In contrast, the maximum size of the testes was related with highest gonadal DNA concentrations. Gonadal growth was not related with liver RNA : DNA ratios of male, while it was with female ratios. White muscle DNA concentrations indicate that somatic cell volume decreased during gonadal growth in both sexes. This suggests the translocation of several elements toward the gonad. Moreover, during gonadal maturation period adult barbel showed no change in their somatic growth. However, males apparently were in poorer metabolic condition (lower muscle RNA : DNA ratio) than females. Females may have allocated additional energy to self-maintenance rather than to breeding effort and it might influenced the cumulative annual growth. One might reasonably assume that there is a trade-off between investment in current versus future reproductive success in female. This result was not seen in males.     

The evolution of gonad expenditure and gonadosomatic index (GSI) in male and female broadcast‐spawning invertebrates

Sedentary broadcast‐spawning marine invertebrates, which release both eggs and sperm into the water for fertilization, are of special interest for sexual selection studies. They provide unique insight into the early stages of the evolutionary succession leading to the often‐intense operation of both pre‐ and post‐mating sexual selection in mobile gonochorists. Since they are sessile or only weakly mobile, adults can interact only to a limited extent with other adults and with their own fertilized offspring. They are consequently subject mainly to selection on gamete production and gamete success, and so high gonad expenditure is expected in both sexes. We review literature on gonadosomatic index (GSI; the proportion of body tissue devoted to gamete production) of gonochoristic broadcast spawners, which we use as a proxy for gonad expenditure. We show that such taxa most often have a high GSI that is approximately equal in both sexes. When GSI is asymmetric, female GSI usually exceeds male GSI, at least in echinoderms (the majority of species recorded). Intriguingly, though, higher male GSI also occurs in some species and appears more common than female‐biased GSI in certain orders of gastropod molluscs. Our limited data also suggest that higher male GSI may be the prevalent pattern in sperm casters (where only males release gametes). We explore how selection might have shaped these patterns using game theoretic models for gonad expenditure that consider possible trade‐offs with (i) somatic maintenance or (ii) growth, while also considering sperm competition, sperm limitation, and polyspermy. Our models of the trade‐off between somatic tissue (which increases survival) and gonad (which increases reproductive success) predict that GSI should be equal for the two sexes when sperm competition is intense, as is probably common in broadcast spawners due to synchronous spawning in aggregations. Higher female GSI occurs under low sperm competition. Sperm limitation appears unlikely to alter these conclusions qualitatively, but can also act as a force to keep male GSI high, and close to that of females. Polyspermy can act to reduce male GSI. Higher male than female GSI is predicted to be less common (as observed in the data), but can occur when ova/ovaries are sufficiently more resource‐intensive to produce than sperm/testes, for which some evidence exists. We also show that sex‐specific trade‐offs between gonads and growth can generate different life‐history strategies for males and females, with males beginning reproduction earlier. This could lead to apparently higher male GSI in empirical studies if immature females are included in calculations of mean GSI. The existence of higher male GSI nonetheless remains somewhat problematic and requires further investigation. When sperm limitation is low, we suggest that the natural logarithm of the male/female GSI ratio may be a suitable index for sperm competition level in broadcast spawners, and that this may also be considered as an index for internally fertilizing taxa.     

Age, growth and reproduction of the barbel, Barbus sclateri (Günther, 1868), in a first-order stream in southern Spain

This study was based on examination of 1476 barbels from a first-order stream located in the Guadalquivir River basin (38°N, 4°43'W). This stock comprised nine age groups of male and 11 of females. No growth was recorded between October and March, most occurring in April–June and, to a lesser extent, in July–September. A classification analysis revealed that this stock had the lowest growth rate of 37 different European barbel populations. Length–weight relationships were obtained for 12 barbel categories and used to estimate both condition and instantaneous growth rate. Relative condition (before and after subtracting gonad weight) was similar in both sexes and was affected by gonad growth and environmental summer conditions.
Spawning occurred during the second half of May (15 May is suggested as the birthday). Gonads began to develop in September (females) and in February (males). Males matured at between 7 and 9 cm fork length (f.l.) (2–4 years old) and females between 11 and 16 cm (6–7 years). The fecundity of this stock was represented by the formula F =6.07 10 ⁴ f.l. (mm)^3.0667. Larger and older fish showed a higher fecundity and bigger eggs. The overall sex ratio did not differ from 1:1.     

Energetic composition,reproductive output,and resource allocation of Antarctic asteroids

Summary Thirteen species of antarctic asteroids were collected in McMurdo Sound during the austral spring and summer of 1984–1985 and 1985–1986. Variability in the energetic composition of the body wall, pyloric cecum, and gonad was directly related to differences in levels of protein, carbohydrate, lipid and ash in these tissue types. Reproductive output was generally similar between males and females when expressed in gravimetric units. When considered in energy units, females generally had greater reproductive output than males. Reproductive output values of antarctic asteroids were similar to published values for temperate, tropical, and subpolar species. In a given year, the majority of antarctic asteroids have the greatest energy resources in the body wall, a tissue representing a long-term energy investment. Lesser amounts of energy are devoted to annual nutrient storage (pyloric ceca) and reproduction (gonads).     

Seasonal changes in energy and the energy cost of spawning in Gulf of Alaska Pacific cod

In the Gulf of Alaska, adult Pacific cod exhibited an annual cycle of condition, gonad index and liver index in which maximum values occurred in ripe fish in March and minima in July. About 30–31 % of prespawning stored energy was expended during the spawning effort. The energy associated with spawning derived from liver (24% and 18%), somatic tissue (22% and 33%) and gonad (53% and 48%) for females and males, respectively. Liver index and gonad index at the time of sampling were directly related in females, but in males gonad index was best related to liver index 1–3 months earlier.
The Pacific cod is very similar to the Atlantic cod in terms of energy cycling, maximum gonad sizes, energy expended during spawning and gonadal contribution to energy expenditure. However, in Pacific cod, somatic tissue contributes markedly to energy expended during reproduction. The Pacific cod cod differs from the walleye pollock with respect to gonad index (13% and 20%ν. 20% and 8% for females and males, respectively), spawning weight loss (25%ν. 38%), liver energy loss during spawning (71%ν. 55%) and energy cost of spawning.     

Onset of maturity in male southern rock lobsters Jasus edwardsii in Tasmania,Australia

Summary

The reproductive contribution of male southern rock lobsters Jasus edwardsii has recently become of interest for managing this fishery as changes to the length of seasonal closures have shifted fishing effort towards males. To assess size at onset of maturity, a total of 387 male southern rock lobsters Jasus edwardsii were collected using baited traps from four of the eight fishery regions around the Tasmanian coast during July and August 2000. Sizes at onset of maturity in these samples were estimated by examining the vasa deferentia for evidence of the commencement of spermatid production. In addition, the total combined weight of the vasa deferentia and testis was compared with body size (as a gonad somatic index, GSI). Estimates of size at onset of maturity from the two methods were similar, although estimates from GSI appeared to have broader confidence limits. Although these physiological indices of maturity do not necessarily translate to functional maturity, they indicate that maturity is well below that of the legal minimum size of 110 mm carapace length used for managing the fishery (all estimates by either method for length at 50% sample maturity ≤70 mm carapace length). Production of sperm by males occurs below the size at onset of maturity of females from the same region and spatial patterns in maturity estimated by onset of spermatid production were similar to those of females.