Maturation and Reproductive Biology of Female Wild Carp, Cyprinus carpio, in Victoria, Australia

The reproduction of carp Cyprinus carpio L. (Family: Cyprinidae), from Victorian waters in Australia, is studied with detailed analysis of gonad maturation, spawning season, fecundity and oocyte diameter. Results show that carp has a high annual fecundity (AF) (0.12–1.54 million oocytes per fish), which is positively correlated with caudal fork-length (L, mm) and total weight (W, kg) but not age. The relationships between length or weight and AF were statistically significant and best described with the simple linear or quadratic regressions. Mean relative fecundity was 0.163 million eggs kg^–1 whole weight. Egg size was estimated from oocyte diameter in carp from eight stocks. Egg size was proportional to maternal size but not age. Seasonal trends in gonadosomatic indices, together with the changes in the macroscopic and microscopic condition of ovaries, demonstrated that spawning generally peaks during spring–early summer, but also occurs through until autumn and can even start in late winter at some sites. In Victoria, this species is a multiple spawner with asynchronous oocyte development and a protracted spawning season. Stocks generally contain both females that spawn once, and females that spawn repeatedly, within a spawning season. Implications for management for the control of feral carp stocks are discussed.     

The relationship of gonad and egg size to weight and age in the European and Chinese races of the common carp Cyprinus carpio L.

Gonad development was studied in young carp of the European and Chinese (big-belly) races. The effects of age and body weight on gonad and egg size were partitioned and the following findings were made: (i) Gonad weight was correlated, in both males and females, with body weight, but proportional gonad weight, i.e. the ratio gonad weight divided by body weight, was independent of body weight, (ii) Proportional gonad weight had reached its maximum value in one year old males, but it more than doubled in two year old females, (iii) Egg size of two year old females was more than twice the size of eggs of one year old females, (iv) The European and Chinese races of carp differed in several aspects of gonad development. In particular, the Chinese matured earlier and their relative gonad sizes were considerably larger, (v) Genetic variation in gonad development was also found within the European race, where faster rate of development is dominant over slower rate. The evolutionary implications of the above differences were discussed.     

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.     

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).     

The effects of population density on fecundity and fertility in the guppy, Poecilia reticulata (Peters)

Experiments were performed to examine the influence of population density on the fecundity and fertility in the guppy, Poecilia reticulata . The frequency of females with only ova or with different combinations of reproductive units and developmental stages in the ovary was significantly different between different densities but not between females with one and two gestations from the same population density. The ovarian width of females with one gestation from the lowest density was significantly larger than in the highest population density. Gonad length and height and ovarian volume of females with two gestation periods were larger in the lowest compared with the highest density. After one gestation, females at the lowest population level had greater ovarian length, height, gonad weight, width and volume compared with those from the higher density. Ovarian length, height and volume and gonad weight were also significantly greater among females of two gestation periods in the lowest than in the second lowest density. The average gonadosomatic ratio was highest in the lowest population density. A significant correlation was found between gonad volume and total body weight and between gonad volume and ovarian weight of females at every population level. This was also the case when ovarian weight vs. total body weight was examined in each population density. Only at the lowest population level was there a significant correlation between fecundity and total body length and between fertility and body length. A significant relationship was also found between gonad volume and total body length and between gonad weight and body length at the same population level. No significant difference of fecundity and ovum size was found between females of one and two gestation periods from the same density. Fecundity and fertility were significantly higher among females of one and two gestation periods from the lowest than from the highest density.     

Hypoxia inhibits fish spawning via LH-dependent final oocyte maturation

To evaluate the effects of long term hypoxia exposure on fish spawning, mature common carp, Cyprinus carpio carpio (Linnaeus) were subjected to either normoxia (7.4+/-0.2 mgO(2)mg O(2) L(-1)) or hypoxia (1.0+/-0.2 mgO(2)O(2) L(-1)) for more than two months. Gonadosomatic index (GSI), and concentrations of serum luteinizing hormone (LH), testosterone (T), and estroldiol (E2) were measured and gonad histology examined. Hypoxia inhibits fish spawning even though the gonad and oocytes developed under hypoxia exposure. LH levels of female carp were significantly decreased upon chronic exposure to hypoxia, and the final oocyte maturation in hypoxic females was significantly retarded. The results indicated that hypoxia may inhibit fish spawning through LH-dependent final oocyte maturation. In addition, no courtship was observed in hypoxic males. In conclusion, hypoxia impairs fish ovulation and, therefore, spawning and reproduction. LH levels were reduced leading to a failure of oocyte maturation. This, along with a lack of courtship by males may be the major mechanisms involved in hypoxic inhibition of reproduction in carp.     

Some aspects of the reproductive biology of the clupeid Ilisha africana (Bloch) off the Lagos Coast, Nigeria

The maturation development, spawning period, oogenesis, fecundity and sex ratio of Ilisha africana off the Lagos Coast, Nigeria were investigated. Six macroscopic and seven microscopic stages of gonad maturity were identified. Maturity was attained at 12.0 cm total length. Breeding occurred throughout the year with the peak spawning between May and December. Oogenesis consisted of seven stages and breeding marks were observed in the species. Fecundity ranged between 2098 and 11 687 eggs. There was a slightly higher correlation between fecundity and weight than between fecundity and length. The egg diameter varied from 0.77 to 1.35 mm. The overall sex ratio was 1 male to 0.97 female. The females however predominated over the males during most months in the peak breeding period.     

Population ecology of the paddy field-dwelling fish Channa gachua (Günther) (Perciformes, Channidae) in Sri Lanka

A population of Channa gachua in a small irrigation canal that supplies rice fields was studied by monthly sampling over 2 years. The population density was positively correlated with the rainfall and varied from 0.34 to 0.95 individuals m⁻². The growth parameters of the von Bertalanffy growth equation determined on monthly size–frequency data were L_x = 179 mm total length and K =0.50. Overall male to female ratio was 0.82 and there were more females than males in the middle size classes. Spawning occurred throughout the year, but all evidence indicated enhanced breeding during major rainy periods of May to July and October to December. The length at first spawning was 102 mm, which is reached in about 20 months. Fecundity, which varied between 389 and 2130, was positively correlated with gonad weight, body weight and total length. Longevity and natural mortality were estimated as 6 years and l.27 yr⁻¹, respectively. However, 99% of the population appeared to live for only 3 years. The mean biomass, average annual production and turnover ratio of the population were 7.35 g m⁻², 12.06 g m⁻² and 1.64, respectively.     

Differences in reproductive strategies between obscure puffer Takifugu obscurus and ocellated puffer Takifugu ocellatus during their spawning migration

Reproductive strategies were compared between obscure puffer Takifugu obscurus and ocellated puffer Takifugu ocellatus captured in waters near Yangzhong Island in the lower reaches of the Yangtze River during the spawning migration season from February to June in two consecutive years (1995 and 1996). Results showed that obscure puffer and ocellated puffer have adopted different reproductive strategies, including different spawning times, different size at maturity, and different number and size of oocytes, resulting in two different larval sizes. In detail, the timing of the spawning migration and status of gonad development of obscure puffer was about 1 month earlier than that of ocellated puffer; the obscure puffer was obviously longer and heavier than ocellated puffer in both mature male and female fish; mean GSI of obscure puffer females (15.8%) was significantly higher than that of ocellated puffer females (14.6%); the average diameter of ocellated puffer eggs (1.49 ± 0.12 mm) was significantly larger than that of the obscure puffer (1.22 ± 0.08 mm); and obscure puffer females (320.8 oocytes mg^?1 somatic wet weight) had significantly higher relative fecundity than ocellated puffer females (125.2 oocytes mg^?1 somatic wet weight). These differences in reproductive strategies between two closely related species of the Takifugu genus indicate that both obscure puffer and ocellated puffer fit the r/K dichotomy. Obscure puffer shows K‐selected characters with maturity at relatively large size and r‐selected characters with relatively many and small offspring, whereas ocellated puffer shows r‐selected characters with maturity at a relatively small size and K‐selected characters with relatively few and large offspring.     

Reproductive biology of Astyanax fasciatus (Pisces: Characiformes) in a reservoir in southeastern Brazil

To study the reproductive biology of Astyanax fasciatus (Cuvier, 1819) in Furnas Reservoir, Minas Gerais, Brazil, a total of 1307 specimens were captured in bimonthly sampling from August 1993 to July 1994. The sex ratio showed a predominance of females, representing 65.2% of the total sample. The stages of gonadal maturation and spawning type were determined through macro- and microscopic analysis of the gonads as well as by variation in the gonadosomatic indices. A . fasciatus showed fractionated spawning and reproduction throughout the year, with spawning peaks influenced by water temperature and rainfall. Males and females initiated gonadal maturation at a mean standard length of 6.4 and 6.5 cm, respectively. During gonadal maturation, females showed a reduction in hepatosomatic index (from 1.06 to 0.83), suggesting that hepatic substances are transferred from the liver to the ovaries. In the reproductive period, A. fasciatus had lower stomach repletion and coelomic fat indices, indicating that this species ate less and consumed more fat reserves when reproduction had the maximum GSI (11.3 for females and 2.7 for males). Weight of the ovaries had a positive influence on the condition factor (K1), as the lowest K2 value (weight without gonads) were obtained during maturation, confirming the influence of the ovaries upon the physiological condition of the females. In weight–length relationship, parameter b was 3.02 for females and 2.23 for males, and the correlation coefficient ( r ²) was 0.84 and 0.66, respectively.     

The reproductive traits of Scomber japonicus (Houttuyn, 1782) in the Eastern Adriatic Sea

The aim of the present study was to investigate the fisheries biology of Scomber japonicus (Houttuyn, 1782) in the Adriatic Sea, to enable implementation of appropriate regulatory measures for fisheries. Reproductive biology and sexual patterns of S. japonicus (chub mackerel) were examined from monthly random samples of commercial catches from purse seine nets (10 mm stretched mesh size) in the eastern Adriatic Sea. The annual mean chub mackerel fork length (FL) was examined from 1998 to 2007. Average fork lengths were approximately 25 cm between 1998 and 2002, decreasing to 22 cm between 2003 and 2007. In addition, sex ratios fluctuated as a function of size and month: females were more abundant in the smaller length classes, as well as before and after spawning; males were more abundant in the larger length classes and during spawning. The overall sex ratio was 0.68; females were significantly predominant. 50% of the chub mackerel population was sexually mature at a FL of 18.3 cm, and both males and females reached sexual maturity at FLs of 20.4 cm and 16.8 cm, respectively. Based on gonad maturity stages, gonad weight and gonadosomatic index, chub mackerel spawned from May to August, with peak spawning in June for both sexes. The mean absolute fecundity was 181 277 ± 62 090 oocytes per ovary. A positive exponential correlation was found between fecundity and fork length, gonad weight, and total chub mackerel weight (r² = 0.640; r² = 0.686 and r² = 0.607, respectively).     

The effects of three different dietary protein levels on the fecundity in the guppy, Poecilia reticulata (Peters)

Experiments were performed to investigate the influences of dietary protein on fecundity in the guppy, Poecilia reticulata . No significant differences in total body length and standard length were found between females given food with different protein levels. A significantly higher total body weight was found both in females fed on 47% and 31% protein compared with females given 15% protein. The frequencies of females with only intrafollicular embryos or with only ova or both embryos and ova in the ovary were not significantly different between populations given different protein amounts. There was a significantly lower frequency of females without embryos and ova but with previtellogenic oocytes in the group given highest protein (47%) compared with the group fed on 31%. Significantly higher values of ovarian length, width and gonad weight were found among females given 47% protein compared with females fed on 15% protein. The ovarian height was significantly higher only in females given the highest protein amount when compared with the group fed on 31% protein. No significant difference of ovarian volume was found between groups given different protein contents. The average gonadosomatic ratio was highest in the groups fed on 47% protein. Significant correlations were found between gonad weight and total body weight of females in each group. Ovarian volume versus total body weight as well as versus gonad weight also yielded significant correlations in all groups. No significant differences of fecundity were found either between females given 47% and 31% protein or between females fed on 47% and 15% protein. A somewhat higher number of ova per individual fish was, however, observed among females given 31% when compared with females fed on 15% protein. The ovum size remained the same between groups given different protein contents. No significant correlation was found between fecundity and ovum diameter in the groups.     

The ecology of populations of dace, Leudscus leuciscus (L.), from two tributaries of the River Wye, Herefordshire, England

Samples of dace were taken from the Afon Llynfi and the River Lugg each month for 13 consecutive months. Male dace grew faster than females and rates were comparable with those described for several other populations. Variations in year-class strengths were observed and these differed between the populations although 1959 was strongly represented in both. The sex ratio varied throughout life and disturbances in the expected ratio were attributed to the attainment of sexual maturity at the end of the fourth year of life (3+). Observation of the seasonal gonad cycles indicated that spawning occurred in March or April. Fecundity was estimated at about forty eggs per gram of total weight. A seasonal cycle of feeding activity was established with highest food consumption during summer. There was evidence of a fast during the spawning period. No correlation between temperature and feeding activity could be established. Dace were omnivorous, but aquatic insects and algae were the main components of the diet. Seasonal variations generally followed availability. The importance of aerial insects declined with age. Many differences in feeding habits were detected between the populations.     

Biology of the amphioxus,Branchiostoma belcheri in the Ariake Sea,Japan II. Reproduction

We investigated the reproduction of the amphioxus Branchiostoma belcheri in the southern Ariake Sea, Japan, from 1999 to 2002. Gonads were very small or almost absent from September to December and began to develop in January. Changes in the gonad length index indicated that spawning began in mid June, after the gonads had attained maximum size. Although most gametes were extruded during the first spawning, some remained in the gonads. Shrunken gonads were much smaller but contained gametes, and the second spawning occurred around 10 July. It was unclear whether all adults spawned twice, but a large proportion of individuals did. After the first spawning, both males and females lost 30% of their body weight. Most one-year-old individuals did not spawn, and the minimum size at maturity was ca. 20 mm. We found no termination of reproduction among very large individuals. The average number of gonads was significantly larger on the right side of the body (26.2 in males and 26.1 in females) than on the left side (24.0 in males and 23.5 in females). We found two hermaphrodites in a total of 11,184 specimens examined. Each had four or three ovaries among a total of 45 or 54 gonads.     

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.     

Spatio‐temporal variability in reproductive ecology of sand flathead,Platycephalus bassensis,in three Tasmanian inshore habitats: potential implications for management

Temporal and spatial variability in gonad development, duration of spawning period, and size/age at maturity were investigated in sand flathead, Platycephalus bassensis. A 3‐year study (2001–2003) revealed that variation in gonad weight with somatic weight was a function of an interaction between season and study location (Coles Bay, Georges Bay, and Tamar River estuary). Highest gonad weight was recorded in Coles Bay in early summer, which in comparison to Georges Bay, was approximately 50% higher and occurred 3 months later. Tamar River mature individuals were reproductively inactive during the spawning season. The proportion of mature individuals at different stages of maturity differed significantly among the three locations across all times. Coles Bay individuals were reproductively active from October to March, while in Georges Bay females with hydrated oocytes were seen in September but were absent from the population by November. Interannual variation in initiation and duration of spawning activity was evident in each location. Apart from male size at maturity, for both sexes the size and age at maturity (L₅₀) was higher in Georges Bay compared to Coles Bay. The results emphasize the necessity of temporal and spatial management based on population differences in reproductive ecology.     

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.     

Quantitative studies on the reproduction of black scallop,Chlamys varia (L.) from lanveoc area (Bay of Brest)

The values of gonad index calculated by the allometric relationships of dry gonad weight over dry body weight of the black scallop, Chlamys varia (L.), gave peak values for May and September during 1976 and 1977. Histological studies on gonads from natural populations and periodic biopsies of gonads from marked animals in experimental cages showed that, in addition to intermittent spawnings between May and September, two peak spawning periods were noteworthy. These were a partial spawning in May–June and a complete spawning in September–October. The energy spent as gametes for these two spawnings has been calculated for individual males and females in each age group of the population and the cost of spawning by the whole population has been estimated. The population spent more energy for spawning in 1977 (144.49 kcal/100 m²) than in 1976 (83.70 kcal/100 m²) and the reproductive effort of an average adult was 0.04 in 1976 and 0.10 in 1977.     

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.     

The cost of sex: quantifying energetic investment in gamete production by males and females

The relative energetic investment in reproduction between the sexes forms the basis of sexual selection and life history theories in evolutionary biology. It is often assumed that males invest considerably less in gametes than females, but quantifying the energetic cost of gamete production in both sexes has remained a difficult challenge. For a broad diversity of species (invertebrates, reptiles, amphibians, fishes, birds, and mammals), we compared the cost of gamete production between the sexes in terms of the investment in gonad tissue and the rate of gamete biomass production. Investment in gonad biomass was nearly proportional to body mass in both sexes, but gamete biomass production rate was approximately two to four orders of magnitude higher in females. In both males and females, gamete biomass production rate increased with organism mass as a power law, much like individual metabolic rate. This suggests that whole-organism energetics may act as a primary constraint on gamete production among species. Residual variation in sperm production rate was positively correlated with relative testes size. Together, these results suggest that understanding the heterogeneity in rates of gamete production among species requires joint consideration of the effects of gonad mass and metabolism.