Effects of varying temperature and food availability on growth and reproduction in first‐time spawning female Atlantic cod

The somatic growth, sexual maturation and fecundity of individually marked first‐time spawning female Atlantic cod Gadus morhua were examined under different varying temperature and feeding regimes over the months preceding spawning. A negative correlation between somatic and oocyte growth was found, reflecting the changing energy allocation pattern. Nevertheless, the somatic growth of mature individuals was at least as high as those of immature fish over the period of vitellogenesis. Potential fecundity was positively correlated with body size, but neither temperature or feeding regime significantly affected this relationship. Consequently, fish with unlimited feeding opportunity invested more energy into somatic growth during vitellogenesis compared to those held under a restricted ration. This indicated that once Atlantic cod had made the decision to invest in first reproduction, they allocated a certain amount of energy relative to their size into egg production and any surplus was invested into somatic growth. Low temperature led to an arrest in the onset of vitellogenesis and significantly affected the number of females that matured.     

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.     

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.     

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.     

The feeding, growth and behaviour of juvenile cod, Gadus morhua L., in cold environments

Most studies on feeding-related activities of fish in cold water (0° C) have not used eurythermal species or examined a broad suite of activities. In this study we report on the feeding, growth, behaviour and conversion-efficiency of 0 + Atlantic cod in response to water temperature and food availability. In one experiment, activity, opercular beat and growth decreased with decreasing temperature over the range 8·3–0·6° C. However, the conversion efficiency increased with decreasing temperature, a result partly explained by the relative change in activity and opercular rate. In another experiment, temperature had a linear effect on growth in cod fed different food rations. Both food availability and temperature were found to influence growth in group 0 + cod. The specific growth rate of cod in this experiment exceeded 2% day^-1. Growth data collected from cod sampled in the field during the cold-water period showed a progressive increase of mean length over the season. Overall, these results demonstrate that 0 + cod have adapted to life in cold-water environments, in contrast to adult cod which mostly inhabit warmer environments.     

Temperature experienced during vitellogenesis influences ovarian maturation and the timing of ovulation in common wolffish

Temperature treatment of common wolffish Anarhichas lupus during vitellogenesis affected the time of final maturation; ovulation in fish held at 8 and 12° C from mid-April to October was about four and five weeks delayed, compared with a 4° C group. Fish in the 8° C group had significantly larger eggs than those in the 4° C and the 12° C groups, and a significantly higher egg production than fish in the 12° C group. Temperature treatment did not affect either fertilization rate or relative fecundity, but absolute fecundity was significantly lower in the 12° C group than the other groups due to poor growth of the fish at high temperature. This did not affect the numbers of spawning individuals. There was a trend towards lower egg survival to the eyed stage in the 12° C group compared to the 4 and 8° C groups, although the effect was not statistically significant. The results indicate that both the timing of final maturation and investment in ovarian growth in common wolffish are affected by temperature experienced during vitellogenesis.     

The reproductive cycle of Coregonus lavaretus (L) in Loch Lomond, Scotland, inrelation to seasonal changes in plasma cortisol concentration

The reproductive cycle of Coregonus lavaretus (L.) in Loch Lomond, Scotland, was investigated by monthly sampling. Spawing takes place during the first 3 weeks of January, on offshore gravel banks. Males congregate on the spawning-grounds throughout the spawning period, while ripe females cruise in unisexual shoals in deeper water. Individual females migrate to the spawning-grounds as ovulation takes place, spawn, and return to deep water. After spawning, the gonadosomatic ratio and somatic condition factor of both sexes decrease until June or July. Gonad recrudescence occurs in males between July and October, and in females between July and December. The somatic condition factor of both sexes rises from its minimum in June or July to its maximum in September. A competitive protein binding assay was used to determine cortisol levels in 0–1 ml plasma samples of individual fish. The lowest cortisol levels occurred in fish caught by seine-netting and killed immediately by anaesthesia in MS 222 (Sandoz). Seine-netting and killing by concussion induced higher cortisol levels, in the range of 3–5 ng/100 ml. Cortisol levels in fish caught by gill-netting for 18 h were 3–6 times higher than in seine-netted fish, and the method of killing was immaterial in this case. The length of time spent in the gill-net had no significant effect on cortisol level, but maintenance of the fish in aquaria for 24–80 h elicited high cortisol levels of over 50 μg/100 ml. Post-mortem delay before blood-sampling resulted in lowered cortisol levels. Seasonal variations in cortisol level were determined in gill-netted fish throughout the year, and in seine-netted fish when available. Cortisol levels were high in both sexes in September; and very high in ovulating females caught on the spawning-grounds, though not in ovulating females caught off the spawning-grounds.     

The influence of feeding regime on sexual maturation, fecundity and atresia in first-time spawning turbot

Exposure of adolescent turbot Scophthalmus maximus to low rations during vitellogenesis, covering the 4 months immediately prior to spawning, led to a drop of 70% in mean ovary weight, and was associated with poor growth of the vitellogenic oocytes or, in a third of cases, the absence of vitellogenic oocytes. Exposure to low rations during the recruitment of vitellogenic oocytes, 4–8 months prior to spawning, produced a more variable response. A few of these females produced well-developed vitellogenic oocytes, but in most, vitellogenic oocytes were absent or only poorly developed and in some cases there was a high incidence of atresia. Exposure to intermediate rations throughout oocyte recritment and vitellogenesis also depressed oocyte development. In maturing females the number of non-atretic vitellogenic oocytes (relative potential fecundity) averaged 998 per g of somatic body weight of fish at the start of the spawning season in mid-June. Maturation of the males was not significantly influenced by the dietary regime. The small size of the testes and the low levels of milt production, even in fish fed high rations immediately prior to spawning, showed that reproductive investment in males was much lower than in females.     

In vitro production of ovarian steroids in yellow perch (Perca flavescens): effects of photothermal manipulation, gonadotropin and phorbol ester

We investigated the capacity of ovaries of yellow perch to produce steroid hormones in vitro and the ovarian response to gonadotropin and phorbol ester during the annual reproductive cycle. The effects of photothermal manipulation on perch gonadal steroidogenesis and its regulation have also been examined. Initially, all females kept indoors were exposed to the same water temperature and photoperiod. By the end of August, following the first sampling, fish were submitted to different photothermal regimes. Group A was maintained under photothermal conditions characteristic for southern Ohio. Group B was submitted to a condensed light/temperature regime designed to accelerate physiological changes that depend on photothermal stimuli. In group A, basal in vitro production of ovarian estradiol (E2) was the highest in October and November, and hCG significantly stimulated E2 secretion during the entire period of vitellogenesis (October-January). In this group, the highest production of basal testosterone (T) was observed before spawning. hCG-stimulated production of T was highest at the beginning of vitellogenesis. Gonadotropin stimulated T production before spawning, a time when gonadotropin was unable to stimulate E2 production. Phorbol ester (PDBu) stimulated E2 and T production during vitellogenesis at the same time points as hCG did (E2: December, January; T: December). hCG-stimulated T production was not mimicked by PDBu in April. Condensing of the photothermal cycle resulted in diminished ovarian production of E2 during vitellogenesis. Moreover, the fish submitted to a condensed photothermal cycle demonstrated augmented T production during the postvitellogenic stage of ovarian development. Ovaries of group B did not respond to PDBu. Generally, the seasonal fluctuations in ovarian capacity to produce E2 and T as well as in gonadal responsiveness to gonadotropin observed in female yellow perch illustrate the dynamic nature of ovarian endocrine function. The lack of response to gonadotropin with regard to E2 production prior to spawning is not due to insensitivity to gonadotropin, but rather due to some deficiency in steroidogenesis (e.g. reduced aromatase activity). It appears also that ovarian steroidogenesis and its regulation are dependent on annual changes of photothermal conditions.     

Migratory and reproductive behaviour of female adult Atlantic salmon, Salmo salar L., in a spawning stream

Migration and spawning behaviour of eight Atlantic salmon adult females were analysed by radio-tracking in relation to the degree of sexual maturity in a spawning tributary of the R. Sélune. Six of them were grilse and four of them were ripe. All the fish migrated upstream until reaching their spawning site at a distance of 4–12 km from the trap. The daily migration rate up to this site was inversely correlated with the length of the female. Spawning occurred in all fish at the same time when the water temperature increased dramatically. Spawning lasted 1–10 days according to the fish. After spawning, females quickly moved downstream only small distances and then stayed in approximately the same location until death. This study concluded that ripeness did not influence behaviour on the spawning migration and describes certain characteristics of the reproductive phase in a spawning tributary.     

Movement of a South American perch Percichthys trucha in a mountain Patagonian lake during spawning and prespawning periods

Movement of the South American perch Percichthys trucha (Perciformes: Percichthyidae) was studied in a set of interconnected mountain lakes in northern Patagonia. Fish were tracked using attached acoustic tags and a fixed hydrophone array over two seasons, corresponding to the spawning (1999) and prespawning periods (2000). Percichthys trucha was found to move throughout the lake system, and to use a small, shallow, connected lake as a principal spawning area. In addition, detailed behavioural information was obtained for the spawning period. Most fish spent the initial 2 weeks descending from the littoral zone to depths of 5–8 m, presumably while feeding. Fish changed depth slowly, spending several days at each depth, and moving throughout the lake. A few days prior to the full moon, there was a change in behaviour thought to be associated with spawning, where the fish moved toward the vegetated littoral areas. Marked daily behaviour also began at this time, with movement to particular near-shore locations at dusk, and return to slightly greater depths offshore at or before dawn. Study of the behaviour of P. trucha during the breeding season has highlighted the conservation significance of the main locations within the lake system.     

Factors affecting gastric evacuation in cod, Gadus morhua L., fed single-meals of natural prey

Functional relationships were derived in order to describe the effects of fish weight (262–2066 g), temperature (1–14° C), prey size and ration size (0.4–8.4% body weight) on gastric evacuation in cod fed four natural prey types. Interindividual (unexplained) differences among voluntary fed cod were relatively large. Power functions were appropriate for describing the effects of fish weight and ration size on the half-time of evacuation. Effects of meal and/or prey size appeared to counterbalance the effects of cod size. It is suggested that when cod of different sizes are fed fixed proportions of their body weight evacuation time is constant and independent of body size. There was found to be an exponential relationship between temperature and gastric evacuation time but, observations made at similar temperatures at different times of the year suggested that seasonal effects were of minor importance.     

Seasonal changes in the day and night activity of adult perch, Perca fluviatilis L.

Perca fluviatilis L. were caught by gill nets every 3 h over a 24 h period monthly from February 1975 until January 1976. The numbers caught increased with rise in temperature (correlation coefficient r = 0.86 P < 0.001). During the winter months perch migrated into deeper water although some fish were still caught in shallow water (1–6 m). Perch were active at very low light intensities. However changes in visual surface irradiance (400–700 nm) and water clarity caused inverse changes in activity. Only June did not conform to this pattern and it was presumed that spawning behaviour was of primary importance. It is suggested that increase in eutrophication with its effect on water clarity enables fish to become more active during the day in the summer months. They are thus able to utilise food sources to a maximum at an optimum period for feeding and growth.     

Reproduction in the greenback grey mullet, Liza subviridis (Valenciennes, 1836)

This paper presents data on the reproductive biology of Liza subviridis , a little studied mullet species. The fish is heterosexual, exhibiting external fertilization. Six maturity stages can be macroscopically identified in the testes and seven in the ovaries. The macroscopic changes in the gonads are manifestations of histological changes occurring in the development of sperm cells and oocytes. First sexual maturity is attained in the length ranges of 9.5–11.5 cm and 10.5–11.5 cm in male and female fish respectively. The fecundity for fish measuring 10.3–13.9 cm in standard length ranged from 40 000–145 000 eggs. The relationship between fecundity ( F ) and length ( L ) can be represented as: F = 1.9044 L ^4.2998. The spawning duration in L. subviridis is restricted to a short and definite period, with all ripe ova being released within a single spawning act. A pronounced spawning season can be detected to extend from June to November. However, during off-seasons, some spawning also occurs. The correlations between spawning, rainfall and air temperature is discussed.     

Comparative study of reproductive biology in single- and multiple-spawner cyprinid fish. I. Morphological and histological features

To clarify the dynamics and regulation of oogenesis in single- and multiple-spawning cyprinid fish with group-synchronous oocyte development, a multidisciplinary approach to their reproduction was undertaken using three species from the River Meuse (Belgium): the roach Rutilus rutilus as a single spawner, and the bleak Alburnus alburnus and the white bream Blicca bjoerkna as multiple spawners. The gonadosomatic index (GSI) and histomorphometric changes (distribution of oocyte size, relative proportion of the various oocyte stages) in the ovary are compared. Different patterns of GSI and oocyte growth were observed both between the single- and multiple-spawner fish and between the two multiple spawners. Maximum GSIs were higher in roach (21%) than in bleak and white bream (17.7 and 14.5%, respectively), and compared to the rapid decline of GSI in the roach population, the GSI of multiple spawners decreased progressively during the spawning season. In roach, a short gonadal quiescent period and an early onset of vitellogenesis was recorded from late summer onwards whereas, in bleak and white bream, exogenous vitellogenesis was not systematically observed before winter. A protracted spawning season and/or a low water temperature in autumn are hypothesized to explain this long period of gonadal quiescence. In bleak, during the spawning season, the oocytes recruited arose from the stock of endogenous vitellogenesis and attained the final maturation stage very rapidly. This recruitment occurred during the whole spawning season. In white bream, the differentiation of vitellogenic oocytes from smaller oocytes was completed before the onset of the spawning season. During the spawning period, the proportion of vitellogenic oocytes decreased progressively whereas the percentage of oocytes in the final maturation stage remained approximately constant.     

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

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

Climate effects on size‐at‐age: growth in warming waters compensates for earlier maturity in an exploited marine fish

Over the past 3 decades, North Sea Atlantic cod (Gadus morhua) have exhibited variable length‐at‐age along with declines in spawning stock biomass and timing of maturity. Multiple factors affecting growth and development in fish acted on this economically important stock over the same period including warming waters and an intensive fishery. Here, we employ North Sea cod as a model population, exploring how a physiologically relevant temperature metric (the growing degree‐day, GDD; °C day) can be used to compare year‐classes on a physiologically relevant time‐scale, disentangling influences of climate (thermal history) on observed length‐at‐age trends. We conclude that the trends in North Sea cod length‐at‐age observed during the last three decades can be explained by a combination of temperature‐dependent growth increases and a trend toward earlier maturation, the latter likely induced by the intensive fishing pressure, and possibly evidence of fisheries‐induced evolution.     

Coping with resource variation: Effect of constant and variable intervals between feeding on reproductive performance at first spawning of female three-spined sticklebacks

Female three-spined sticklebacks Gasterosteus aculeatus exposed to spring conditions (14) C and a photoperiod of 16L: 8D), and fed 6% of initial body weight per day of enchytraeid worms (controls) over 56 days, or either at regular 3-day or variable 1–5-day intervals between feeds, did not differ significantly in the proportion spawning (72%), in time to first spawning (24·5 days), in fecundity at first spawning (34) or in egg diameter (1·4 mm) at first spawning. There was no significant effect of feeding regime on lipid concentration of females immediately after first spawning (18·8% dry weight), but the experimental females had a slightly lower dry matter content and lower fresh and dry liver weight than controls. Eggs of females on the variable-interval regime had a higher lipid content than eggs from other groups. Days to spawning was correlated inversely and fecundity positively with initial length. Females adjusted their food intake to compensate for short periods of food deprivation, allowing them to maintain their reproductive performance at first spawning.     

The influence of photoperiod on oocyte growth and its role in the control of the reproductive cycle of the polychaete Harmothoe imbricata (L.)

The date of spawning of the first of the two batches of oocytes produced by Harmothoe imbricata each year is known to be influenced by both temperature and day length conditions during the preceeding period of vitellogenesis, acting directly on the rate of oocyte growth. The influence of day length is shown to be in the form of a critical day length effect, with days of 11 h of daylight or more acting as long days and advancing the date of spawning. It is suggested that this critical day length effect serves to synchronize the spawning of the first cohort of oocytes, and it may also entrain an endogenous time sense underlying the reproductive cycle of this species.     

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.