Seasonal changes in condition, nutrition, gonad maturation and energy content in barbel, Barbus sclateri, inhabiting a fluctuating river

The objective of this study was to investigate the seasonal cycle of condition, nutrition and gonad development, as well as the magnitude of seasonal variations in energy content of somatic and gonad tissues in juveniles, males and females of Sclater's barbel, Barbus sclateri, from the upper Guadalete River (south Spain). The influence of reproductive cycle on somatic changes was also investigated and discussed. Measurements of condition factor (K), somatic index (SI), gonadosomatic index (GSI) and somatic and gonad energy content (J g^-1 dw) were made in individual specimens taken from the Guadalete River monthly for 12 months. This freshwater ecosystem is characterised by strong seasonal fluctuations in water and flow level, temperature and food supply. It was found that in general juveniles, males and females of barbel exhibited a similar condition, nutrition and somatic energy cycle throughout the year, with the highest values in spring and the lowest in summer. Depletion of K, SI and somatic energy storage from spring until mid-summer seems to be associated with high metabolic demands during this period, and in adult fish also with spawning-related activity. The gonad index and energy content of the gonad were the highest in April and the lowest in summer for both sexes. Spawning took place during late spring – early summer, with fish quiescent by mid summer. The energy required for ovarian development (3970 J g^-1 dw) was greater than the one for testes development (2763 J g^-1 dw). Data on gonad energy content indicated a period (March to April) of intense energy accumulation (64% males and 37% females) which was related to the decline in the average somatic energy content in males and females. The somatic energy content was linearly related to K and SI. In the same way, GSI correlated positively with gonad energy. Linear trends were found between somatic parameters (K, SI and energy content) and gonad parameters (GSI and energy content). This revised version was published online in July 2006 with corrections to the Cover Date.     

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.     

Feeding behaviour of cod (Gadus morhua) in relation to spawning

The food consumption and egg production of 26 adult (13 female and 13 male) Atlantic cod ( Gadus morhua ) were monitored during prespawning, spawning and postspawning periods. Females spawned from late January to mid-April. Feeding activity occurred from December to early January and ceased for females, on average, 36 days (15–54 days) before the onset of spawning. The duration of spawning by females was, on average, 42 days (10–61 days) and feeding was suppressed by both sexes during the first three-quarters of each female's spawning period. Mature females went, on average, 70 days or 19% of the year without eating. An abrupt increase in feeding activity, particularly by females, occurred during the last quarter of spawning or shortly after the release of the last egg batch (on average, feeding started again after 91% of a female's eggs had been released or 82% of egg batches). Females consumed greater quantities of food than males during both winter and postspawning feeding periods. During spawning, females lost, on average, 29% of their body weight and males 14%. Fecundity ranged from 0.75 to 3.97 million eggs per female. The volume of eggs produced by four individual females (range = 1285–5995 ml in four to 11 batches) ranged from 99 to 195% (mean 150%) of a female's postspawning body volume. Six immature cod fed throughout the experimental period and gained, on average, 8% of initial body weight. Laboratory results were supported by stomach fullness index values of Georges Bank cod exhibiting different maturity states.     

Interspecific differences in growth of somatic and reproductive tissues during the breeding season in Menidia menidia and M. beryllina

The annual atherinid fishes, Menidia menidia and M. beryllina , are extremely similar morphologically, but appear to differ in their energetics and life histories. They emerge from winter in relatively emaciated condition. Menidia menidia males and females showed little change in carcass weight but significant increases in gonad weight in March and April prior to spawning in May. Female liver weight increased in March and remained high. Thus, this species had reached its reproductive size in the autumn and allocated its energy to gonadal growth during the spring. Menidia beryllina showed no significant increases in weight until early May, when carcass, liver and gonad all increased simultaneously, a pattern very different from its congener. The later spawning of M. beryllina is probably due both to its need for carcass growth in spring prior to spawning and to its apparent requirement for warmer water for growth. Females of both species invested more into liver and gonad than did males. Maximum gonadosomatic index was about twice as large for M. menidia males as for M. beryllina males, but the index was similar for females of both species.     

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.     

The spawning activity of cod, Gadus morhua L.

Aspects of the reproduction of reared cod, Gadus morhua L., with special emphasis on the females, were studied under laboratory conditions. The fecundity and condition factor were 2–5 and 1–5 times, respectively, that of wild cod. A total of 18 spawning females were kept in separate tanks/ chambers, each with one or two males. Seven of the 18 females were classified as stressed, based upon behaviour, irregular spawning intervals and low fertilization rates of the eggs. The reared cod were found to spawn 17–19 batches. The number of eggs liberated in each batch normally followed a smooth, dome-shaped curve with time. The fertilization rate was normally 100%. Egg size decreased from first to last batch and the egg dry weight decreased by about 20–30%. The reared cod showed the same egg diameter to dry weight relation as wild cod. Egg diameter of first batch and maternal fish length were significantly positively correlated. The mean spawning interval for the female and the mean water temperature during its spawning were negatively correlated. The reared cod spawned in both the night and the day for about 50–60 days.     

The multiple spawning pattern of weakfish in the Chesapeake Bay and Middle Atlantic Bight

Weakfish Cynoscion regalis were collected from commercial fisheries in the Chesapeake Bay and the Middle Atlantic Bight (n=4380) during 1989–1992 and their reproductive biology assessed using the gonadosomatic index, macroscopic gonad stages, oocyte diameter distributions, microscopic whole oocyte analysis and histology. Sex ratios were approximately 3:1, females to males, in 1990–1992. Most fish (90%) attained sexual maturity by age 1 and at a small size. Estimated mean length at first maturity was: 164mm total length (TL) for males, and 170 mm TL for females. Weakfish spawn within the Chesapeake Bay, as far north as the Virginia/ Maryland border. Although spawning occurred during May–August and gonad development and initiation of spawning was synchronous, cessation of spawning was asynchronous. There was no indication that older fish exhibited a more extended spawning season than younger fish. Weakfish are multiple spawners with indeterminate fecundity. Oocyte development is asynchronous with oocytes of all stages being present in developed ovaries. Because of the complex and dynamic weakfish ovarian cycle, typical methods of assessing reproduction, such as the GSI and macroscopic gonad stages, are inadequate for this species if not used in conjunction with more detailed methods such as histology.     

Energetic cost of spawning in male and female Atlantic salmon (Salmo salar L.)

Energetic expenditure during spawning of male and female 1 -sea-winter Atlantic salmon, Salmo salar L., was measured. Before spawning, the somatic energy content per unit weight did not differ between the sexes. The testes content offat was 0.24, of carbohydrate 8.89, of water 1.21 and of ash 1.61 times as high as that of the ovaries. Just prior to spawning, mean gonadosomatic index (GSI; wet weight, %) in males was 4.36 and in females 20.26, and expressed as energy ratios (kJ, %) 4.47 and 27.75, respectively. During spawning the energy loss of male soma was higher (35.57%) than that of females (25.00%). This was a result of higher loss of fat in males than in females. Total energy cost of spawning in males and females was on average 51.8 and 51.4%, respectively.     

Energy expenditures during spawning by chum salmon Oncorhynchus keta (Walbaum) in British Columbia

Physiological telemetry and proximate tissue analyses were used to assess energy expended by chum salmon Oncorhynchus keta on various behaviours during spawning in Kanaka Creek, British Columbia, Canada, and results were compared with published data on Fraser River sockeye salmon Oncorhynchus nerka , the only other species for which both types of measurements have been taken. Chum salmon arrived at the spawning grounds with body energy densities of 4·84 MJ kg⁻¹ in males and 4·62 MJ kg⁻¹ in females, lower than most sockeye salmon populations, and died with energy densities of c . 4 MJ kg⁻¹, similar to that observed in sockeye salmon and other salmonids. Moisture levels generally increased in body tissues over the spawning life, particularly in female gonads, and lipid levels decreased. Declines in protein observed over the spawning life of other Pacific salmon Oncorhynchus sp. were less evident in Kanaka Creek chum salmon. Holding behaviour constituted the dominant component of the activity schedule and energy budget of both sexes. After holding, the most expensive behaviours were nest digging in females and aggressive displays in males. Dominant males expended the most energy on behaviours each day, as indexed by oxygen consumption (3600 mgO₂ kg⁻¹), while satellite males expended nearly as much (3504 mgO₂ kg⁻¹) but females expended considerably less (2327 mgO₂ kg⁻¹). Kanaka chum salmon engaged more frequently in energetically expensive reproductive behaviours than Stuart River sockeye salmon.     

Sexual dimorphism in the energetics of reproduction and growth of North Sea plaice, Pleuronectes platessa L.

The chemical composition and energy content of North Sea plaice during the spawning period were examined in mature males and females and in immature fish, to study differences in the allocation of energy over reproduction and somatic growth between the sexes. At the beginning of the spawning period mature males and females had equal dry weights of lipid that were 70% higher than in immatures. Protein content in mature males was equal to that in immatures but was 23 % higher in mature females. Immature males and females did not differ in chemical composition. At the end of the spawning period, spent and immature fish had equal lipid contents, but protein content in spent females was 10% lower than in spent males, and 17% lower than in immatures. Gross energy content of the body decreased by 44% (65·2 to 36·3 J cm^-3) in mature females, 27% (55·0 to 40·OJ cm^-3) in mature males, and 9% (48·7 to 44·2J cm^-3) in immatures. Energy content of plaice eggs was estimated at 6·60 kJ per 1000 eggs. Reproductive investment was estimated from the energy loss during the spawning period and included the energy of sex products and spawning metabolism. Somatic growth comprised the annual increase in energy content of fish. The pattern of energy allocation over reproduction and somatic growth differed between males and females. Males started their reproduction at a smaller length and a younger age and allocated a higher proportion of the available energy into reproduction than females. Available energy resources for somatic growth and reproduction (surplus production) were equal between the sexes up to a length of about 30 cm. Beyond this length male surplus production levelled off whereas female surplus production continued to increase. The differences in surplus production and the allocation patterns are discussed. For female plaice the energy allocated into egg production was estimated as between 48 and 64% of the total amount of energy lost during spawning. The remaining energy is used for metabolism during the spawning period, yielding an estimate of the metabolic rate of mature females of between 6·4 and 9·1 kJ day^-1. A maximum estimate of the metabolic rate of mature males was 7·4 kJ day^-1.     

Gonadal development and associated changes in liver size and sexual steroids during the reproductive cycle of captive male and female Atlantic cod (Gadus morhua L.)

Gametogenesis in female and male Atlantic cod (Gadus morhua L.) was investigated by sampling blood plasma and gonadal tissue from 19 to 33-month-old fish. The reproductive cycles of both female and male Atlantic cod are characterized by distinct annual variations in gonadal size and developmental stage and these are associated with changes in sex steroids and liver size. I(H) did not change during early gonadal development, but both spent females and males had lower I(H) than late maturing females and spermiating males, respectively. In females I(G) was correlated to plasma E2 levels and they were highest in spawning females. The lowest levels during the reproductive cycle were observed in spent females. Plasma T levels were low throughout ovarian development, and were at a minimum in spent females. 11-ketotestosterone in plasma of males increased rapidly during spermiation, while T increased at earlier testicular stages and reached maximum during spermiation. High plasma levels of steroids in male and female cod during spawning serve to promote further development and growth of less advanced stages of germ cells.     

Reproductive investment in the Silurus meridionalis

A comparison of pre- and postspawning Silurus meridionalis showed that 20·7% of body stored energy was utilized during spawning for a standard male (74·5 cm) and 23·8% for a standard female (85·3 cm). About one-third of the loss of the stored energy was released as eggs by females, and almost all of the energy loss for males and about two-thirds for females were expended in metabolism. Stored lipid as fuel for metabolism supplied 90·0% of energy in males and 95·2% in females, and protein supplied the rest of the energy. Models for predicting energy in released gametes ( G _g), deposited in the body as somatic growth ( G _s), utilized in spawning activity ( S _a), expended in maintenance ( M , including metabolism, faeces and excretion), and food energy ( C ) were developed, and annual energy budgets were compiled. The balanced budget for a male aged 4 was: 100 C =0·06 G _g+11·17 S _a+19·5 G _s+69·2 M , and for a female aged 5: 100 C =5·48 G _g+8·51 S _a+15·8 G _s+70·2 M .     

Sexual maturity cycle and spawning of Greenland halibut Reinhardtius hippoglossoides in the Davis Strait

Female sexual maturation cycle and the main spawning time of Greenland halibut Reinhardtius hippoglossoides in the Davis Strait were studied through regularly collected samples during 1 year starting in spring 2003. Samples were collected from the southern slope of the Davis Strait Ridge between Canada and Greenland in the depth range 1000–1500 m. Female sexual maturation was described using different approaches: gonado‐somatic index, visual macroscopic maturity stage index, histological microscopic maturity index and oocyte diameter measurements. A significant increase in the gonado‐somatic index was seen from September onwards until February with a maximum estimated value of 18%. The proportion of mature fish increased from December until March. At the same time, the proportion of females with a low gonado‐somatic index also increased from February, indicating that spawning had occurred and females were recovering. Oocyte diameter distribution revealed a leading cohort development during autumn through to December to February. A coupling between sexual maturity and fish condition was seen for females in maturing condition indicating a steady build up of stored energy in the liver from June to November.     

Biochemical composition of the Atlantic bonito Sarda sarda from the Aegean Sea (eastern Mediterranean Sea) in different stages of sexual maturity

The content (% wet mass) in water, ash, lipid, crude protein, DNA and RNA of different tissues was determined during sexual maturation of bonitos Sarda sarda from the Aegean Sea. A total of 220 specimens were collected in the following stages of sexual maturity: immature, resting, developing, mature, spawning and spent. Highest lipid levels in the white muscle, red muscle and liver were measured in immature specimens, while lowest levels were found in spawning bonitos. The gradual percentage of lipid reduction from immature to spawning bonitos was relatively higher in the liver (females 71·2% and males 64·4%) than in the white (females 59·2% and males 53·5%) and red (females 62·1% and males 51·7%) muscle. Lipid levels in the gonads increased gradually from the immature to spawning stage. The decrease of lipid in the somatic tissues was more intense in females than in males, and gonadal lipid content was higher in females than in males. There was a strong reverse correlation between water and lipid percentage in all tissues. Protein content decreased significantly only in spawning bonitos. The percentage of protein reduction from immature to spawning stage was relatively higher in males than in females in both white (females 3·4% and males 4·6%) and red (females 4·6% and males 5·1%) muscles. Protein content in the liver was significantly lower than in the other tissues, being highest in mature females. Gonadal protein content in females increased with maturation and decreased after spawning. The content in ash exhibited considerable stability. The RNA:DNA ratio exhibited a similar pattern of variation in both muscles. The RNA:DNA ratio increased during gonadal development gradually from the developing to spent stage. It was concluded that in S. sarda during gonadal development, there was an increase in gonadal lipid accompanied by a decrease in somatic tissue lipid reserves. Thus, reproductive inactive bonitos have more lipid in their edible part and a higher nutritional value than active ones.     

Intersexual differences in energy expenditure of Anolis carolinensis lizards during breeding and postbreeding seasons

Although the amount of energy that males and females invest in reproduction is an integral component of theories explaining the evolution of particular mating strategies, few studies have actually determined the amount of energy that each sex allocates to reproduction. We compared how energy is expended by male and female Anolis carolinensis lizards during both the breeding and postbreeding seasons. We used laboratory respirometry to determine resting metabolic rates (RMRs) of inactive, freshly captured lizards and the doubly labeled water technique to determine field metabolic rates (FMRs) of free-ranging lizards. Both RMRs and FMRs were influenced by body mass but not by sex. Season did not influence FMRs; however, RMRs of both sexes increased approximately 40% from the breeding to the postbreeding season. The seasonal increase in RMRs was attributed to a postreproductive increase in feeding rate and specific dynamic action. We used RMRs, FMRs, and thermal profiles of lizards to calculate energy budgets for breeding and postbreeding seasons. Energy budgets partitioned daily field energy (DFE; calculated from FMRs) into daily activity energy (DAE) and daily resting energy (DRE; calculated from RMRs). Energy expended for reproduction was estimated as DAE during the breeding season plus egg production (for females). Despite males having 40% greater body mass, females expended 46% more energy for reproduction than did males (906 and 619 J/d, respectively). Total metabolizable energy (TME=DFE+egg production for females) expended during the breeding season was similar for males and females (1,280 and 1,365 J/d, respectively). Although TME of females decreased 44% from the breeding to the postbreeding season (1,365 vs. 766 J/d), TME of males was similar during both seasons (1,280 vs. 1,245 J/d). There were both seasonal and sexual differences in DRE and DAE. Compared with most lizards from semiarid/desert habitats, A. carolinensis in a temperate habitat expends more total energy during the breeding season, allocates more energy to eggs, and appears to have more total energy available for reproduction.     

Biochemical content, energy composition and reproductive effort in the broadcasting sea star Asterias vulgaris over the spawning period

To assess sex differences in reproductive effort, we examined the biochemical composition and energetic content of the principal body components of the broadcast spawning sea star Asterias vulgaris in the Mingan Islands in the northern Gulf of St. Lawrence, eastern Canada. The body wall was the most stable body component, showing no variations in mass or in lipid and protein content (and total energetic content) between sexes or during spawning. Patterns in the gonads differed between sexes and with spawning. The lipid, protein and carbohydrate content of the ovary dropped during spawning, while only the protein content of the testis decreased significantly. Reproductive effort, expressed as loss of energy in the gonads during spawning for an individual weighing 10 g in underwater mass (8.2 cm in radius), was six times greater in females (49.5 kJ) than males (7.9 kJ). The energetic content of the pyloric caeca also decreased during spawning, by 17.7 kJ in females and 21.5 kJ in males, mainly due to a decrease in lipids. If this decrease is included as reproductive effort, it lessens the gender difference. The caecum decrease possibly represented expenditures due to formation of aggregations or the expulsion of gametes during spawning. Effectively, we observed aggregations during a massive spawning in this population. The sex ratio did not differ from 1:1 in all size classes sampled. This suggests that, unless males suffer higher mortality, females manage to allocate as much energy to somatic growth as males, possibly by feeding at higher rates to compensate for their higher reproductive effort. Stomach protein content tended to be higher in females than males and may indicate greater muscular development to facilitate digestion.     

Changes in structure of tissues and in plasma cortisol during the spawning migration of pink salmon, Oncorhynchus gorbucha (Walbaum)

Changes in the structure of the gonad, skin, interrenal, liver, kidney, stomach, gill and pituitary gland, as well as blood cortisol and haematocrit values were investigated in adult pink salmon during their migration through the Fraser and Thompson Rivers to the spawning grounds. At the commencement of their freshwater migration the gonads of both males and females were in an advanced state of development, the pituitary contained a large complement of well-granulated gonadotrops, and hypertrophy was evident in the interrenal tissue and in the epidermis of the skin. At this time, no change from the normal sexually immature salmon was evident in the structure of the gill, liver or stomach. Sclerosis of the glomeruli was noted in the kidney. The plasma cortisol level was consistent with concentrations in unstressed salmon.
Migration of the fish through a turbulent section of the Fraser River evoked a marked increase in both blood cortisol concentration and in interrenal nuclear diameters.
On arrival at the spawning grounds, 10–15 days after entry into fresh water, a general but not marked deterioration of the tissues was evident. The results are discussed in relation to the spawning migration of other species of Pacific salmon.     

Interannual variation in the reproductive cycle of the New Zealand snapper Pagrus auratus (Bloch & Schneider) (Sparidae)

Changes in reproductive condition in the New Zealand snapper Pagrus auratus (Bloch & Schneider) were monitored in a wild population over three successive years. Recrudescence occurred in spring with spawning beginning in October and continuing for 3 to 5 months. The initiation of spawning varied by up to 3 weeks and was associated with sea surface temperatures of 15–16° C. Theconclusion of spawning was associated with temperatures of 19–21° C but showed greater interannual variation than the onset of spawning. Changes in the gonadosomatic index were accompanied by parallel changes in hepatosomatic index in females but not males, reflecting the role of the liver in vitellogenesis in females.     

Time of start of spawning in Atlantic cod (Gadus morhua) females in relation to vitellogenic oocyte diameter, temperature, fish length and condition

Spawning time of female Atlantic cod ( Gadus morhua L.) can be predicted to within 3 days using vitellogenic oocyte diameters measured over the last month before spawning. Simulations show that a l°C drop in temperature during vitellogenesis delays spawning by about 8–10 days. Similar results were obtained with fish on low ration for 4–8 months before spawning. Moderately fed females held over two to four consecutive spawning seasons demonstrated a significant delay (⋍3–40 days) in the date of first spawning as the fish grew older and more fecund. Forecasting studies on spawning migrating Arcto-Norwegian cod gave no evidence of a significant effect of fish length on the time of spawning. Warmer water temperature during vitellogenesis for the larger fish appears to compensate for the negative effect of fecundity On spawning time.     

Seasonal changes in energy allocation to somatic and reproductive body components of the common cold temperature sea urchin <Emphasis Type="Italic">Loxechinus albus</Emphasis> in a Sub-Antarctic environment

Monthly variations in the somatic indexes and energetic content of organs were investigated in Loxechinus albus from the Beagle Channel. Samples were collected monthly from May 2004 to May 2005. Lantern and test indexes did not vary significantly. A major peak of gonad index (GI) was observed in winter (sexual maturation period), with a strong declination in November suggesting a spawning period in spring. In coincidence a shortening of feeding activities was expressed by the lower values of gut index (Gut I), suggesting that the gut is a storage organ. The values of gonad energetic content (GEC) and total gonad energetic content (TGEC) showed minimum values in winter (ripe stage) and the maximum in spring (spawned stage). The TGEC reached higher monthly average values (50–200 kJ) than total gut energetic content (TGuEC) (20–40 kJ). These differences indicate that the gonads constitute the most important store organs in L. albus. Moreover, organic stores are built up in the gonads after spawning, and then utilized during gamete production.