Differences in growth between maturing and non-maturing male Atlantic salmon, Salmo salar L., parr

Sibling male Atlantic salmon parr that matured tended to be the larger fish in January, but their monthly specific growth rates between January and July did not differ from those of non-maturing fish. Maturing fish had lower condition factors in March, but greater increases in condition factor during April, exceeding those of non-maturing males by May. In maturing males, feeding rates between July and September, and specific growth rates in August and September, were lower than those of immature fish. Consequently, the mean size of immatures equalled or exceeded that of maturing males by October. Maturation rates were strongly correlated with increases in mean condition factor only during April.     

Annual variation in energy reserves in male three-spined stickleback, Gasterosteus aculeatm L. (Pisces, Gasterosteidae)

Wet and dry weights of tissue were measured and concentrations of glycogen, lipid and protein were estimated for the liver, gonad and carcass of male sticklebacks from an annual population collected each month over one complete year. In young-of-the-year there is one period of rapid weight gain, in all three body regions (liver, carcass and gonads) but particularly of the carcass, in the autumn and a second in spring and early summer. This is accompanied by an increase in the water content of all three body regions. The gonadosomatic index also increases sharply in spring and early summer. Young sticklebacks accumulate lipid and glycogen slowly during the autumn and winter of their first year of life and more rapidly from late winter to early summer. Thus, the period of most rapid accumulation of these reserves coincides with the time when body weight and gonad maturation are also increasing sharply. Lipid and glycogen levels fall during the reproductive season in those males that breed, so that by July they are reduced to 43% and 37% (respectively) of their peak values in May. Levels of protein increase throughout the year as the fish grow, but in breeding males by July the concentration of protein in the carcass falls to 70% of pre-breeding levels. Breeding males therefore reach the end of the reproductive season with their total energy reserves severely reduced, and consequently they suffer a very high mortality. In contrast, adult males that fail to reproduce survive beyond the breeding season. They continue to gain weight and to accumulate lipid and glycogen from August to September, but these energy reserves fall (to levels comparable to those of post-breeding fish) in December, when these fish probably die. These results demonstrate that in male sticklebacks, growth and gonad maturation can be sustained in parallel with the accumulation of energy reserves, which are subsequently extensively depleted as a result of reproductive activities.     

Seasonal changes in the levels of 11-oxotestosterone and testosterone in the serum of male salmon, Salmo salar L., and their relationship to growth and maturation cycle

Levels of 11-oxotestosterone (17 β-hydroxyandrost-4-ene-3, 11-dione) and testosterone in the blood serum of individually marked adult male Atlantic salmon held in captivity, were measured by radioimmunoassay at approximately monthly intervals for periods of up to 18 months. In addition to peak concentrations of both hormones shown by all the maturing fish at the time to full sexual maturation during October and November, a majority of maturing fish also showed a significant elevation of 11-oxotestosterone during the early months of the year. The possible involvement of this early elevation of 11-oxotestosterone in controlling the mitotic multiplication of spermatogonia is discussed. Weight and length increases expressed as specific values G_W and G_L and weight to length relationships for the maturing males for each sampling period are presented and compared with those of non-maturing fish.     

Effects of sexual maturation, castration, and androgen implants on growth in one- and two-year-old parr in a Baltic Atlantic salmon (Salmo salar L.) stock

The effects of maturation, castration, and androgen implants on growth in 1- and 2-year-old Baltic salmon ( Salmo salar L.) were studied in a stock originating from the Umeälven (Ume river) in northern Sweden. 1 + male parr that later matured were larger than non-maturing parr in the spring. Later in the summer, during the period of gonadal growth, the growth rate of maturing parr was depressed compared to that of non-maturing males. Implantation of capsules with 11-ketoandrostenedione (OA) or testosterone (T) in sexually immature 1-year-old fish, accelerated the specific growth rate in spring and early summer, particularly considering growth in weight, resulting in an increase in the condition factor. In contrast, the growth rate was depressed from late June onwards, the period when the gonads normally develop in 1 + parr. T, but not OA, also promoted sexual maturation. Castration or OA treatment did not affect total summer growth in 2-year-old salmon. This study suggests a causal relationship between androgens and the growth pattern observed in maturing 1 + male parr.     

Qualitative and quantitative models relating otolith zone deposition to growth and condition in sexually mature male and female capelin (Mallotus villosus)

Summary Correspondence analysis (CA) is used to examine the relationship between months, between selected body components/organs and between months and components/organs for male and female capelin from 12 to 37 months of age in Balsfijorden, northern Norway (69°21N:19°06E). The CA profiles have been used to quantify condition with regard to age and season. The major features determining the condition of fish depositing hyaline or opaque otolith zones are itemized using CA, and the data modelled by Multiple Regression (MR) using a cosine curve incorporating the chief mass variables with linear age-dependency. The highest proportion of fish having outer opaque zones are found from ca. June to October, whilst the highest proportion with exterior hyaline zones are found from ca. December to May. There is a clear annual cycle in the deposition of opaque and hyaline otolith zones in both sexes, but there is a significant difference in phase of ca. 12 days between the sexes. There is a significant trend for decreasing proportions of fish having outermost opaque otolith zones with increasing age; this trend is similar in both sexes. The mean level and amplitude of the cycles are similar in both sexes. On the other hand, CA shows a 12-month cycle of storage and utilization of body materials with obvious differences between sexes. In the female protein in the fillet plays a more dominant role than lipid in the fillet in accounting for anabolic and catabolic variations in mass, whereas in the male the opposite is true. Sexual maturation is negatively correlated with fillet (somatic) growth. Variations in stomach weight (with contents) and otolith zone deposition are not significantly correlated. Nevertheless, in both sexes there is a clear positive correlation between stomach weight and liver protein and lipid implying an association between stomach weight and feeding activity. In females, however, maximum liver condition is more positively correlated with exponential ovarian growth, probably through mobilization of lipoproteins for eggs. Qualitative (CA) and quantitative (MR) models have related changes in growth and condition to the type of otolith zone deposited. The fillet of capelin is the major somatic growth site. As CA indicates a negative correlation between fillet reserves and gonad maturation, and the best MR models describing the cycle of otolith zone deposition in males and females (accounting for 93% and 70% respectively of the variation) underlined and quantified the interplay between somatic and gonad growth, it is implicit that the energy balance of the fish will be an important determinant of otolith zone deposition in sexually maturing and mature capelin.     

Gonadal histology and plasma sex steroid concentrations in maturing and mature spring migrants of Caspian lamprey Caspiomyzon wagneri in the Shirud River,southern Caspian Sea,Iran

Maturing degree-days, gonadal histology, and changes in serum sex steroids (progesterone, P; testosterone, T; and 17β-estradiol, E2) were examined in maturing and mature spring migrant Caspian lamprey Caspiomyzon wagneri (Kessler, 1870) in the Shirud River (southern Caspian Sea). Blood and gonad samples were collected from ten fish when they first entered the river (maturing stage) and from ten fish that showed spawning readiness after being held in cages in the river (mature stage). The maturing degree-days of Caspian lamprey from the start of upstream migration to maturation was 208–470°C.day. Serum P and E2 concentrations in maturing females were significantly higher than in maturing males, but in the mature stage, serum P and E2 concentrations of females were lower than males. In both stages, there were no differences in serum concentration of T between females and males. In both males and females, P increased significantly with maturation; T levels likewise appeared to increase, but the difference was not significant. E2 increased significantly with maturation in males, but females showed a significant decrease. Maturing females had similar stage gonads with the germinal vesicle in the polar position. Maturing males had testes that primarily contained secondary spermatocytes with an occasional occurrence of spermatozoa. These results suggest that males mature earlier than females, which is a pattern similar to that found in the sea lamprey.     

Reproduction in puyen, Galaxias maculatus (Pisces: Galaxiidae), in the southernmost extreme of distribution

Reproductive aspects of a peripheral population of Galaxias maculatus are described and the relationship with the physiology and handling of energetic reserves under marginal environmental conditions is investigated. The G. maculatus population of Tierra del Fuego has an extended reproductive season, with differences in timing and duration compared to other populations of continental Patagonia, New Zealand and Australia. Elevated gonadosomatic indexes (I_G) were observed during this period in both sexes (maximum I_G = 33.49% males; 35.94% females). The high abundance of mature males (with high I_G values) on the spawning grounds during the reproductive season suggests that they were waiting for the return of the mature females. Larger females reached total maturation at the beginning of the reproductive season, whereas the size of maturing females diminished toward the end (mean TL = 96 mm, October; 70 mm, February). Both sexes showed an extremely high investment in reproduction, reaching a maximum I_G of about 35% for both sexes. At the beginning of the reproductive season females reached the maximum median I_H (3.37%) and males the minimum (0.96%), suggesting differences in the role of the liver in the management of energetic reserves during sexual maturation. The variation in the fat index (I_F) suggests that fat reserves were used to survive winter (maximum median I_F > 1%, autumn; minimum about 0.2%, spring).     

Reproductive phenology of the Azorean rock pool blenny a fish with alternative mating tactics

Both male and female Parablennius sanguinolentus parvicornis were ready to mate by the end of May. Seventy-five per cent of the 1-year-old males were mature, but only 25% of the females reached maturity at this age. In females, gonads formed an increasingly greater proportion of the total weight with age, but in males the proportion decreased. A bimodal distribution of gonadosomatic indices classes was observed in 2-year-old males (intermediate size), which reflected the lower opportunity that intermediate size males have to spend the ejaculates than have the younger and older males. The liver played an important role in the storage of energy before the reproductive season and reserves were utilized during the breeding season. Physical condition was at its lowest in July in both sexes, but in May, just before the breeding season began, large maturing females had significantly lower eviscerated condition than males. This could be due to females investing in oocyte development, while the priority for the males was to reserve their energy for the parental activities to come. In both males and females, older and bigger animals had lower hepatosomatic and body conditions than younger ones.     

Fat cycling in the mosquitofish (Gambusia affinis): fat storage as a reproductive adaptation

Summary We argue, based on reviewed literature covering reptiles, amphibians, birds, and fish, that fat storage may represent a life history adaptation because it enables an organism to shift in time when resources are allocated to reproduction. We applied these arguments to fat and population cycles in three populations of the mosquito fish, Gambusia affinis. For males, there appeared to be a constant size at maturation during the reproductive season. Mature males became scarce late in the summer. At the same time, immature males delayed maturity and attained much larger sizes; they matured in large numbers in the fall. The amount of stored fat tended to be equal for immature and mature males at all times except in the late summer. In the August samples, when mature males were relatively rare, they also had the lowest level of fat reserves. It appears that the older generation of mature males did not store fat and did not overwinter. At the same time, immature males registered a two to three fold increase in fat reserves. These differences in fat content between mature and immature males disappeared by September, probably because of the recruitment of a new generation of mature males. The reserves were gradually utilized during the winter. Females reproduced from the late spring through mid- to late-summer. They stopped reproducing in the late summer, when there was ample time to produce an additional litter of young. There was an inverse relationship between resources devoted to reproduction and fat reserves. As reproductive allotment decreased in the late summer, fat reserves increased. The magnitude of the change in fat reserves was similar to that displayed by males. The reserves were depleted over the winter. Significant reserves remained at the beginning of the reproductive season the following spring. Reproducing females utilized the remaining reserves significantly more rapidly than non-reproducing females. An analysis of resource availability revealed an overall decrease in food availability in the late summer, coincident with the increase in fat reserves. These cycles are therefore not attributable to changes in resource availability. They instead indicate a change in how resources are allocated by the fish. The trends in the data indicate that fat reserves are used to shift investment in reproduction from the late summer to the following spring. In males, deferring maturity, rather than maturing in August, allows them to store the necessary reserves to survive the winter so that they can mate the following spring. In females, a subset of the fat reserves is intended for producing the first clutch of eggs the following spring. The female pattern corresponds to those reported for a diversity of organisms. The possible advantages of shifting reproductive effort from the fall to the following spring include higher fecundity and higher offspring fitness. The limitations of the methodology and potential directions for future research are discussed.     

Gonad development and sperm motility of the diving beetle Cybister brevis Aubé, 1838 (Coleoptera: Dytiscidae) in response to seasonal changes in Japan

Gonad maturation in wild Cybister brevis Aubé, 1838 from Japan was investigated to determine the physiological mating season. Breeding experiments showed that hatching occurred from May to September, and the peak was observed in July. The gonadosomatic index (GSI) in collected females (ovary development) showed the highest value in May. The GSIs were low in other seasons. On the other hand, the GSIs in collected males (testes and accessory glands) remained at constant values during the experimental period. Interestingly, high sperm motility was exhibited in May and September, whereas it was low in December and March. This indicates the maturation difference between females and males. Females mature only at the beginning of the breeding season (May), but males do so from May to September. This result suggests that males have a wide-ranging maturation period and earlier sexual maturation than females, ensuring a successful fertilization.     

Seasonal patterns of growth, lipid deposition and lipid depletion in anadromous Arctic charr

Seasonal patterns of growth, lipid deposition and lipid depletion were studied in anadromous Arctic charr from a north Norwegian population. Samples were collected in late May when fish were migrating between fresh water and the sea, and in mid-July when the fish re-entered fresh water. A sample of maturing fish captured in mid-July was held in captivity until late September to assess lipid mobilization linked to the final stages of maturation. The fish increased substantially in weight during their 40–50-day summer residence in sea water (immatures from c . 300 to 600 g; maturing fish from c . 500 to 800 g), and body lipid stores were increased approximately fivefold. The carcass (head, skeleton and skin) was the major lipid depot, accounting for c . 50% of the total lipid content when the fish re-entered fresh water from the sea. The muscle of the ascending charr contained 35–40% of the body lipids, whereas the gut and liver each held 4–5%. Body lipid decreased 30–40% during the period between the re-entry of the fish to fresh water and spawning; although lipids were depleted from all depots the carcass and muscle were quantitatively the most important. By the time of spawning, the gonads of the females held almost 25% of the body lipid, whereas in the males the gonads accounted for <3% of the total lipids. Females lost c . 80% of their body lipid during spawning and overwintering, and, consequently, the lipid depots were severely depleted by the early spring. By contrast, lipid depletion in the males amounted to 50–55% of total lipid in the same period. These data suggest that the combination of spawning and overwintering in fresh water imposes a greater load on the females than on the males. It may be that the severity of the depletion prevents females from spawning in successive years.     

Life-history patterns in a southern population of Atlantic salmon

Juvenile Atlantic salmon Salmo salar in the river Esva, Asturias, north-west Spain, developed a bimodal growth pattern during their first growing season. Segregation between the two modes was apparent by late autumn. All fish in the upper modal group (UMG) grew throughout the year and migrated downstream in the following spring (by April) at the age of 14 months. Some lower modal group (LMG) fish (56% of the main stem, 31% and 50% of the two tributaries) silvered like smolts and apparently migrated downstream the same spring, although c. 1·5 months after UMG fish. Larger LMG fish appeared more likely than smaller ones to migrate. LMG fish did not grow in winter, but they grew fast between March and May. Maturing male parr were detected first at 5 months old in July, and they occurred initially among the faster growers, >6·8 cm long. In spite of this, length of maturing males did not change from late summer and throughout the winter, whereas that of non-maturing UMG and LMG fishes continued to diverge from September onwards. These findings suggest that favourable conditions for growth (very early start of a long growing season) at the southern limit of the species' range may influence the life-history pattern of this population.     

Seasonal variation of mass, condition and gonadosomatic index in fourhorn sculpin (Myoxocephalus quadricornis) with normal and malformed vertebrae in the Gulf of Bothnia

It may be expected that vertehral anomaties would reduer the viability and reproductive success of fish. In the present study. the relationship of vertebral defects to body mass. Iength, age, condition, gonadosomatic index and growth in fourhorn sculpin, Myoxocephalus quadricornis , in the Gulf of Bothnia was studied. There were no differences in growth between normal and defective lish, for either males or females. While normal and defectives males were similar in age and length. defective females were older and longer than normal females. Defective males and females had a significantly greater somatic mass that normal fish. However defective females may be later than in normal fish. Hence, vertebral defects do not necessarly reduce the reproductive potential of fourhorn sculpin.     

Reproductive injury in English Sole (Pleuronectes vetulus) from the Hylebos Waterway, Commencement Bay, Washington

The effect of exposure to xenobiotic compounds on ovarian development was investigated in prespawning female English sole (Pleuronectes vetulus) from the Hylebos Waterway, an industrial site in Commencement Bay, WA, contaminated with aromatic hydrocarbons (AHs), polychlorinated biphenyls (PCBs)and other chlorinated compounds, including hexachlorobutadiene (HCBD) and hexachlorobenzene (HCB). Reference sole were collected from Colvos Passage, a nearby site with minimal sediment contaminant concentrations. English sole from theHylebos Waterway had significantly higher concentrations of fluorescent aromatic compounds (FACs) in bile, polycyclic aromatic compound-DNA adducts in liver, and dioxin-like and other selected PCB congeners in liver than sole from Colvos Passage. The Hylebos Waterway animals also showed significant alterations in their pattern of reproductive development when compared to Colvos Passage sole. Hylebos Waterway sole entered vitellogenesis at a nearlier age than Colvos Passage sole, with about 50%of fish below 5 years of age maturing in the Hylebos Waterway as compared to 20% of Colvos Passage sole in this age range, with corresponding increases in plasma estradiol concentrations and GSI in Hylebosfish. However, while the proportion of maturing Colvos Passage females increased with age to over70% for fish 5 years of age or greater, the proportion of maturing females in the Hylebos Waterway remained at about 50%. Moreover, plasma estradiol concentrations and gonadosomatic indices in these sole were depressed. Inhibited reproductive development and increased oocyte atresia in adult fish were correlated with elevated concentrations of FACs in bile. Enhanced growth, as well as exposure to both aromatic and chlorinated hydrocarbons, were associated with precocious maturation in sub adult Hylebos Waterway sole. This revised version was published online in July 2006 with corrections to the Cover Date.     

Sex-Specific Aggression and Antipredator Behaviour in Young Brown Trout

Sex differences in adult behaviour are often interpreted as consequences of sexual selection and/or different reproductive roles in males and females. Sex-specific juvenile behaviour, however, has received less attention. Adult brown trout males are more aggressive than females during spawning and juvenile aggression may be genetically correlated with adult aggression in fish. We therefore tested the prediction that immature brown trout males are more aggressive and bolder than immature females. Because previous work has suggested that precocious maturation increases dominance in salmonids, we included precocious males in the study to test the prediction that early sexual maturation increase male aggression and boldness. Aggression and dominance relations were estimated in dyadic contests, whereas boldness was measured as a response to simulated predation risk using a model heron. Independent of maturity state, males initiated more than twice as many agonistic interactions as females in intersexual contests. However, males were not significantly more likely to win these contests than females. The response to a first predator attack did not differ between sex categories, but males reacted less to a second predator attack than females. Sexual maturity did not affect the antipredator response in males. Since there is no evidence from field studies that stream-living immature male and female salmonids differ in growth rate, it appears unlikely that the sex differences demonstrated are behavioural consequences of sex-specific investment in growth. It seems more likely that sex-specific behaviour arises as a correlated response to sexually selected gene actions promoting differential behaviour in adult males and females during reproduction. Alternatively, sex differences may develop gradually during juvenile life, because a gradual developmental program should be less costly than a sudden behavioural change at the onset of sexual maturity.     

Reproductive biology, length-weight relationship and relative condition of pond raised Tilapia zilli (Gervais)

In 'runted' populations of Tilapia zilli positive correlations were found between maturation stages and the following: gonadosomatic index, gonad weight, fish weight and fish length. The minimum size at maturity was 9·0 cm in males and 11·0 cm in females. The mean fecundity in the 'stunted' females was 2359 eggs and it increased with length ( L ) weight (HO and depth ( D ) of the fish. There was, however, no correlation between fecundity and egg size. The results indicated that growth in this 'stunted' T. zilli population was not isometric. The relative condition factor ( Kn ) which approached one indicates good condition.     

Changes in reproductive strategy in the ruffe during a period of establishment in a new habitat

The pattern of maturation, body size and fecundity was examined in a population of ruffe ( Gymnocephalus cernuus L.) three times during a period of rapid growth, and eventual stabilization, following its introduction to a new habitat. When the ruffe were less common, maturing ruffe were relatively large and immature ruffe relatively small, compared with when the ruffe were abundant. Intermediate ruffe population size showed a maturation pattern intermediate between these two extremes. It is suggested that this pattern of maturation is a response of the ruffe population to changing growth opportunity induced by changing intraspecific competition. This fluctuating maturation pattern is interpreted in terms of a threshold-dependent maturation trigger, operating on the rate of accumulation of energy and a trade-off between somatic growth and gonad development. When the ruffe population was large, high intraspecific competition resulted in low opportunity for growth; only fish with the highest rate of food acquisition were able to mature in a given year–the investment in gonadal tissue reducing somatic growth. When the ruffe population was low, the high rate of energy acquisition in the population resulted in the triggering of maturation, even at small size, only the very smallest fish remaining immature. High growth opportunity allowed maturing fish to develop gonad and maintain somatic growth. The pattern of size related fecundity also changed over the three periods. When growth opportunity was low, size related fecundity was greater than when opportunity for growth was high. This suggests that maturing females faced with poor growth conditions compensated by increasing egg number for a given body size either by decreasing egg size or by increasing total investment in ovarian tissue.     

The influence of photoperiod on sexual development in underyearling rainbow trout, Salmo gairdneri Richardson

Photoperiods of 19 h light or more induced sexual maturation in male underyearling rainbow trout but did not influence maturation in the females. Early maturation was not a consequence of the increased growth under long photoperiods. The rate of mortality was much higher in the early maturing males than in the immature fish.     

Sex‐specific reproductive investment of summer spawners of Illex argentinus in the southwest Atlantic

Energy investment in reproduction and somatic growth was investigated for summer spawners of the Argentinean shortfin squid Illex argentinus in the southwest Atlantic Ocean. Sampled squids were examined for morphometry and intensity of feeding behavior associated with reproductive maturation. Residuals generated from length‐weight relationships were analyzed to determine patterns of energy allocation between somatic and reproductive growth. Both females and males showed similar rates of increase for eviscerated body mass and digestive gland mass relative to mantle length, but the rate of increase for total reproductive organ weight relative to mantle length in females was three times that of males. For females, condition of somatic tissues deteriorated until the mature stage, but somatic condition improved after the onset of maturity. In males, there was no correlation between somatic condition and phases of reproductive maturity. Reproductive investment decreased as sexual maturation progressed for both females and males, with the lowest investment occurring at the functionally mature stage. Residual analysis indicated that female reproductive development was at the expense of body muscle growth during the immature and maturing stages, but energy invested in reproduction after onset of maturity was probably met by food intake. However, in males both reproductive maturation and somatic growth proceeded concurrently so that energy allocated to reproduction was related to food intake throughout the process of maturation. For both males and females, there was little evidence of trade‐offs between the digestive gland and reproductive growth, as no significant correlation was found between dorsal mantle length‐digestive gland weight residuals. The role of the digestive gland as an energy reserve for gonadal growth should be reconsidered. Additionally, feeding intensity by both males and females decreased after the onset of sexual maturity, but feeding never stopped completely, even during spawning.     

Expression of IGF-I and IGF-I receptor in male and female sterlet, Acipenser ruthenus--evidence for an important role in gonad maturation

Recent in vitro studies suggest that insulin-like growth factor I (IGF-I) is involved in cell differentiation and steroidogenesis in the gonad and could therefore function as an important trigger in vivo. In this study, sensitive real-time RT-PCR assays were used to determine IGF-I and the IGF-I receptor (IGF-IR) mRNA expression in maturing male and female sterlet (Acipenser ruthenus) over a period of two years: In the first year, females entering vitellogenesis (maturing female group, MFG) revealed an increase of IGF-I expression in the ovaries in contrast to females that did not enter vitellogenesis (non-maturing female group, NMFG). Congruently, IGF-IR expression was elevated in females at the onset of vitellogenesis (MFG), decreased towards the first winter, and increased to similar levels at late vitellogenesis in the second winter just prior to spawning. In the second year, NMFG reached the onset of vitellogenesis. Here, IGF-I and IGF-IR reached similar levels as previously observed in the first year in MFG. In males, low and constant IGF-I expression was observed in the testis, whereas IGF-IR was expressed at a constant high level comparable to those of females entering vitellogenesis. These findings suggest an involvement of IGF-I as an important paracrine regulator of gonad maturation, particularly in the ovary.