Patterns of oocyte growth, vitellogenin and gonadal steroid concentrations in greenback flounder

The pattern of oocyte development in association with changes of plasma concentrations of vitellogenin (Vtg), 17β‐oestradiol (E₂) and testosterone (T) was investigated in maturing female greenback flounder Rhombosolea tapirina over the first part of a reproductive season (February to June). Examination of oocyte size‐frequency distributions showed that the oocyte developmental pattern in R. tapirina is multiple group synchrony, and that reproductively mature fish were present at all sampling times. There were no significant temporal variations in the gonado‐somatic index ( I _G), hepato‐somatic index ( I _H), or plasma concentrations of Vtg, E₂ and T during the sampling period, which indicates that reproductive development is not synchronized within the population. Significant increases in I _G, I _H and plasma concentrations of Vtg, E₂ and T, however, were observed in vitellogenic fish, and in fish undergoing final maturation. A positive relationship was also found between the growth of oocytes and plasma concentrations of Vtg, E₂ and T, although the patterns of increase were different for each variable. Plasma concentrations of Vtg and E₂ rose steadily across oocyte sizes from 100 to 450 μm, but the rate of increase of plasma E₂ was slower than that of Vtg, and both reached a saturated concentration at oocyte sizes of c . 450 μm. In contrast, plasma concentrations of T showed no marked increase until oocytes grew beyond 400 μm.     

Latitudinal variability in lunar spawning rhythms: absence of a lunar pattern in the northern mummichog Fundulus heteroclitus macrolepidotus

In this study, the natural spawning season of the northern mummichog Fundulus heteroclitus macrolepidotus was examined from May to July in the lower St John River, New Brunswick, Canada (45° N), through measurements of gonado-somatic index ( I _G), liver-somatic index ( I _L) and condition factor ( K ). I _G increased during the prespawning phase (mean ± s.e. 5·49 ± 0·20%), peaked during the spawning phase (14·68 ± 0·51%) and dropped during the regression phase (3·23 ± 0·29%). A single peak of spawning was observed in F. h. macrolepidotus , contrasting with multiple peaks observed in studies south of 41° N in North America. Within the prespawning phase, there were significant differences between sampling dates in female body mass, gonad mass, liver mass and K . On the basis of results from this study, water temperature is believed to be the predominant factor controlling initiation of spawning in F. h. macrolepidotus .     

Reproductive cycle and sexual maturity of the anglerfish Lophiomus setigerus in the East China Sea with a note on specialized spermatogenesis

The reproductive cycle and sexual maturity of the anglerfish Lophiomus setigerus were examined. Spermatids were released from the germinal cysts into the lumina of the seminal lobules, and both spermatids and spermatozoa were present in the lumina of the seminal lobules and sperm ducts. Spermatogenesis and vitellogenesis occurred throughout most of the year. The testes of males were full of spermatozoa throughout the year, with spawning from May to November. Males and females reached sexual maturity at a mean total length and age of 178 mm, 3.3 years and 303 mm, 6.1 years, respectively. There were clear seasonal cycles in the gonadosomatic index (I_G) and hepatosomatic index ( I _H) in females. The mean I_G of females increased rapidly with ovarian development while the mean I _H decreased from the middle of vitellogenesis to final maturation. Mean values of I _G and I _H in males increased with testicular development.     

Effects of fasting on temporal changes in plasma concentrations of sex steroids, growth hormone and insulin‐like growth factor I, and reproductive investment in Arctic charr

Proportions of maturing fish and reproductive output [egg size, relative fecundity, spermatocrit and gonado‐somatic index ( I _G)] were studied in repeat‐spawning (+4 year old) male and female Arctic charr Salvelinus alpinus , subjected to periods of fasting. Groups of individually tagged Arctic charr were fasted for c . 3, 6, 7 and 9 months, from November 1998. In the period February to November 1999, size (fork length and mass), specific growth rate ( G ), condition factor ( K ) and plasma concentrations of growth hormone (GH), insulin‐like growth factor I (IGF‐I), oestradiol‐17β(E₂; females) and 11‐ketotestosterone (11KT; males), were monitored monthly. Maturing fish in each group started to gain mass soon after food was made available, and both sexes reached the highest K and G c . 2 months after the onset of feeding. The fasting regimes resulted in different growth patterns during spring and summer when energy stores are normally replenished in Arctic charr, and K prior to the breeding season was significantly higher in the groups fasted for 3 and 6 months compared to the groups fasted for 7 and 9 months. There were significant positive correlations between K during the period prior to the breeding season and reproductive output in terms of the I _G, spermatocrit and relative fecundity. There was, however, no clear relationship between the length of starvation and the proportion of maturing fish. Likewise, no clear relationships were found between reproductive development and plasma levels of GH and IGF‐I, although both showed marked seasonal changes, being 'down‐regulated' during winter months and 'up‐regulated' throughout summer months.     

Effects of fish size on spawning time in Norwegian spring-spawning herring

During October-January in a northern Norwegian fjord system and in February-March at the main western Norway spawning grounds, an index I _G of gonad weight of spring-spawning herring Clupea harengus in percentage of expected gonad weight at full maturity for a given total length L _T, tended to increase with L _T in the range 27–31 cm as the proportion of recruit spawners decreased. Insignificant differences in I _G were found between L _T groups in the range 32–37 cm (repeat spawners). This is contrary to that suggested in other studies and signifies that generally the stock spawns in only two waves, repeat spawners first and the recruit spawners second. I _G also increased with somatic and total condition factor, which signifies that spawning time may be influenced by summer feeding conditions.     

The reproductive biology and ecology of the Port Jackson shark Heterodontus portusjacksoni in the coastal waters of eastern Australia

The reproductive biology and ecology of the Port Jackson shark Heterodontus portusjacksoni was investigated at three locations on the central and southern coast of New South Wales (NSW), Australia from January 2002 to December 2005 using underwater visual census surveys and samples obtained from a commercial fishery. Adults displayed sexual dimorphism in total length ( L _T) at sexual maturity, with males maturing between 762 and 772 mm L _T and females between 902 and 905 mm L _T. The mean ovarian fecundity was estimated at 16 offspring per female but was unrelated to female L _T. Male gonado-somatic ( I _G) and hepato-somatic ( I _H) indices and female I _G declined from July to November as did maximum ovarian follicle diameter and the diameter of the three largest follicles. Adults were absent from inshore reefs between December and July. Hence, H. portusjacksoni has a synchronous annual breeding season in NSW, which occurs between July and November (the austral winter to spring), with a peak in oviposition from August to October. Heterodontus portusjacksoni copulatory and ovipository behaviour are reported for the first time. Copulation was observed and involved oral grasping of the female's pectoral fin by a single male, which wrapped his body around hers to insert one clasper. Ovipositing females appeared to search crevices in the reef prior to delivering a single capsule, which was washed into the crevice by water movement, with the female departing very soon after oviposition. This study represents the first rigorously quantitative analysis of H. portusjacksoni reproductive biology and ecology in NSW waters.     

Maturation of walleye by age, size and surplus energy

The probability of annual sexual maturation by male and female walleye Stizostedion vitreum was related to age, size and an index of condition, I _VF=[arcsine(visceral fat)^0·5(body mass)^−0·5]. Most males first matured at ages 2 and 3 years; size explained first maturation, but condition explained later maturation. In contrast, most females first matured at ages 4 and 5 years; maturity of females was more dependent upon condition. Maturity of females at ages 4 and 5 years was significantly correlated with average I _VF of the population ( PI _VF). The size reached by age 2 years (early growth) was correlated with the PI _VF. Growing degree-days, Secchi depth, latitude and lake morphology were not correlated with the PI _VF. Annual variations in female spawning stock size were related to the condition of the females, presumably reflecting the net acquisition of energy in the preceding growing season. Annual variations within lakes in the net acquisition of energy may exceed the variations in energy availability between lakes, dictated by lake morphology and geography. Thus, assessment of condition could be used to predict annual potential spawning stock size and egg production.     

Histological classification of oocyte growth and the dynamics of ovarian recrudescence in Tilapia zillii

Serum levels of 17β-oestradiol and testosterone peaked and fell within 6 days of spawning in Tilapia zillii suggesting that vitellogenic growth began as early as day 2 or 3 postspawning. As early as day 8, stage 6/7 (late nitellogenic and maturing) oocytes occupied 60–70% of the ovary. From day 8 onwards the proportion of stage 6/7 oocytes changed little, though I _G increased (as oocytes grew) to reach maximal levels of ∼ 5·5% by day 14. I _G was correlated significantly to the proportion of stage 6/7 oocytes. Postovulatory follicles were observed immediately following spawning occupying up to ∼ 7% of the ovary but were not present from day 3 onwards. Atretic oocytes were found throughout the time period monitored (generally occupying <2% of the ovary) but were more prevalent from day 18 onwards. Data suggest that previtellogenic oocytes are recruited into vitellogenic growth immediately after spawning and can complete vitellogenesis as early as day 8 postspawning. Knowledge of this timing is likely to be important in the development of spawning induction programmes in T. zillii and other related species.     

Effect of temperature changes on the reproductive cycle of roach in Lake Geneva from 1983 to 2001

In Lake Geneva, the surface water temperature has increased by 1° C over 20 years probably as a result of climate change. The effects of changes in temperature on the reproductive cycle of the roach Rutilus rutilus were assessed in a 19 year survey. Over time, spawning tended to begin earlier. The consequences of temperature changes were assessed on two different stages of the female reproductive cycle: the development of the ovaries from the beginning of autumn to ovulation, and the onset of the spawning period. The development of the ovaries was studied for 7 consecutive years from October to June. From 1 October to the onset of spawning, it was possible to assess the gonado-somatic index ( I _G) of females in terms of time expressed as a sum of degree-days. The correlation between I _G and the sum of degree-days was +0·97. The onset of the roach spawning period in Lake Geneva was triggered by a thermal threshold (median and range 190 ± 10 degree-days for the 15 previous days). From October to April, climate warming accelerated the development of gonads, then in May, a thermal threshold that triggered the onset of roach spawning occurred earlier.     

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.     

Sex differences in pectoral muscles but not in pectoral fins in the three‐spined stickleback Gasterosteus aculeatus

The pectoral muscle index ( I _PM)( I _PM = 100 M _PM M⁻¹, where M _PM and M are the pectoral muscle and body masses, respectively) fin‐area and fin ray length were studied over a year in male and female three‐spined stickleback Gasterosteus aculeatus from a marine population (Öresund, Sweden) kept under simulated natural light and temperature conditions. A castration‐replacement experiment was used to test androgen effects on the I _PM, fin‐area and fin ray length. Non‐breeding males were castrated or sham‐operated in winter ( i.e . the fish had low levels of androgens). Castrated control and sham‐operated fish were implanted with empty Silastic capsules and castrated groups with capsules containing the androgens testosterone or 11‐ketoandrostenedione into the abdominal cavity. The experiment was terminated after 41 days, when the controls had matured. No morphological differences were found in pectoral fins between sexes during the year, except during the peak breeding season (May), where females showed larger fin‐area and longer fin ray in length compared to males. No effects of androgens treatment or of castration on pectoral fin‐area or fin ray length was observed. Breeding and non‐breeding males showed higher I _PM compared to females. The lower I _PM in females than in males could not be explained by the larger gonads in the former alone, as a sex difference in I _PM was still present after deduction of the ovaries from the female body mass. The I _PM was higher in sham‐operated compared to castrated fish. No effects of androgens treatment on I _PM was observed.     

Variations in the spawning periodicity of eight fish species in three English lowland rivers over a 6 year period, inferred from 0+ year fish length distributions

The spawning periodicity of eight fish species was investigated in three English lowland rivers over a 6 year period from patterns in 0+ year fish standard length ( L _S) distributions. A single cohort of 0+ year dace Leuciscus leuciscus , roach Rutilus rutilus and perch Perca fluviatilis was observed each year, suggesting that these species spawned only once annually. By contrast, populations of chub Leuciscus cephalus , bleak Alburnus alburnus , bream Abramis brama , gudgeon Gobio gobio and minnow Phoxinus phoxinus were inferred to spawn on more than one occasion each year. Annual and intercatchment variations occurred in the L _S distribution patterns of some of the fish species. In chub, for example, although a minimum of two 0+ year cohorts occurred in all years in the River Trent, 'multiple' spawning (either at the individual or population level) was most apparent in 1999, 2003 and 2004. By contrast, 'multiple' spawning events were not evident in all years in the Warwickshire Avon and Yorkshire Ouse, with recruitment presumably based upon a single spawning event in some years. There is effectively a trade-off between early spawning (extended growing season), and the possibility that environmental conditions will impact upon recruitment success, and the potential for reduced overwinter survival of smaller individuals with lower lipid resources from later spawning events. Notwithstanding, fishes as small as 15 mm L _S survived the winter in some years, suggesting that progeny from later spawning events may make important contributions to fish recruitment success.     

Lipid content and fatty acid composition of target tissues in wild Perca fluviatilis females in relation to hepatic status and gonad maturation

Variation in liver ultra‐structure and composition in relation to energy mobilization was investigated in female perch Perca fluviatilis from the Meuse River between August 2001 and June 2002. In April, just before spawning, the lipo‐somatic index ( I _F) was 0·3%, the gonado‐somatic index ( I _G) was 28% and the total lipid content of the liver was 2·53%. The average areas of lipid droplets and mitochondria were 0·05 and 0·06 μm², respectively. Glycogen supply reached 7·9% of the total area of the hepatocyte. During the sexual resting period, females accumulated energy in perivisceral fat and in the liver to reach 1·6% I _F and 4·85% of liver lipid content in August with lipid droplets average size of 0·09 μm² and glycogen average area of 15%. Liver cells contained a weakly developed rough endoplasmic reticulum (RER) and a great number of small mitochondria (average size 0·02 μm²). The I _G was 0·6% at this time. During the whole annual cycle, the average lipid content of female liver never exceeded 3·9 ± 1·9%. The concentration of docosahexaenoic (DHA), linolenic and linoleic acids increased in mature gonads while linolenic and linoleic acids decreased in the liver during the same period. Fatty acid composition of muscles of perch was characterized by a high content of DHA.     

Onset of maturity and cohort composition at spawning of Baltic sprat Sprattus sprattus on the basis of otolith macrostructure analysis

Data on catch per unit effort (CPUE) from the Danish commercial fishery showed dense aggregations of sprat Sprattus sprattus born in 1997 in the main spawning area of the Baltic Sea during May 1999. After the analysis of their otolith macrostructure, these fish were found to have a shorter distance to the beginning of the first winter zone compared to fish from the same cohort caught in other months. At the same time, the number of fish with large otoliths at the period of winter zone formation tended to decrease or even disappear. The otolith size at winter zone formation ( O _W) was found to be a good predictor of sprat total length ( L _T) and mass ( M ) in age group 0 years. Changes in the otolith size distribution over time were related to variations in cohort composition depending on the L _T and M attained during the juvenile stage. Examination of the gonads suggested two life history patterns: 1) individuals achieving a larger L _T and M during the first growing season may contribute at age 1 years to the spawning stock and the fishery, whereas 2) individuals attaining smaller L _T and M contribute in larger amounts to both the spawning stock and the fishery later, when they are at age 2 years. Moreover, changes in the relationship between O _W and L _T and M were related to differences in growth trajectories for sprat born during the same spawning season. Smaller sprat as group 0 years continued growing during the second growing season delaying maturity compared to larger individuals from the same cohort.     

Intra-annual variation in the batch fecundity of sardine off Portugal

Atresia in the ovaries of sardine Sardina pilchardus did not show clear seasonal patterns. There was high prevalance ( c. 60%) and low intensity ( c. 1% of gonadal area occupied by oocytes) in all months of the spawning season studied. The batch size to ovary-free mass relationship was significantly different in the mid-spawning season, with higher relative fecundity and gonadosomatic index (I_G) in January than in November and March. There was no additional effect on batch fecundity due to age or condition factor. Data from monthly market samples during 1998–1999 revealed a continuous decline of the condition factor throughout the spawning season. The market samples also revealed that the duration of the spawning season is not equal for all fish. In any month, pre-spawning fish tended to have significantly higher I_G than spawners, indicating that delayed initiation of spawning can lead to larger relative fecundity. The increase in sardine relative fecundity during the first half of the spawning season is probably due to the continuous influx of new spawners and possibly to body mass loss that is initially not translated into gonad mass loss. After January when all visible fat reserves within the body cavity have disappeared, the influx of new spawners practically ceases.     

Reproductive behaviour of a temperate serranid fish, Paralabrax clathratus(Girard), from Santa Catalina Island, California, U.S.A.

The reproductive behaviour of the kelp bass Paralabrax clathratus was studied on Santa Catalina Island, California, U.S.A. from April 2000 to September 2002. Adults formed aggregations of three to > 200 individuals, and spawning occurred within subgroups of three to 23 individuals that contained a single female. The gonado‐somatic index ( I _G) of collected ripe males (mean = 5·8%, range = 0·5–13·1%) indicated a large investment in sperm production that is common in group‐spawning fishes characterized by intense sperm competition. Spawning occurred 32 min before sunset to 120 min after sunset, and both males and females were capable of spawning multiple times during a single evening. Behavioural observations of adults and estimates of spawning periodicity from the collection of females with hydrated oocytes suggested that spawning occurred continuously throughout the summer months and showed no significant relationship with the lunar cycle. In general, the spawning behaviour of kelp bass was similar to other functionally gonochoric, group‐spawning serranids. The dynamics of P. clathratus spawning aggregations, however, were inconsistent with that of tropical reef fish spawning aggregations, including the transient spawning aggregations of some tropical serranids. Aggregation spawning appeared to be an important component of the annual reproduction of this species.     

Changes in ovarian development and plasma levels of sex steroid hormones in the wild female Japanese sardine (Sardinops melanostictus) during the spawning period

In order to clarify the spawning pattern of Japaneses sardine ( Sardinops melanostictus ) during the spawning period, changes in ovarian histology, frequency distributions of oocyte diameter and plasma levels of oestradiol-17β (E₂) and 17 a ,20β-dihydroxy-4-pregnen-3-one (17 a ,20β-diOH-P) were examined in female fish captured in the region off Kyushu and Shikoku in Japan. With the development of the ovary, the gonadosomatic index (GSI) increased gradually and the size of oocytes became larger. When the GSI exceeded 13, the translucent hydrated eggs began to dominate and ovulation subsequently occurred. After ovulation, GSI decreased to less than 3. Post-ovulatory follicles were found only in the lower GSI ranges and atretic oocytes were always observed in the ovary. Plasma E₂ levels increased along with ovarian development but decreased after completion of yolk accumulation. On the other hand, 17 a ,20β-diOH-P showed the highest level just before and after ovulation. Ovarian histology and hormonal profiles suggest that the sardine spawns repeatedly during the spawning period. Based on the concentration of 17a,20β-diOH-P, the female Japanese sardine was estimated to ovulate before the middle of the night.     

Lunar spawning in Siganus canaliculatus

A major spawning of the seagrass rabbitfish Siganus canaliculatus occurred 4 days after the new moon in both May and June 1993, and 7 days after the new moon in 1994. The gonadosomatic index ( I _G) and serum vitellogenin (VTG) levels fluctuated according to the lunar cycle; IG and VTG levels showed peaks at around the new moon and the waning moon, respectively, suggesting that spawning of this species is synchronized with the lunar cycle. Vitellogenic oocytes appeared on day 2 after the first spawning and were fully mature on day 30. When a greater percentage of the most advanced oocytes attained the tertiary yolk stage, they formed a batch and separated from the adjacent group of smaller pre-vitellogenic oocytes, indicating that S. canaliculatus is a multiple spawner with an ovary belonging to the group-synchronous type of oocyte development. Batch fecundity, assessed using batches of oocytes at and after the tertiary yolk stage, ranged from c. 0·52 to 2·56 million eggs. The relationship between batch fecundity (F) and fork length (1) can be represented as F=0·0536854L^5·07292.     

Seasonal growth of Atlantic cod: effects of temperature, feeding and reproduction

Growth of 2659 Atlantic cod Gadus morhua aged 4 to 9 years examined in Placentia Bay, Newfoundland, peaked in most cases in June and was at a minimum in October or November. Water temperature, partial fullness index ( I _P) and gonado‐somatic index ( I _G) explained between 31 and 52% of the monthly variability in growth. Temperature and I _P of capelin Mallotus villosus had significant effects on growth of all age groups and explained most of the variance for ages 6–8 and 4–5 years, respectively. The I _P of large invertebrates (ages 4 to 7 years), sandlance ( Ammodytes sp. age 6 years) and demersal fishes (age 9 years) had age‐specific effects in the model. Overall, amphipods, decapods and echinoderms dominated the Atlantic cod diet in most seasons, but fish consumption by Atlantic cod was high in June and July, particularly on capelin. The rapid increase in somatic mass during June and July occurred despite cold water temperatures ( < 3° C at 50 m) and moderate to high gonado‐somatic index. The findings of this study suggest that when food was not a limiting factor, growth tended to increase even when Atlantic cod occupied colder waters, but when food was limiting, the opposite may have occured.     

Ovarian morphology of the Japanese eel in Mikawa Bay

Annual changes in gonadal maturation of female Japanese eel Anguilla japonica in sea water were investigated histologically over 5 years in the Mikawa Bay, Japan, where they occurred throughout the year except in March. Almost all immature Japanese eels (yellow eels) occurred mainly from April to September, and they were rare after November. In contrast, maturing Japanese eels (silver eels) occurred from October to February. The gonado‐somatic index ( I _G) and oocyte diameters of yellow eels were <1·0 and 150 μm, respectively, and oocytes were at the peri‐nucleolus or the oil droplet stages. The I _G and oocyte diameters of silver eels were greater than those of yellow eels and most oocytes developed to the primary yolk globule stage. The numbers of silver eels lacking oocytes at the primary yolk globule stage increased after January in Mikawa Bay, although I _G and oocyte diameters remained unchanged. In contrast, silver eels caught at the mouth of the bay in January possessed oocytes that had advanced to the secondary yolk globule stage. These observations indicate that oocyte development changes seasonally, especially after winter in Mikawa Bay.