Reproductive potential of the eastern baltic cod <Emphasis Type="Italic">Gadus morhua callarias</Emphasis> L. population

Trends in interannual variation in maturation and spawning terms of various age cohorts in the Eastern Baltic cod population in 1997–2009 were studied. Specific features in the age structure of the mature population part that were established by the end of the first decade of the 21st century were clarified. The role of cod age cohorts in the current population reproduction was considered taking into account the data on cod recruitment and fecundity.     

The effect of growth rate,size, and season on oocyte development and maturity of Atlantic cod (Gadus morhua L.)

The effect of growth rate, body weight, age, and season on ovarian development and maturation was investigated for Atlantic cod (Gadus morhua L.) reared in the laboratory over 10 months, for each of two consecutive years, 1978–1980. Cod were also collected from the Gulf of St. Lawrence, the Scotian Shelf, Georges Bank, the Flemish Cap, and the N.E. Newfoundland Shelf.The state of maturity was recognized by oocyte size. Stage I oocytes did not vary in size with growth rate, season, age, or maturity, while the size of stage II oocytes was positively correlated with maturity and negatively correlated with maximum stage of development achieved. Ovarian wall thickness was positively correlated with age and maturity.The frequency distribution of stage I and II oocytes distinguished the state of maturation, with cod that would mature by the next spawning season having a minimum of 20% ($\text{x}\text{?}\text{= 42\%}$) of their oocytes at stage II or greater level of development.Maturing 3-yr-old cod had greater life specific growth rates than immature 3-yr-olds, but growth rates during the third year itself were not significantly different. A hypothesis of a three-part density-dependent mechanism controlling fecundity is postulated. Future reductions in partial recruitment and total fecundity are predicted for the Gulf of St. Lawrence cod stock based on calculated growth rates for Gulf cod in 1979.     

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.     

Sex‐specific differences in gonopore and gonadal growth trajectories in the brooding sea urchin,Abatus cavernosus (Spatangoida)

The heart urchin Abatus cavernosus shows sexual dimorphism characterized by the development of external brood pouches and the enlargement of gonopores in brooding females. Relationships between body size, gonopore size, and gonadal maturation in each sex were examined for inflection points using piecewise regression models (PRM). Opening of the gonopore occurred at 15.5 mm test length. Inflection points in the gonadal growth and gonopore diameter trajectories were clustered at smaller sizes in males (23 and 24.2 mm, respectively) than in females (25.1 and 25.9 mm), indicating sex‐specific differences in sexual maturation. Gonopore growth showed positive allometry at pre‐adult stages of development in both sexes, but isometry and negative allometry in adult females and males, respectively. Gonadal growth was initiated at smaller sizes and proceeded at a higher rate with increasing body size in males than in females. Identification of inflection points in gonopore and gonadal growth trajectories, using objective PRM, allows the determination of life stages and sexual maturation for individuals, thus providing a complementary tool for population studies.     

Population variation in thermal growth responses of juvenile Atlantic cod (<Emphasis Type="Italic">Gadus morhua</Emphasis> L.)

Controlled environment experiments were carried out to investigate thermal influences and population differences on growth of wild-caught juvenile Atlantic cod Gadus morhua L. from two regions of differing thermal regime off Scotland; the Clyde Sea on the west coast and St Andrews Bay on the east coast. Cod from the Clyde demonstrated significantly higher growth rates than cod from St Andrews. In both populations the growth rate was greater at 12°C than at 8°C. These population and temperature effects act to reinforce one another and it could therefore be predicted that the growth differences between the two areas in the wild should be even more pronounced. The results are consistent with the suggestion that cod may be locally adapted to their thermal environment.     

External and internal control of patterns of feeding, growth and gonadal function in a temperate zone anuran, the toad Bufo bufo

Young toads were brought into the laboratory before hibernation in the autumn and exposed to alternating periods of low temperature and high temperature and feeding, thus simulating climatic cycles. In newly metamorphosed, non-hibernating toads, food intake was initially low and growth was stagnant, but subsequently food intake and lean growth increased strongly. Presumably, growth was initiated independently of feeding. In one-year-old, non-hibernating toads, food intake and growth declined with time, but high rates of feeding and growth were restored in the toads exposed to simulated hibernation. In both metamorphosed toads and one-year-old, hibernating toads, a period of predominant fat deposition succeeded predominant lean growth before food intake again declined to low levels. Initiation of sexual maturation in the female toads coincided with suspended growth, whereas sexual maturation in males was not correlated with changes in growth rate. In the female, but not the male toads, sexual maturation was greatly advanced under the experimental conditions, resulting in adult female sizes far below those typical of the wild population.     

Aspects of reproductive biology of wild-caught polar cod (<Emphasis Type="Italic">Boreogadus saida</Emphasis>) from Svalbard waters

Polar cod (Boreogadus saida) is considered a key species in the Arctic marine ecosystems. Yet detailed or even basic knowledge regarding its biology and adaptations, especially during the polar night, is in many cases poor. Data are presently unavailable in Western literature on the gonad development of polar cod and its reproductive biology in wild specimens. Accordingly, gonad development of wild-caught polar cod from fjords of the Svalbard archipelago was studied across seasons (April, August, September, November and January). Histological analyses of polar cod showed strong indication of a group-synchronous oocyte development with determinate fecundity and iteroparity. Females started gonadal development prior to April and had not yet reached the final stage of maturation in January. Testes matured more rapidly, with males ready to spawn in January. Furthermore, our data show that polar cod were able to reach sexual maturity at age 1+. Based on our data and previous reports, we hypothesise that polar cod is a total spawner.     

Countergradient variation in behaviour,growth and lipid storage in Atlantic cod; environmental and genetic effects

We housed offspring from northern (70° N) and southern (60° N) coastal cod (Gadus morhua) together in a ‘common garden’ rearing experiment at a temperature and light regime representative of the southern population. Through a more active feeding behaviour and a higher success, the northern cod achieved a larger food share and a higher growth rate and condition than their southern conspecifics. This is contrary to what was demonstrated by field data of fish from their natural habitats. The northern cod also allocated more energy to the liver throughout the experiment. Our results agree with the theory of countergradient variation, suggesting that genetic influences on growth and condition have been opposed by environmental constraints in their natural habitat. The observation that the offspring from these populations differ in behavior and growth when housed together support the idea that the growth response to selection would be through a behavioral response. The field data suggest that density‐dependent population process and high juvenile density relative to prey limit the growth and condition in the wild and not necessarily the length of the growth season per se as assumed in the literature. The topographic distance (over 2000 km) limit mixing of early life stages of cod from the northern and southern population, and the different environmental stimuli (seasonality, temperature, food‐web interactions and habitat heterogeneity) in north and south are likely to evolve genetic differences.     

Haemoglobin polymorphism inGadus morhua: Genotypic differences in maturing age and within-season gonad maturation

276 specimens of Atlantic cod (Gadus morhua L.) were caught during spawning in a restricted area of the Trondheimsfjord, Norway, in April and May 1979. Genotypes at the polymorphic haemoglobin locusHbl differed significantly with respect to mean age at maturation (in males) and mean gonadic development stage (in females). There was no indication of population mixing in the genotypic composition atHbl or at any of the 4 polymorphic tissue enzyme loci investigated (LDH-3, IDH-1, PGM, andPGI-1. The findings obtained were considered with regard to temperature-related differences in the functional properties of Hbl molecules, and genotypic differences in growth, age at maturation, and fishing mortality. At the present stage of investigation, the natural selection pattern seems directional and strong. However, theHbl allele frequencies observed in cod from the examined areas reveal no detectable changes over a period of two decades (～ 4 generations). The current pattern of commercial exploitation causes, through size selection, a modification of the rate of erosion of the inferior allele, but additional factors must be in force, which play a role in its current abundance in an evolutionary perspective. The observedHbl genotypic differences in the exact within-season time for spawning might be one such factor. A potential sexual difference in genotypic fitness might be another, but this has yet to be confirmed. The apparent existence of considerable natural and artificial selection forces acting upon cod haemoglobin genotypes makesHbl allele frequencies unreliable for use in population structure analyses.     

Specific growth rate and proximate body composition of Atlantic cod (Gadus morhua L.)

The effect of growth rate and maturation on the proximate composition and energy content of Atlantic cod, Gadus morhua L., was investigated over 10 months for each of two consecutive years, 1978–1980 at 5 and 8 °C. Relative energy and lipid content of whole cod increased with specific growth rate for all three sampling periods (November, January, March), each at 5 and 8 °C. Relative water content decreased with specific growth rate and temperature, and was lower in March than in January and November. Relative protein content was positively correlated with specific growth rate, but to a lesser degree than with temperature and age. Relative ash content was negatively correlated with specific growth rate. The effect of season and temperature on the proximate content of gonad, liver, muscle, and carcass was also determined. The major energy and lipid source in cod was the liver. Energy, lipid, and water were highly correlated to each other, and regressions are provided to allow for their prediction, given one of the components. Energy budgets for cod at 5 and 8 °C are calculated and the effect of increased ration size on the budget is estimated. The prediction of short-term specific growth rates of cod from the proximate composition is proposed. The proximate composition of cod is affected by growth rate and thus feeding level, and in turn directly affects behaviour. The relative proximate content of maturing and immature 3-yr-old cod was not found to be significantly different. Keywords: specific growth rate; proximate; bioenergetics; Atlantic cod; energy budget; temperature; season     

ECOLOGY AND NATURAL HISTORY OF IRIDAEA CORDATA (GIGARTINALES,RHODOPHYTA) GROWTH1

The in situ growth, maturation and senescence, of central California Iridaea cordata (Turner) Bory was studied. This ontogenetic progression was quantified by measuring development and growth of: i) individually tagged blades, and ii) populations of blades within cumulative (1-yr duration) and seasonal experimental plots. Greatest growth and longest life span were exhibited by winter-spring initiated blades, and were correlated with a rapid increase, in irradiance, but not with either seawater temperature or nutrients. Tagging studies showed that reproductive maturation and senescence of blades occurred throughout the year, irrespective of date tagged, growth rate or size. Moreover, the majority of blades continued to elongate following maturation, and some matured within 3 mo of initiation at all seasons but winter. At the population level maturation took place primarily during summer-autumn when 90 ±2% of the population was mature. The majority of the population senesces or “dies back” during autumn-winter. It is concluded that in situ, the blades are derived almost totally via vegetative means involving perennation. This indirectly suggests that sexual reproduction or the success of sporeling development are marginal. Additionally, the species is perennial with annually deciduous blades, characterized by both rapid growth and maturation.     

Biology of the minnow Phoxinus phoxinus and its interactions with brown trout Salmo trutta in Øvre Heimdalsvatn, Norway

Minnows have recently been introduced into the mountain lakes of southern Norway. In Øvre Heimdalsvatn, an intensively studied lake, the minnow population has increased rapidly during the last decade. Studies were performed on feeding habits, habitat preferences, age, growth, energy content, densities, parasites, predation and spawning activities. Several aspects were found to be different from other studies, e.g. higher age of sexual maturation, lower growth rate, no feeding on plants, and low infection of parasites. The interaction with the only other fish, the brown trout, is discussed in relation to the minnow population. In 1978 the minnows constituted little more than 10% of production and food consumption of the trout population, and only limited food competition and predation was documented. Increasing competition is likely with a greater minnow population.     

Substantial Downregulation of Myogenic Transcripts in Skeletal Muscle of Atlantic Cod during the Spawning Period

Gonadal maturation is an extremely energy consuming process for batch spawners and it is associated with a significant decrease in growth and seasonal deterioration in flesh quality. Our knowledge about the molecular mechanisms linking sexual maturation and muscle growth is still limited. In the present study, we performed RNA-Seq using 454 GS-FLX pyrosequencing in fast skeletal muscle sampled from two-year-old Atlantic cod (Gadus morhua) at representative time points throughout the reproductive cycle (August, March and May). In total, 126,937 good quality reads were obtained, with 546 nucleotide length and 52% GC content on average. RNA-Seq analysis using the CLC Genomics Workbench with the Atlantic cod reference UniGene cDNA data revealed 59,581 (46.9%) uniquely annotated reads. Pairwise comparison for expression levels identified 153 differentially expressed UniGenes between time points. Notably, we found a significant suppression of myh13 and myofibrillar gene isoforms in fast skeletal muscle during the spawning season. This study uncovered a large number of differentially expressed genes that may be influenced by gonadal maturation, thus representing a significant contribution to our limited understanding of the molecular mechanisms regulating muscle wasting and regeneration in batch spawners during their reproductive cycle.     

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.     

Stock-specific changes in growth rates, food conversion efficiencies, and energy allocation in response to temperature change in juvenile Atlantic cod

Under common environments, populations of laboratory reared (Grand Banks, GB and Gulf of Maine, GOM) and wild caught (Fortune Bay, FB and Bonavista Bay, BB; Newfoundland) juvenile cod Gadus morhua responded similarly to temperature change in specific growth rates, food conversion efficiencies, condition factors, liver water content, and muscle water content. However, GOM cod had higher condition factors, and showed differences from GB cod in phenotypic plasticity of hepatosomatic index to temperature. These differences were not present in a different population comparison between FB and BB cod. All populations had higher growth rates and food conversion efficiencies at warmer temperatures, and exhibited compensatory growth when temperature was increased. The results suggest relatively larger genetic differences between GB and GOM cod than between FB and BB cod, and indicate that the faster growth of southern populations in the wild is not due to a higher genetic capacity for growth rate.     

Temporal dynamics of oocyte development, plasma sex steroids and somatic energy reserves during seasonal ovarian maturation in captive Murray cod Maccullochella peelii peelii

The temporal dynamics of oocyte growth, plasma sex steroids and somatic energy stores were examined during a 12 month ovarian maturation cycle in captive Murray cod Maccullochella peelii peelii under simulated natural photothermal conditions. Ovarian function was found to be relatively uninhibited in captivity, with the exception that post-vitellogenic follicles failed to undergo final maturation, resulting in widespread pre-ovulatory atresia. Seasonal patterns of oocyte growth were characterised by cortical alveoli accumulation in March, deposition of lipids in April, and vitellogenesis between May and September. Two distinct batches of vitellogenic oocytes were found in Murray cod ovaries, indicating a capacity for multiple spawns. Plasma profiles of 17β-oestradiol and testosterone were both highly variable during the maturation period suggesting that multiple roles exist for these steroids during different stages of oocyte growth. Condition factor, liver size and visceral fat stores were all found to increase prior to, or during the peak phase of vitellogenic growth. Murray cod appear to strategically utilise episodes of high feeding activity to accrue energy reserves early in the reproductive cycle prior to its deployment during periods of rapid ovarian growth.     

Reproductive potential of the Eastern Baltic cod Gadus morhua callarias L. population

Trends in interannual variation in maturation and spawning terms of various age cohorts in the Eastern Baltic cod population in 1997-2009 were studied. Specific features in the age structure of the mature population part that were established by the end of the first decade of the 21st century were clarified. The role of cod age cohorts in the current population reproduction was considered taking into account the data on cod recruitment and fecundity.     

Dynamics of the fecal corticosterone content in males of red,gray-sided,and bank voles (<Emphasis Type="Italic">Myodes</Emphasis>, Rodentia,Cricetidae) upon sexual maturation

The dynamics of the corticosterone content in feces of males are analyzed in red (M. rutilis), graysided (M. rufocanus), and bank (M. glareolus) voles. The ontogenetic dynamics of the corticosterone content in feces of these species collected on the 20th and 40th days are shown to depend differently on the month of their birth. At the same time, the fecal corticosterone content is similar in males of all species that originated from litters with various sizes and shares of males in it. The fecal corticosterone content in the 40-day-old animals is related to the month of birth for all three species. The species-specific features of adrenal activity are found on the 20 and 40 days after the birth of animals. The males of the May and August generations have the highest corticosterone level in feces. The fecal corticosterone content in the red vole males also correlates with the social environs; in addition, socially isolated single males have a higher rate of maturation. The fecal corticosterone in the gray-sided vole males related to the season of start maturation and to the date of birth negatively correlates with sexual maturation. The mature males of those species are found only among the spring–early summer generation. Thus, population factors are important only for maturing males that were born in the current year. Moreover, sexual maturation at a high population density is accompanied by a smaller decrease in the adrenocortical activity.     

Sex‐specific plasticity and genotype × sex interactions for age and size of maturity in the sheepshead swordtail,Xiphophorus birchmanni

Responses to sexually antagonistic selection are thought to be constrained by the shared genetic architecture of homologous male and female traits. Accordingly, adaptive sexual dimorphism depends on mechanisms such as genotype‐by‐sex interaction (G×S) and sex‐specific plasticity to alleviate this constraint. We tested these mechanisms in a population of Xiphophorus birchmanni (sheepshead swordtail), where the intensity of male competition is expected to mediate intersexual conflict over age and size at maturity. Combining quantitative genetics with density manipulations and analysis of sex ratio variation, we confirm that maturation traits are dimorphic and heritable, but also subject to large G×S. Although cross‐sex genetic correlations are close to zero, suggesting sex‐linked genes with important effects on growth and maturation are likely segregating in this population, we found less evidence of sex‐specific adaptive plasticity. At high density, there was a weak trend towards later and smaller maturation in both sexes. Effects of sex ratio were stronger and putatively adaptive in males but not in females. Males delay maturation in the presence of mature rivals, resulting in larger adult size with subsequent benefit to competitive ability. However, females also delay maturation in male‐biased groups, incurring a loss of reproductive lifespan without apparent benefit. Thus, in highly competitive environments, female fitness may be limited by the lack of sex‐specific plasticity. More generally, assuming that selection does act antagonistically on male and female maturation traits in the wild, our results demonstrate that genetic architecture of homologous traits can ease a major constraint on the evolution of adaptive dimorphism.     

Faster or slower: has growth of eastern Baltic cod changed?

Recent environmental changes have influenced the ecology and biology of eastern Baltic cod. Declining somatic condition, maturation at smaller size and restricted size distribution of the population suggest that growth rates have decreased between the early 2000s and the 2010s. Extensive age estimation problems have until now precluded testing of this hypothesis. This study presents evidence for a decrease in somatic growth rate of Baltic cod. Temporal patterns of growth, condition and maturation were analysed based on two complementary analyses: length frequency mode progression derived from DATRAS bottom trawl survey data and known-age samples, where size at age was back-calculated from daily otolith growth patterns. In the known-age samples, growth was positively related to somatic condition at capture with maturity dependent differences. Immature individuals had experienced significantly lower growth and were in lower condition at capture than mature individuals. Growth rates in the known-age samples were estimated at 9.5, 7.8 and 5.7?cm per year for age classes 1, 2 and 3 respectively. Growth between age 2 and 3 decreased significantly from 8.8?cm in the 1997 year class to 7.6?cm in the 2010 year class. While the 2001 and 2004 known-age samples were representative for the population, the 2013 sample was biased towards individuals with a higher condition and growth. Complementary length frequency analysis following the length mode of fish from age 2 to age 3 confirmed growth estimates from the early 2000s, while suggesting a 37.5% lower growth in 2013 compared with 2005.