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.     

Differential utilization of energy during wintering and spawning migration in Norwegian spring-spawning herring

The energy utilization in non-feeding Norwegian spring-spawning herring Clupea harengus was examined during periods of wintering in a fjord system in northern Norway and southward spawning migration (800 km distance) to the main spawning area off western Norway. At the onset of wintering the gonadosomatic index (IG) was higher and the amount of mesenteric fat was lower in male than in female herring, whereas the subsequent increase in IG and decrease in mesenteric fat occurred at a higher rate in females than in males. However, no differences were recorded between sexes in fat content (%) of whole fish at any time of the year. The absolute energy loss increased, whereas the relative energy loss decreased, with fish length. Furthermore, the absolute and relative length specific energy losses were two to three and three to four times higher, respectively, during spawning migration than during wintering. The distinct energy loss was attributed mainly to fat metabolism, whereas the storage protein only was transferred from the soma to the gonads and remained relatively constant in whole herring throughout the wintering and spawning migration. © 1999 The Fisheries Society of the British Isles     

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.     

The biochemical composition and calorie density of the walleye pollock <Emphasis Type="Italic">Theragra chalcogramma</Emphasis> in the Sea of Okhotsk

In tissues of the walleye pollock Theragra chalcogramma the dry matter content averages 18.5%. The lipid content of the raw material is 0.7%, the protein content is 15.3%, carbohydrates are 0.6%, and ash is 1.3%. The average calorie density is 940 cal/g wet weight and 5080 cal/g dry weight. The dry matter content of gonads varies within 14.9–28.0% in females and 14.5–17.0% in males. The lipid content of the raw material is 0.9–3.0% in females and 1.3–1.8% in males; the protein content is 10.2–21.5% and 10.7–13.4%, respectively. The calorie density of female gonads is 702–1537 cal/g wet weight and 4426–5482 cal/g dry weight; for the male gonads it is 760–960 cal/g wet weight and 4952–5641 cal/g dry weight. The dry matter content of the liver varies within 42.2–62.2% for females and 34.4–62.4 for males. The lipid content of the raw material is 25.6–44.5% for females and 16.6–41.3% for males; the protein content is 6.3–9.8% and 8.1–12.3%, respectively. The calorie density of the liver in females varies within 2918–4601 cal/g wet weight and 6370–7395 cal/g dry weight; in males it is 2291–4357 cal/g wet weight and 6392–7492 cal/g dry weight. The minimum calorie density of the liver is observed in juvenile pollock: 963 cal/g wet weight and 2045 cal/g dry weight. The dry-matter content of feces in different size groups varies within 15.0–18.4%. Values of the average lipid content of raw material range from 1.1 to 1.6%; the protein content is from 1.8 to 3.8% and carbohydrates are from 0.9 to 1.4%. The calorie density of feces from variously-sized walleye pollock varies within a narrow range, from 308 to 362 cal/g wet weight. The energy equivalent ranges, depending on body size, from 259 to 2377 cal. The share of energy concentrated in the somatic (muscle) tissue of variously-sized walleye pollock during ontogenesis constitutes 56.5–93.9%; in female gonads it is 0.9–26.6%; in male gonads it is 0.4–7.3%, in the female liver it is 7.9–27.2%, and in the male liver it is 5.7–26.9%. The amount of energy (cal), concentrated in the female liver and gonads is on average 1.5 and 3 times as high as that in the male liver and gonads, respectively. The maximum total energy loss (15–30%) in mature walleye pollock of various-sizes occurs in the spawning period, during the transition from the maturity stage 5 to stage 6. The total amount of energy accumulated during the lifecycle from small juveniles (<17 cm) to very large individuals (>60 cm) averages 1964 kcal for females and 1465 kcal for males. The difference in the amount of energy is explained by the fact that oogenesis requires more energy than spermatogenesis.     

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.     

Sperm and milt characteristics and male v. female gametic investment in the Caribbean reef fish, Thalassoma bifasciatum

Individual sperm cells produced by two male morphs of the bluehead wrasse Thalassoma bifasciatum did not differ in size (i.e. cell volume). Initial phase (IP) males (high sperm competition) had a 60% higher sperm concentration in their milt than did terminal phase (TP) males (low sperm competition), which may reflect differences in how accurately the two male morphs need to allocate sperm to their spawns. The energy density of milt was about 16% lower than that of eggs. Estimates of gametic energy investment based on (a) the difference in testis weights between the beginning and the end of the spawning period and (b) the number of sperm released in natural spawns (determined in other studies), suggested that, on a daily basis, IP males invest about 65% of that of females. Estimates based on stripping milt from IP males at the beginning and the end of the spawning period, however, indicated that their daily energy investment in gamete production is about 10% of that of females Gametic investment by TP males is lower than that by both IP males and females.     

Seasonal variation in fatty acid composition of silver pomfrets,Pampus argenteus (Euphrasen), in Kuwait waters

Proximate and fatty acid composition of wild silver pomfrets, Pampus argenteus, were studied in Kuwait waters for a period of 1 year (November 2007–October 2008) to see whether there were any seasonal compositional differences between males and females. Ten adults (five males, five females) were sampled each month during (i) Pre‐spawning (March–May), (ii) Spawning (June–August), (iii) Post‐spawning (September–November), and (iv) Winter (December–February). Both sexes had significantly (P < 0.05) higher whole body moisture and lower crude protein and lipid contents in winter compared to the respective males and females sampled in other seasons. However, females had significantly higher (9.1%) lipid content during the pre‐spawning season than females in other seasons (7.0–8.2%). The most abundant fatty acid in whole body lipid in both sexes was C16 followed by C18:1n‐9, which accounted for about 31–35% and 22–24% of total lipids, respectively. Males in the pre‐spawning and spawning seasons had significantly higher total monosaturated fatty acids (MUFA) than males and females in post‐spawning and winter. Males had significantly higher total polyunsaturated fatty acids (PUFA) during post‐spawning seasons than females in pre‐spawning and winter seasons. However, there were no significant differences (P > 0.05) in total saturated fatty acids (SFA), PUFA, EPA (eicosapentaenoic acid), DHA (docosahexaenoic acid) or n‐3/n‐6 ratios between respective males and females in different seasons. Livers in males had significantly (P < 0.05) higher MUFA, SFA, PUFA, EPA and DHA than respective females in all months during the spawning season. Female gonads had significantly (P < 0.05) higher MUFA and PUFA but lower SFA content than males in different months during the spawning season. In contrast to the liver, the gonad DHA content and n‐3/n‐6 ratios in females were significantly higher than in males. The gonads from both sexes contained more than double the amount of EPA present in liver; in the case of DHA this was more than three‐fold higher in female gonads, but not in males. Thus, the presence of higher proportions of PUFA, EPA and DHA in gonads, particularly in eggs of silver pomfret, indicates their need for these fatty acids, which may be used as a guideline for dietary essential n‐3 fatty acid requirements for feed formulation of this species. A higher content of DHA in eggs also indicates the higher requirement for DHA in the broodstock diet of silver pomfret.     

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.     

The seasonal changes in somatic energy content of Gulf of Alaska yellowfin sole, Pleuronectes asper

Seasonal measurement of body energy content was made for Pleuronectes asper (Pallas, 1814) from the Gulf of Alaska. Whole body energy content of complete fish (～ 3472 J g^?1 wet wt) was minimal in May for females as their overwintering phase ended, then increased to ～ 4456 J g^?1 prior to spawning. The ovarian index [(g.w./t.w.) × 100] and energy content of ovaries (J g^?1) was highest in June and May respectively, and then declined markedly by August as spawning occurred. Throughout the year male whole body energy content of complete fish ranged from 3351 to 4590 J g^?1 with the lowest values in May and highest values occurring during June to September, the feeding season. The testes index [(g.w./t.w.) × 100] and total energy content of testes (J g^?1) were high in March and lowest during June and July. On a weight-specific basis, males and females had similar whole body energy values throughout the year. Juveniles followed the same seasonal trends in energy storage as adults and had similar whole body energy values. Whole body energy content was linearly related to wet and dry weight condition factor with r² values of 0.70 and 0.87, respectively. Dry body weight as percent of wet body weight was the best predictor of body energy (r²=0.91). Yellowfin sole had an annual energy cycle with energy accumulation and growth from May to September. Thereafter they utilized stored energy for metabolic and reproductive needs. Spawning began in late May or early June and fish were spent by August. Whole body energy content increased by 28, 33 and 35% between May and June, for females, juveniles, and males, respectively, the most dramatic change during the year long survey. This suggests that intense feeding in May must be an important aspect of their energy storage cycle.     

Reproductive cycle and reproductive output of the sea urchin Loxechinus albus (Echinodermata: Echinoidea) from Beagle Channel, Tierra del Fuego, Argentina

Reproductive cycle, frequency and duration of spawning, energetic content of gonads, and reproductive output of the common green sea urchin Loxechinus albus were analyzed in the Beagle Channel (Tierra del Fuego) between May 2004 and May 2005. Gonad indices (GI, percentages of gonad mass in total body mass) were significantly higher in March, April, July, and August than in November and May, thus showing a negative correlation with the photoperiod. Highest GI values of mature individuals were observed in August, and spawning occurred from September to December. In females, the mass-specific energy content of gonads (ECG) was highest in spawned gonads and lowest in mature ones, while in males ECG values were higher in immature stage and lower in premature and mature stages. High ECG values can be explained by the abundance of nutritive phagocytes. Both ECG and total gonad energy content (TECG) were higher in females than in males. Mean reproductive output was 7.28% for females and 6.15% for males (expressed as the difference between mean GI of mature and spawned gonads) and 25.02 kJ for females and 19.26 kJ for males (expressed as the difference between mean TECG of mature and spawned gonads).     

Sex differences in brown adipose tissue thermogenic features during caloric restriction.

Caloric restriction (CR) studies have shown that females rats conserve energy more efficiently, showing a higher resistance to weight loss and higher protection of vital organs mass than male rats. Gender-dependent inactivation of thermogenesis in brown adipose tissue (BAT) has been proposed as one of these possible energy conserving mechanisms. To study the changes underlying this inactivation in rats, a three month study with 40% CR was undertaken to unravel the effects on BAT recruitment. Under ad libitum conditions female rats had greater BAT recruitment and greater oxygen consumption than their male counterparts. Total and mitochondrial protein, as well as triglyceride and DNA content were more reduced in restricted female rats than in restricted males. Similarly, the levels of key BAT functional proteins (UCP1, LPL, HSL, TFAM) were more reduced in restricted females, whereas no changes were found in mitochondrial DNA levels (mtDNA) and OXPHOS activities in males and females. Furthermore, alpha (2A)/beta (3) adrenergic receptor ratio remained constant in male rats whereas in female rats CR increased 60%. In conclusion, our results suggest that female rats, whose BAT thermogenic activity is higher in ad libitum conditions, is depressed during CR. This inactivation involves the mitochondrial differentiation process and lipolytic system and could be due, at least in part, to the unfavourable adrenergic receptor balance for thermogenic activation.     

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 .     

Plasma corticosterone level of rats after bilateral lesion of the globus pallidus and elimination of the sex-dependence of the body weight loss following pallidal lesion by bilateral adrenalectomy

Plasma corticosterone (CST) level and body weight were studied in male and female rats after bilateral pallidal lesion (GPL). The effect of high dose of glycocorticoids on body weight of food and water deprived, non-lesioned rats was also studied since high CST levels due to the stress caused by the aphagia and adipsia could be anticipated. Weight changes in lesioned-adrenalectomized animals were in investigated as well. A weight loss, higher in males than females, developed after the pallidal lesion and the CST level was higher in lesioned females than in lesioned males. This sex-difference was present in non-lesioned, food and water deprived animals as well. The weight loss was higher in lesioned males than in food and water deprived male/female controls. Adrenalectomy eliminated weight loss differences between the sexes after lesion, by reducing the weight loss in males. CST administration reduced body weight loss in food and water deprived males but not in familes, and cortisol facilitated it in both sexes.     

Reproductive success in male Japanese minnows,Pseudorasbora parva: observations under experimental conditions

To evaluate the spawning success of male Japanese minnows,Pseudorasbora parva, and female mate choice, spawning behaviour was observed under both artificial and experimental conditions. Larger males had larger territories and greater reproductive success. The body weight of territorial males decreased during the maintenance of territories, while that of non-territorial males increased significantly. When the weight of non-territorial males exceeded that of territorial males, the former began to establish new territories on the substrate, suggesting a conditional strategy by non-territorial males to trade off immediate reproductive success with growth and hence improve future reproductive success. Females chose males with larger body size, probably based on dominance rank rather than the quality (or size) of territory. It was concluded that females choose males of higher dominance rank and that males compete for large territories, both of which play an important part in male reproductive success.     

Epizootic cutaneous papillomatosis, cortisol and male ornamentation during and after breeding in the roach Rutilus rutilus

The prevalence of epidermal papillomatosis in roach is known to peak during the spawning period and to be higher in males than in females. The high occurrence of papillomatosis in polluted waters suggests that stress may contribute to the outbreak of the disease. However, little is known about breeding-induced stress in fish and its relationship with diseases. In this study, plasma cortisol concentration, hematocrit and the relative size of the spleen were determined in healthy and diseased male and female roach Rutilus rutilus during and shortly after spawning in a wild population. In addition, the sexual ornamentation (breeding tubercles on the lateral sides and on the frontal) of male roach during spawning was examined. Plasma cortisol concentration was higher during than after the spawning period, and higher in males than in females during spawning, indicating a spawning-induced stress and higher spawning stress among males. There was no correlation between cortisol concentration and the intensity of papillomatosis (number of scales under papilloma tumors) among the diseased fish. However, the significant interaction sex x disease status revealed by ANCOVA suggested that diseased males could be more prone to increased cortisol levels than diseased females or healthy males. Hematocrit values (ratio of the volume of red blood cells to total volume of blood) but not condition factor were lowered in papilloma-diseased fish after spawning. The relative size of the spleen was greater in males than in females. The number of frontal breeding tubercles correlated negatively with the intensity of papillomatosis. Experimental studies are needed to investigate the association of papillomatosis with stress and cortisol.     

Synchronous female spawning and male mating behavior in a land-locked population of Japanese charr, Salvelinus leucomaenis japonicus

In resource-based promiscuous mating systems, synchronous spawning of females affects competition among males and variation in the reproductive success of males. We documented the mating behavior of Japanese charr (Salvelinus leucomaenis japonicus) through an annual breeding season to examine the relationship between female spawning synchrony and male mating behavior. Females spawned highly synchronously in the population studied, i.e., approximately half the spawning was finished within the first three days of the entire spawning season (11 days). The daily operational sex ratio (OSR) was nearly 1:1 through the spawning period. The number of males around a spawning female was very small (1.21+/-0.49 males per female) over the spawning ground and period, suggesting that a competitive male could effectively chase subordinate males away from a spawning female. A few males attempted to sneak near the oviposition site of females (16%; 9 of 57 breeding groups), while some males adopted sneaking tactics in the initial phase of females' spawning (24%). We did not observe any males to succeed in sneak fertilizations. We conclude that in this Japanese charr population, the synchronous spawning of females was related to the unbiased daily OSR, male aggregation around females, and consequently whether and how efficiently males engaged in sneak mating behavior.     

Female mate choice and spawning behaviour of Chinook salmon under experimental conditions

After spawning their first nest, female chinock salmon Oncorhynchus tshawytscha paired with small males ( c. 46% of female weight) spent an average of 16·2 h between spawning of successive nests compared with 9·6 h for females paired with large males ( c. 112% of female weight) ( P< 0·05). Neither frequencies of female nest construction behaviours (digging and probing) nor male courtship behaviour (crossovers and quivers) differed between large- and small-male pairs. Male quivering frequencies were correlated significantly with female digging and probing frequencies, whereas the crossover frequencies were not. It is suggested that delayed spawning by females in the presence of relatively small males is a primary mechanism by which females in the genus Oncorhynchus exhibit mate choice.     

Plasma non-esterified fatty acid profiles in wild Atlantic salmon during their freshwater migration and spawning

Total plasma non-esterified fatty acid (NEFA) levels increased significantly in adult Atlantic salmon during the first months of their upstream migration and spawning in the Exploits River, Newfoundland, Canada. The highest levels occurred in May and were 5467±270·43 nmol ml⁻¹ for females and 4617±334·70 nmol ml⁻¹ for males. Significantly higher levels were maintained by females compared with males for most of the upstream migration. Between August and October, total plasma NEFA levels declined by 61% in females but only 23% in males. The decline in plasma monounsaturated and polyunsaturated fatty acid levels accounted for 74% of the loss of NEFAs in females. Specific plasma NEFAs such as 16: 0 (palmitic), 16: 1 (palmitoleic), 18: 1n9 (oleic) and 20: 5n3 (eicosapentaenoic acid) differed significantly between males and females during migration and spawning. The mean gonadosomatic index ( I _G) values of females in May and just prior to spawning were 0·37±0·01 and 10·25±0·32, respectively. The rapid decline in the plasma NEFA content of females coincided with the largest increase in their I _G (1·85±0·02–10·25±0·32). Corresponding I _G values for males were 0·34±0·01 in May and 3·33±0·78 prior to spawning. Plasma NEFA levels of spent salmon did not differ between sexes and were significantly lower than those of salmon preparing to spawn.     

Steroid hormone profiles and correlative gonadal histological changes during natural sex reversal of sobaity kept in tanks and sea-cages

The gonadosomatic index of sobaity Sparidentex hasta fits well with the natural spawning season of February-March. At 12 and 24 months all the population matured as males. However, during this time, between April-September intersex gonads were seen. From October of the third year of life, females were seen for the first time. These changes in the histology of the gonads were correlated with the hormonal profile. Testosterone showed no significant correlation with sex but 11-ketotestosterone correlated very well with the mature male gonads and thus can be used for sexing. Oestradiol-17β higher in sex reversed individuals (females) in the spawning season than in males. The sequence of events in the protandrous sex change of this fish during the first two spawning seasons is male → intersex → male. However, during the third spawning season (age 36–37 months) this sequence is male →intersex → male or female (10–17%).     

Behavioral Tactics of Male Sockeye Salmon (Oncorhynchus nerka) under Varying Operational Sex Ratios

Previous studies have demonstrated several reproductive-behavior patterns in male salmon, including competitive and sneaking tactics, the formation of hierarchies, and non-hierarchical aggregations around ripe females. Through behavioral observations at varying spatial and temporal scales, we examined the hypothesis that operational sex ratio (OSR) determines male sockeye salmon (Oncorhynchus nerka) distribution and breeding tactics. Patterns of male distribution and behavior varied over both coarse and fine scales, associated with apparent shifts in reproductive opportunities, the physical characteristics of the breeding sites, and the deterioration of the fish as they approached death. Females spawned completely within a few days of arriving on the spawning grounds, whereas males courted the available ripe females from the date of their arrival on the spawning ground until their death. This difference in reproductive lifespans tended to elevate late-season OSRs but was partially counterbalanced by male departures and the arrival of other ripe females. The proportion of males able to dominate access to ripe females decreased and the number of large courting groups increased over the course of the season, apparently related to both increasing OSR and the deteriorating physical condition of males. However, great variation in OSR was observed within the spawning sites on a given day. OSRs were generally higher in shallow than in deep water, perhaps because larger females or more desirable breeding sites were concentrated in shallow water. The aggregations of males courting females were not stable (i.e. many arrivals and departures took place) and male aggression varied with group size. Aggression was most frequent at low OSRs and in groups of intermediate size (2–4 males per female), and much less frequent in larger groups, consistent with the needs of maximizing reproductive opportunities while minimizing unproductive energy expenditure. These results indicate that, while OSR strongly influences male distribution and behavior, other factors such as physical condition, limited movement and habitat choice may also affect male reproductive opportunities.