Effect of spawning number and ration on reproductive performance of the batch-spawning three-spined stickleback Gasterosteus aculeatus

An experiment quantified the effect of food ration and spawning number on the breeding season reproductive performance of batch-spawning, female three-spined sticklebacks Gasterosteus aculeatus . Individually housed females were maintained on fixed rations of 2, 4, 8 or 16% of their initial postspawning mass of live enchytraeid worms from immediately after their first spawning until they ceased spawning. Number of spawnings correlated positively with ration. Total breeding season reproductive investment (total wet mass of eggs produced as a percentage of initial postspawning female mass) ranged from 38% at the 2% ration to 147% at the 16% ration. At the lower rations, postspawning mass, batch fecundity and wet and dry masses of the batch declined over successive spawnings, with the rate of decline inversely related to ration. At the highest ration, there was no decline in batch fecundity and postspawning mass increased over successive spawnings. Mean reproductive investment per inter-spawning interval was higher at the highest ration, but at all rations declined over successive spawnings. Mean reproductive effort per inter-spawning interval (wet mass of eggs spawned as a percentage of the wet mass of food consumed over the inter-spawning interval) was inversely related to ration. At the higher rations, reproductive effort showed no trend over successive spawnings. Neither spawning number nor ration had a systematic effect on egg diameter, wet mass per egg, dry mass per egg or total lipid content of the eggs. If the rate of food intake was insufficient, although batch fecundity declined, the main adjustment was a reduction in the number of spawnings in the breeding season.     

Effect of variable food levels on reproductive performance of breeding female three-spined sticklebacks

The study examined the response to variation in food ration within a breeding season on components of the reproductive performance of mature female three-spined sticklebacks Gasterosteus aculeatus , a batch spawner. The main effect of food availability was on the duration of the inter-spawning interval (ISI). Low ration resulted in an increase in the number of days until the next spawning. A low ration during one inter-spawning interval increased the length of the next ISI, even if there was a high ration in the latter. Females that experienced high rations for two successive ISIs showed an increase in batch fecundity over the successive spawnings. There was no tendency for fish on low rations to produce bigger eggs or eggs with higher lipid concentration. Females on low ration had a lower carcass dry matter concentration and a lower liver dry weight immediately after their third spawning. Female sticklebacks responded to a low ration with little change in egg number or size, but a slight reduction in physiological condition. Egg characteristics and, to a lesser extent, batch fecundity are insensitive to current rate of food consumption, but the interval between spawnings is sensitive to both current and previous rates of consumption.     

Do random fluctuations in the intervals between feeding affect growth rate in juvenile three-spined sticklebacks?

This study measured the effects of regular and irregular intervals between feeds on the growth performance of juvenile Gasterosteus aculeatus . The experimental period was 21 days at 14) C and photoperiod of 10L: 14D. The fish were housed individually. The control fish received a constant ration every day. Fish on a constant interval were fed on days 1, 5, 9, 13, 17 and 21. Fish on a random regime were fed with the same average interval between feedings, but the interval varied randomly between 1 and 5 days. The rations were calculated so that over the 21-day period, all fish were supplied with the same total quantity of food. The two ration levels were: 2% (maintenance ration) and 6% of the initial body weight per day. At a given ration, the feeding interval had no significant effect on specific growth rate, RNA/DNA ratio and lipid contents. The percentage dry matter was slightly, but significantly lower in treatment groups than in the control group. Groups receiving a mean 2% ratio consumed all the food supplied. At a mean ration of 6%, the control group ate 100%, the regular interval group 95·4% and the irregular interval 98·3% of the total food supplied. For the temporal patterns of feeding used, the fish were able to adjust their food intake, when food become available, to compensate for short periods of food deprivation and maintain their growth performance except for a decrease in dry matter content.     

Effects of overripening on spawning behaviour and reproductive success of Atlantic salmon females spawning in a controlled flow channel

The influence of overripening on salmonid egg quality has often been described in artificial breeding. However, no information exists on the impact of this factor on fish reproducing naturally, although it is likely to be critical, especially in disturbed environments. Pairs of Atlantic salmon in which the size of the male and the female was evenly matched were released in a spawning channel from 1 to 12 days after ovulation. Handling did not affect egg quality. Overripening had a negative effect on egg viability. Egg mortality, infertility and malformations increased with the number of days after ovulation. In females which had just matured, 16·7% of their recovered eggs were dead, 3·1% were unfertilized and 1·7% were malformed, as compared to 25·4, 9·7 and 5·4%, respectively, in females that had been mature for at least a week. Overripening played a role in both female spawning behaviour and capacity to spawn, as females which had been mature for over a week released their eggs more quickly while they retained more ova in their abdominal cavity than early spawning females. Thus, in studies of reproductive behaviour and in particular of sexual selection, it is critical to take into account the maturation status of females since the duration of spawning activity is used by various authors to determine female sexual motivation. Alternatively, overripening may decrease the female selectivity for mates.     

Reproductive biology of the endangered Oxleyan pygmy perch Nannoperca oxleyana Whitley

The reproductive biology of the Oxleyan pygmy perch Nannoperca oxleyana is described from simultaneous studies of wild populations in north-eastern New South Wales and mature fish held in aquaria. In the wild, 50% of males and females matured at total lengths of 24·0–25·9 and 28·0–29·9 mm, respectively. The species displays sexual dichromatism during the spawning season, with males developing more intense red and brown fin and body colouration, and black pelvic fins. Captive male broodfish displayed territoriality during the breeding season, closely guarding sites within artificial, plant-like substrata in which pairs of fish spawned adhesive eggs. Protracted serial spawning of wild and captive fish occurred from September to April and May at mean water temperatures ≥16·6° C and day length ≥10·7 h. Captive broodfish spawned on an average of 57% of days during the 256 day spawning period. Gonado-somatic indices averaged 0·7% for all ripe males and 4·1–4·2% for all ripe females collected. Mean total and batch fecundities of captive females were 1323 eggs per fish and 7·8 eggs per fish per day, respectively, and relative fecundity was 587 eggs g⁻¹ of body mass. Batch fecundity of wild females was estimated at 7·8 eggs per fish. The adaptive significance of this reproductive strategy in a harsh, variable environment is discussed.     

The induced spawning of roach, Rutilus rutilus (L.), with the antioestrogens clomiphene and tamoxifen

The induced spawning of gravid roach was investigated using the antioestrogens clomiphene and tamoxifen. Three dose levels ofeach were used: 0·1, l or 10 mgkg^-1, given twice with a 4-day interval to groups of eight fish with saline controls. Running males were randomly distributed. Tamoxifen, when injected at a rate of l mg kg^-1 was found to be most successful. This treatment induced ovulation in five of the six females, and profuse spawning on 3 consecutive days, 4 days after the first injection. Clomiphene induced ovulation and spawning in one of six females at 2 × 10 mg kg^-1, and two of eight females at 2 × l mg kg^-1, respectively 6 and 7 days after the first injection. The eggs produced showed normal devel-opment. No control fish ovulated or spawned. Both drugs probably act by indirect mechanisms, blocking sex steroid feedback inhibition of gonadotropin (GtH) secretion at the pituitary, thereby inducing a plasma GtH surge. The results of this experiment suggest that tamoxifen may be an effective substitute for pituitary preparations in the induced spawning of fish.     

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.     

Fecundity of the American eel Anguilla rostrata at 45° N in Maine, U.S.A.

The northern portion of the geographic range of the American eel Anguilla rostrata may contribute a great proportion of the reproductive potential to this panmictic species because of apparent increases in average female size and female percentage with latitude. The regressions of fecundity on body length and on body weight of 63 female eels captured at about 45° N latitude on their spawning migration to the sea were log F= 1·2601 + 2·9642 log L and log F= 4·1646+0·9153 log W , where F is fecundity, L is total length (cm), and W is total weight (g). Length and weight each explained about 90% of the variation in fecundity. Estimates of fecundity from counts of aliquots of eggs ranged from 1·84 million to 19·92 million eggs for eels ranging in length from 45 to 113 cm, nearly the range of sizes of migrating females reported in the literature. Fecundities of the American eel were greater than reported in one study at about 37° N and greater than reported for the European eel, A. anguilla , shortfin eel, A. australis , and longfin eel, A. dieffenbachii . If a geographic cline in fecundity does exist in American eels, it is established anew each generation because the species forms a single panmictic population.     

Reproductive biology of the threatened golden galaxias Galaxias auratus Johnston and the influence of lake hydrology

Golden galaxias Galaxias auratus (31–235 mm fork length, L _F) were collected monthly from littoral habitats in Lakes Crescent and Sorell, Tasmania, Australia, between July 2000 and December 2002. Spawning habitats were identified and monitored in both lakes, and surveyed in Lake Crescent. Trends in gonado-somatic indices and reproductive stages of development indicated that gonad development in both sexes begins in midsummer and peaks in late autumn to early winter. Males mature at smaller sizes (50% at 52 mm L _F) than females (50% at 76 mm L _F), larger individuals are predominately females (95% of fish ≥138 mm L _F), and overall male to female ratios are female biased ( c . 1:2). Spawning occurs late autumn to early spring (water temperatures = 1·4–9·7° C) with peaks in spawning activity in winter (mean water temperatures <5° C). Demersal adhesive eggs ( c. 1·5 mm diameter) were found on cobble substrata ( c. 20–250 mm diameter) in littoral areas ( c. 0·2–0·6 m deep) and fecundity of fish 71–181 mm L _F ranged from 619 to 14 478 eggs. The rate of change in water level over the 20 days prior to monthly sampling was important in explaining the occurrence of spent fish and this accounted for temporal differences in spawning between the populations. Lake hydrology influences the reproductive cycle of G. auratus by possibly providing a stimulus for spawning and it controls the availability of spawning habitat in Lake Crescent. Seasonal hydrological cycles ( i.e. rises during late autumn to winter) and a minimum water level of 802·20 m Australian Height Datum in Lake Crescent during autumn (above which littoral areas of cobble substratum are inundated) are critical to G. auratus populations.     

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 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.     

Fecundity, lifespan and egg mass in butterflies: effects of male-derived nutrients and female size

1. Effects of larval reserves and nutrients received as adults on fecundity and lifespan in female Danaus plexippus (the Monarch Butterfly) were measured to determine the relative importance of different sources of nutrients for reproduction and somatic maintenance.
2. Egg-laying lifespan was correlated with female size but not with the amount of male-derived nutrients or adult food concentration.
3. Lifetime fecundity was higher when females received a large first spermatophore, but was not affected by female size when lifespan was controlled or by adult food concentration.
4. At the end of their lives, females contained unlaid eggs and retained, on average, 88% of their initial mass. This proportion was unchanged in two years, although mean egg-laying lifespan varied from 22·5 to 28·7 days.
5. Egg mass decreased over the female lifespan, and was correlated with female size.
6. These results suggest that larval reserves are more important for somatic maintenance than adult income, but that the protein-rich nutrients received from males contribute to egg production. This supports theoretical predictions and empirical studies of other Lepidoptera showing that larval reserves are less likely to affect fecundity when the adult income can contribute substantially to egg production.     

Sex structure and potential female fecundity in a Epinephelus guttatus spawning aggregation: applying ultrasonic imaging

Ultrasonic imaging was used to determine the spawning population structure and develop a fecundity estimation model for a red hind Epinephelus guttatus spawning aggregation within the Red Hind Bank Marine Conservation District, St Thomas, U.S.V.I. The spawning population showed considerable within‐month and between‐month variation in population size‐ and sex‐structure. In the spawning season studied, males appeared to arrive at the aggregation site first in December although females represented a large proportion of the catch early in the aggregation periods in January and February. Spawning occurred in January and February, and size frequency distributions suggested that an influx of small females occurred during the second spawning month. An overall sex ratio of 2·9 : 1 (female : male) was recorded for the whole reproductive season. The sex ratio, however, differed between months and days within months. More females per male were recorded in January than in February when the sex ratio was male biased. Fecundity estimates for this species predicted very high potential fecundities (2·4 × 10⁵−2·4 × 10⁶ oocytes). The ultrasound model also illustrated a rapid increase in potential female fecundity with total length. Ultrasonic imaging may prove a valuable tool in population assessment for many species and locations in which invasive fishing methods are clearly undesirable.     

Batch fecundity and spawning frequency of invasive Hemiculter leucisculus (Basilewsky, 1855) in Erhai Lake,China

Hemiculter leucisculus are multiple spawners with indeterminate fecundity, although previous fecundity estimates have assumed a determinate spawning pattern. Batch fecundity and spawning frequency of H. leucisculus were studied in Erhai Lake, China, in which the fish, as an exotic species, has become a successful colonizer. Spawning frequency was 16.05 %, as determined from the percent of females with postovulatory follicles 12 to 36 h old during the peak reproductive period (from May 27 to August 8). The average interval between spawning was 6.2 days and there were more than 16 total spawning batches during the peak reproductive period. The average batch fecundity (mean ± SD) was 11,934?±?5,921 hydrated oocytes in 40 females (standard length or SL: 9.1–14.4 cm), while relative batch fecundity was 560?±?137 eggs g^?1 wet weight. The total potential annual fecundity was approximately 190,944 oocytes over the whole spawning season and was much higher than the estimated standing stock (31,585 oocytes) of yolked oocytes (36 females, SL: 9.5–16.2 cm) at the beginning of the spawning season in April 2010. This high annual fecundity is likely to have contributed to successful invasion of Erhai Lake by H. leucisculus.     

Egg cannibalism in Barents Sea capelin in relation to a narrow spawning distribution

The spawning distribution of Barents Sea capelin Mallotus villosus off northern Norway in 2002 was restricted to a narrow area at the easternmost spawning grounds. There was an increase in abundance and a marked shift in aggregation area from east towards west during 11 March to 4 April, as the capelin went from prespawning through spawning to spent stages. The capelin stomachs were either empty or contained capelin eggs, often mixed with sand. Both the occurrence of cannibals and stomach fullness increased with stage of maturity, being highest in spent fish, and higher in males than in females. No differences were found between cannibals and non-cannibals of the same sex when adjusted for total length ( L _T), age, L _T at age and condition factor. At similar stomach fullness, the females had consumed more eggs than the males. This was attributed to a higher proportion of empty and broken eggs and less sand in females compared to males. In the full stomachs, the mean consumption in females and males was 623 and 334 eggs, respectively, whereas the respective maximum egg consumption was 871 and 683 eggs. The minimum estimates (given no digestion) of mean ± 95% CL egg consumption in spent females and males were 75·4 ± 6·9 and 58·4 ± 12·0 eggs, respectively, implying a minimum mortality of 1–2% of the total egg production caused by cannibalism.     

The influence of previous feeding regime on the compensatory growth response of maturing and immature Arctic charr, Salvelinus alpinus

Alternating periods of food deprivation with those of unlimited provision of food depressed the growth of Arctic charr, Salvelinus alpinus , below that of controls. Fish that were deprived of food and then fed on alternate weeks (1:1) were larger than those that were exposed to periods of 1 5- or 3-week deprivation and feeding (1·5:1·5 or 3:3). On receiving excess food supplies following 24 weeks on the restricted feeding regimes the previously-restricted fish grew more rapidly than the controls. The greatest compensatory growth was displayed after the 3:3 regime, followed by the 1·5: 1·5 and then the 1:1 feeding regime. At the termination of the experiment there were no significant differences in body weight between fish fed according to the different regimes during the period that food restriction was imposed. Growth patterns of the immature males and females were similar, but mature males were significantly lighter than the immature fish by the end of the experiment. Both immature and maturing fish displayed a compensatory growth response on return to adequate feeding. Beginning food restriction in May did not influence the proportions of male fish ( c . 60%) which were mature in the autumn.     

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.     

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 .     

Spawning characteristics of the anadromous brown trout in a small Swedish stream

The spawning pattern of the anadromous brown trout Salmo trutta was studied in Själsöån, a small stream in Gotland, Sweden, during eight winters between 1992–1993 and 1999–2000. The total length ( L _T) at spawning was normally distributed (185–890 mm) for females and multimodal for males (300, 400 and 550 mm most frequent length classes). Spawning males were significantly younger (2+ to 4+ years) than females (3+ to 5+ years). The sex‐ratio at the beginning and at the end of the spawning season favoured males. The mean ±  s . d . number of spawners was 70 ± 16 individuals per year. Migration into and out of the stream occurred between November and June. The highest number of spawning fish was found in the stream at the end of November or at the beginning of December. Migration mainly occurred during high water flow and at night. The majority of the females entered the stream and spawned the same (29·3% of all the females) or the next night (32·8% of all the females) while males may have stayed for 2 to 3 weeks (21·3% of all the males) in the stream before spawning. Males usually remained much longer in the stream (mean ±  s . d . 45 ± 56 days) than females (16 ± 30 days). Females lost more mass in the stream (mean ±  s . d . 17·3 ± 8·6%) than males (7·7 ± 9·6%). For both sexes, mass loss was positively correlated with the time spent in the stream. Only 7·3% of the males and 5·7% of the females occurred in the stream for >1 year. Spawning took place only during the night.     

Reproductive biology of the mosquitofish in a permanent natural lagoon in south-west Spain: two tactics for one species

Introduced Gambusia holbrooki in a natural lagoon of southern Spain consisted of two age groups: 1992 cohort, 7-11 months old and 1993 cohort, <4 months old. In the 1992 cohort, females grew even during the gestation period at about 0·30 mm day^-1. In the 1993 cohort, females displayed a high growth rate (0·55 mm day^-1) and reached reproductive size in a few weeks, but stopped growing when they matured. All the 1992 cohort reproduced from mid-May to mid-June, but only 50% of the 1993 cohort reproduced, from mid-August to mid-September. Reproducing females were significantly larger in the 1992 cohort (39·8 mm) than in the 1993 one (34·8 mm). The largest 1992 females cohort had reproduced previously; the 1993 cohort had not. The mean dry weight of intra-ovarian embryos decreased to a minimum immediately before birth. These metabolic costs represented 29·8 and 31·4% of the initial weights of the 1992 and 1993 embryos, respectively. Mean dry weight of full-term embryo was significantly higher in the 1992 cohort (0·80 ± 0·129 mg; 95% CL) than in the 1993 one (0·70 ± 0·086 mg; 95% CL). With more females in 1992, cohort fecundity was considerably higher (number of embryos=7151; 63%) than in the 1993 (4193; 37%) cohort. The population completed two generations each year (spring and summer). The spring stock grew slower than the summer one but continued to grow during the gestation period, reaching larger final lengths, with more synchronous reproduction and clearer evidence of a second reproductive event. Each reproductive stock displayed its own life-history characteristics, with significant differences between mean length of reproducing females, growth rate, mean brood size, offspring size, standard fecundity and minimum length at reproduction.