Adaptive winter survival strategies: defended energy levels in juvenile Atlantic salmon along a latitudinal gradient

Current knowledge suggests that patterns of energy storage and depletion in animals are governed by behavioural trade-offs between risks associated with feeding and future energy demands. However, the length of adverse periods varies over geographical or climatic gradients. To explore the potential for genotypic sources of variation in behavioural trade-offs, we compared the winter energy-depletion patterns among 13 wild populations of juvenile Atlantic salmon (Salmo salar L.) along a latitudinal gradient (58–70°N) and performed common-environment experiments of energy-state-dependent feeding. In the wild, winter lipid-depletion rates were lower for northern than for southern populations. The variation in spring lipid levels among the population was lower than autumn variation, with storage lipid levels clustered close to critical limits for survival. In semi-natural stream channels with natural food supply, hatchery-reared fish originating from northern populations showed a positive scaling of feeding activity with decreasing energy levels, whereas southern populations did not. In conclusion, juvenile Atlantic salmon from northern populations defend their energy levels more strongly than fish from southern populations. Adaptive variation in feeding activity appears important for this difference. Thus, the present study shows a link between geographical patterns in storage energy trajectories and adaptive differences in state-dependent feeding motivation.     

The allometry of energy reserve depletion: test of a mechanism for size-dependent winter mortality

We experimentally tested the hypothesis that energy reserve depletion varies inversely with size in the fish Menidia menidia, an estuarine fish known to exhibit size-dependent winter mortality. Individuals in two size groups were starved at two winter temperatures (4°and 8°C) and sacrificed at a range of time intervals (up to 127 days). Lipid levels and lean tissue were analyzed to estimate somatic energy storage. As predicted, energy depletion was greater at high temperatures, and proportionally greater in small than in large fish. After 60 days of starvation at 4°C, small fish retained an average of 67% of their original energy reserves (vs 53% at 8°C), while large fish retained an average of 80% (vs 66% at 8°C). At 4°C, fish that were fed depleted their energy reserves as rapidly as unfed fish, but at 8°C, fish that were fed maintained reserves at higher levels than unfed fish. A high proportion of unfed fish (56% at 4°C, 27% at 8°C) died before they were to be sacrificed. Survival probability did not vary with size, nor was it influenced by the amount of energy reserves. The rate of energy depletion (equivalent to routine metabolic rate) decreased gradually over time, particularly in small fish. Routine metabolism did not conform to a single scaling relationship. Within each temperature-size group, the routine rate declined more rapidly than metabolically active mass (lean mass). At 8°C, the difference between size groups in energy depletion rate conformed closely to the expected allometry exponent of 0.8. In contrast, at 4°C, the estimated allometry exponent increased over the experiment (−0.19 to 2.5). We conclude that strategies to minimize energy loss may often modify bioenergetic scaling relationships. Received: 30 September 1998 / Accepted: 10 February 1999     

Winter energy allocation and deficit of juvenile walleye pollock <Emphasis Type="Italic">Theragra chalcogramma</Emphasis> in the Doto area,northern Japan

Seasonal energy allocation and deficits of marine juvenile fishes have considerable effects on their survival. To explore the winter survival mechanism of marine fishes with low lipid reserves in their early life, juvenile walleye pollock Theragra chalcogramma were collected along the continental shelf of northern Japan over a 2-year period, and energy allocation and deficit patterns were compared between wild and laboratory-starved fish. Contrary to expectations, wild fish generally continued to accumulate protein mass and concurrently tended to reduce lipid mass from late autumn through winter. The most plausible explanation for the continuous structural growth is that juvenile pollock give priority to reducing mortality risk from size-selective predators under quasi-prey-limited conditions. Exceptionally, inshore small fish reduced both constituents during a winter. The inshore fish consumed 2.5 times more lipid energy than protein energy in November–December, but protein was more important than lipids as a source of energy in December–January and in February–March. However, dependence upon protein reserves was lower for the wild fish than for the laboratory-starved fish, suggesting milder nutritional stress of the wild fish than that observed in the starvation experiment. Moreover, the lipid contents of mortalities in the starvation experiment were mostly <1%, whereas few wild fish had such lipid contents in the field. These results suggest that juvenile pollock are able to avoid both starvation and predation by accumulating protein reserves.     

Rate of energy depletion and overwintering mortality of juvenile walleye pollock in cold water

The winter energy deficit and mortality of juvenile walleye pollock at extremely cold temperature were examined by field observations and laboratory experiments. In the Doto area, along the northern coast of Japan, juvenile walleye pollock resided on the continental shelf despite extremely cold temperatures (mean 0·4° C) during the latter half of winter (March to April). Measurements of the rate of energy depletion (equivalent to the routine metabolic rate) revealed that juvenile walleye pollock consumed 37% less energy at 0·5° C than at 2·0° C, suggesting an energetic benefit of residence in cold water (<1·0° C) over the shelf during winter. Prior to the starvation experiments, temperatures and ration level in the holding tanks were adjusted to create two different body condition groups of fish. Under the thermal condition of the latter half of winter (0·5° C), fish with a mean condition factor of 0·6 and 0·5 suffered 19·1 and 74·5% mortality, respectively, at the end of the experiments (after 56 days). The residual analysis of total body energy demonstrated that the cause of mortality was mainly associated with the depletion of energy reserves. When a logistic regression model for mortality derived from the experiments was applied to wild fish collected in March, the estimated overwintering mortality in 2004 and 2005 was 25·4 and <2·3%, respectively, assuming no feeding during the winter. Considering that juvenile walleye pollock feed during winter as shown in previous studies, intense overwintering mortality induced by energy depletion is improbable during the latter half of winter in the Doto area.     

Energy acquisition and retention by age‐0 and age‐1 paddlefish Polyodon spathula (Walbaum, 1792) in relation to size,growth, and rearing conditions in two Great Plains reservoirs and hatchery ponds

The objective of this study was to investigate lipid accumulation and storage in age‐0 and age‐1 paddlefish Polyodon spathula (Walbaum, 1792) in relation to age, stock, year, and growth. Juvenile paddlefish were collected from three locations in North Dakota and Montana, USA, during July and August of 2011 and 2012 and proximate analysis was used to determine lipid content. RNA/DNA ratios were used as an index of growth rates. Differences in age‐based lipid accumulation and storage in juvenile paddlefish suggest a split allocation between growth and lipid storage, with growth being the highest initial priority and emphasis on energy storage occurring at a larger size, later in life. Differences in lipid allocation between stocks indicate that allocation is influenced by hatchery/wild rearing conditions. Differences within and between year‐classes are consistent with field evidence observed in 2012 of a strong 2011 year‐class, and indicate that during productive times, paddlefish may allocate energy to both body growth and lipid reserves, and that allocation differs among years. The lack of a relationship between RNA/DNA ratio and lipid does not support a physiologically exclusive allocation strategy between growth and lipid. Evidence from this and other studies suggests rather that an emphasis on growth, some energy storage, and a large rostrum size in relation to overall fish length in age‐0 and age‐1 fish, may be adaptive in avoiding predation while accruing necessary energy reserves for overwintering. Although this study also provides reference information regarding proximate composition of wild and hatchery origin juvenile paddlefish, much more study is needed into the relationships among growth, low and high lipid groups, lipid allocation in juvenile paddlefish as well as the existence and timing of allocation changes between growth and storage. To aid in understanding paddlefish survival and year‐class strengths, these relationships also need to be linked to inter‐annual differences in early rearing environments for age‐0 and age‐1 fish.     

Saving for the future: Pre‐winter uptake of algal lipids supports copepod egg production in spring

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Annual variation in energy reserves in male three-spined stickleback, Gasterosteus aculeatm L. (Pisces, Gasterosteidae)

Wet and dry weights of tissue were measured and concentrations of glycogen, lipid and protein were estimated for the liver, gonad and carcass of male sticklebacks from an annual population collected each month over one complete year. In young-of-the-year there is one period of rapid weight gain, in all three body regions (liver, carcass and gonads) but particularly of the carcass, in the autumn and a second in spring and early summer. This is accompanied by an increase in the water content of all three body regions. The gonadosomatic index also increases sharply in spring and early summer. Young sticklebacks accumulate lipid and glycogen slowly during the autumn and winter of their first year of life and more rapidly from late winter to early summer. Thus, the period of most rapid accumulation of these reserves coincides with the time when body weight and gonad maturation are also increasing sharply. Lipid and glycogen levels fall during the reproductive season in those males that breed, so that by July they are reduced to 43% and 37% (respectively) of their peak values in May. Levels of protein increase throughout the year as the fish grow, but in breeding males by July the concentration of protein in the carcass falls to 70% of pre-breeding levels. Breeding males therefore reach the end of the reproductive season with their total energy reserves severely reduced, and consequently they suffer a very high mortality. In contrast, adult males that fail to reproduce survive beyond the breeding season. They continue to gain weight and to accumulate lipid and glycogen from August to September, but these energy reserves fall (to levels comparable to those of post-breeding fish) in December, when these fish probably die. These results demonstrate that in male sticklebacks, growth and gonad maturation can be sustained in parallel with the accumulation of energy reserves, which are subsequently extensively depleted as a result of reproductive activities.     

The influence of life-history strategy on lipid metabolism in overwintering juvenile Atlantic salmon

From November to May, the lipid mass in the viscera and carcass of juvenile Atlantic salmon Salmo salar that were undergoing smolt transformation prior to seaward migration ('early migrants') were significantly greater than those of their siblings that would delay migration for at least a further year. During winter (November-February), the depletion of lipid associated with the viscera was significantly greater in early migrants, whilst lipid depletion in the remaining carcass was greater in delayed migrants. Early migrants continued to deplete both lipid compartments in spring (February-May), whereas delayed migrants depleted visceral lipid but replenished carcass lipid over the same period. Fatty acid accumulation rates (a measure of storage lipid synthesis rates) were two to six times greater in visceral than in carcass lipid throughout the study, suggesting that lipid turnover is much more rapid in the viscera. There were no differences in fatty acid accumulation rates between migrant groups in November, despite the much lower food consumption rate of delayed migrants at that time, suggesting that these fish allocated a larger proportion of their nutritional resources to lipid synthesis. In the carcass lipid of early migrants, and in both the visceral and carcass lipid of delayed migrants, the fatty acid accumulation rate was negatively correlated with lipid mass. Fatty acid accumulation rates increased from November to February in both visceral and carcass lipid in the two migrant groups. The fatty acid accumulation rate in carcass lipid was significantly higher in delayed migrants than in early migrants in February, but not in May. These results support the hypothesis that life history strategies involving rapid growth will result in a relatively low allocation of resources to lipid reserves.     

Growth, maturation and reproductive investment in Arctic charr

Size and rates of growth in a cohort of 1 + Arctic charr housed in standard conditions were tracked over 12 months (December to December) and mature and immature males and females compared retrospectively. In both sexes, maturing fish were larger than non-maturing ones. In males, this size differential was the result of differences in growth in winter and early spring, but not in the remainder of the study period. In females, size differentials resulted mainly from growth rate differences immediately prior to breeding. In females but not in males, gonadosomatic index was predicted by growth rates in the months leading up to maturation, and among the females that matured, faster growing fish produced more eggs. Lipid reserves in July were correlated negatively with growth during the previous 7 months and, in females only, lipid reserves were significantly lower in maturing fish than in non-maturing fish, indicating that mobilization of lipid energy reserves in maturing fish had commenced by this time. Variation in investment in gonadal tissue, measured as gonadosomatic index, was not explained by variation in July lipid reserves for either males or females. However, July lipid reserves were negatively correlated with egg number, so females investing more in ova exhibited greater depletion of lipid reserves. These results are discussed in the context of the relationship between body condition and the onset of maturation in salmonids, relative investment in reproduction and sexual differences in the cost of reproduction.     

Comparative energy allocation in two sympatric, closely related gobies: the black goby Gobius niger and the grass goby Zosterisessor ophiocephalus

Seasonal energy allocation of lipid reserves into different body tissues was analysed comparatively in two sympatric, closely related gobies: the grass goby Zosterisessor ophiocephalus and the black goby Gobius niger . Lipid reserves were measured in liver, muscle and ovary and compared between the two species within a given sex and seasonal period (reproductive v . non-reproductive). Furthermore, temporal patterns of lipid reserves were investigated in the two species in relation to gonado-somatic and liver-somatic indices, as well as the relationship between size and lipid content. Results showed that the grass goby allocated more lipid reserves in reproduction while the black goby accumulated more reserves in liver and muscle, at a given size, although the temporal patterns of lipid accumulation and depletion were basically similar. Results are discussed in the light of life-history theories, taking into account both adaptation and evolutionary constraints.     

Male reproductive strategy and reserve allocation in sand smelt from brackish lagoons of southern France

In male sand smelt Atherina boyeri , from three coastal brackish lagoons of southern France, the duration of the reproductive season differed according to fish size: larger fish exhibited an extended reproductive season from February to August while smaller males reproduced only from April to June-July. Smaller and larger fish showed differences in their seasonal pattern of reserves. Through the cold season, energy content was exhausted for over-winter maintenance and reproductive needs, and was restored just after fish ceased reproducing. In smaller fish, however, reserves were more rapidly restored than in larger fish, and exhibited a decrease in autumn. This result suggested that small fish invested their energy into growth and winter survival rather than reproduction while older fish invested in reproduction. These results are discussed in terms of life history Strategies.     

Seasonal variations in the physiological status and energy content of somatic and reproductive tissues of chub

The seasonal fluctuations in condition, nutrition and somatic energy content, and gonad development cycle were investigated in chub Leuciscus pyrenaicus from the headwaters of the Guadalete River, a freshwater ecosystem characterized by strong seasonal fluctuations in discharge and water level, temperature and food supply. The relationship between somatic stage and gonad cycle was also investigated and discussed. Condition, nutrition and somatic energy cycles could be divided into two distinct periods: from April to January with summer decreases and autumn increments, which is common for juveniles and mature fish; and from January to March, when juveniles and mature fish displayed different temporal variations related to the reproductive cycle. Gonad development took place from the end of the winter into the summer, the testes developing before the ovaries. Spawning started in late spring (May) and continued into summer (June and July), with fish quiescent by autumn (September). The results suggest that, for L. pyrenaicus , both environmental factors (e.g. food supply, water temperature) and reproduction needs affect the condition, nutrition and somatic energy storage of fish, which have been used as indicators of the physiological status of the population.     

Genetically based population divergence in overwintering energy mobilization in brook charr (Salvelinus fontinalis)

Investigating the nature of physiological traits potentially related to fitness is important towards a better understanding of how species and/or populations may respond to selective pressures imposed by contrasting environments. In northern species in particular, the ability to mobilize energy reserves to compensate for the low external energy intake during winter is crucial. However, the phenotypic and genetic bases of energy reserve accumulation and mobilization have rarely been investigated, especially pertaining to variation in strategy adopted by different populations. In the present study, we documented variation in several energy reserve variables and estimated their quantitative genetic basis to test the null hypothesis of no difference in variation at those traits among three strains of brook charr (Salvelinus fontinalis) and their reciprocal hybrids. Our results indicate that the strategy of winter energy preparation and mobilization was specific to each strain, whereby (1) domestic fish accumulated a higher amount of energy reserves before winter and kept accumulating liver glycogen during winter despite lower feeding; (2) Laval fish used liver glycogen and lipids during winter and experienced a significant decrease in condition factor; (3) Rupert fish had relatively little energy reserves accumulated at the end of fall and preferentially mobilized visceral fat during winter. Significant heritability for traits related to the accumulation and use of energy reserves was found in the domestic and Laval but not in the Rupert strain. Genetic and phenotypic correlations also varied among strains, which suggested population-specific genetic architecture underlying the expression of these traits. Hybrids showed limited evidence of non-additive effects. Overall, this study provides the first evidence of a genetically based—and likely adaptive—population-specific strategy for energy mobilization related to overwinter survival.     

Growth hormone-induced effects on mortality, energy status and growth: a field study on Brown Trout (Salmo trutta)

1. Growth hormone (GH) treatment increases the growth rate and competitive ability of salmonids under laboratory conditions. Since fast growth should increase fitness, why is endogenous secretion of GH not higher in wild fish? To address this question, three hypotheses were suggested. H₁: high GH levels reduce antipredator responses and may therefore increase mortality from predation. H₂: high GH levels reduce long-term (e.g. over winter) survival by reducing allocation to critical energy reserves. H₃: GH is not beneficial for growth under natural conditions.
2. To test these hypotheses, the performance of GH-treated juvenile Brown Trout ( Salmo trutta ) and control (placebo) trout was compared in an enclosed stream section subjected to natural predation. Four experiments were conducted during winter, spring, summer and autumn, respectively.
3. Mortality rates were not significantly different between GH-treated and control trout in any of the four experiments so H₁ was not supported. Energy reserves were generally lower in GH-treated fish, which is consistent with H₂, whereas growth rates in mass were higher in GH-treated fish than in controls so H₃ was not supported. This suggests that GH promotes growth at the expense of investment in maintenance.
4. Judging from growth and mortality rates, the fitness of GH-treated and control trout appeared similar. Thus, escaped GH-manipulated fish may compete successfully with wild fish.
5. Hatchery-raised trout with higher initial condition index suffered higher mortality rates than more slender fish. This novel finding may be explained by reduced escape ability related to body morphology, reduced behavioural responses towards predators by high-condition trout, or predator preferences for high-condition fish.     

Rhizome dynamics and resource storage in Phragmites australis

Seasonal changes in rhizome concentrations of total nonstructural carbohydrates (TNC), water soluble carbohydrates (WSC), and mineral nutrients (N, P and K) were monitored in two Phragmites australis stands in southern Sweden. Rhizome biomass, rhizome length per unit ground area, and specific weight (weight/ length ratio) of the rhizomes were monitored in one of the stands.Rhizome biomass decreased during spring, increased during summer and decreased during winter. However, changes in spring and summer were small (< 500 g DW m-2) compared to the mean rhizome biomass (approximately 3000 g DW m^–2). Winter losses were larger, approximately 1000 g DW m-2, and to a substantial extent involved structural biomass, indicating rhizome mortality. Seasonal changes in rhizome length per unit ground area revealed a rhizome mortality of about 30% during the winter period, and also indicated that an intensive period of formation of new rhizomes occurred in June.Rhizome concentrations of TNC and WSC decreased during the spring, when carbohydrates were translocated to support shoot growth. However, rhizome standing stock of TNC remained large (> 1000 g m^–2). Concentrations and standing stocks of mineral nutrients decreased during spring/ early summer and increased during summer/ fall. Only N, however, showed a pattern consistent with a spring depletion caused by translocation to shoots. This pattern indicates sufficient root uptake of P and K to support spring growth, and supports other evidence that N is generally the limiting mineral nutrient for Phragmites.The biomass data, as well as increased rhizome specific weight and TNC concentrations, clearly suggests that reloading of rhizomes with energy reserves starts in June, not towards the end of the growing season as has been suggested previously. This resource allocation strategy of Phragmites has consequences for vegetation management.Our data indicate that carbohydrate reserves are much larger than needed to support spring growth. We propose that large stores are needed to ensure establishment of spring shoots when deep water or stochastic environmental events, such as high rhizome mortality in winter or loss of spring shoots due to late season frost, increase the demand for reserves.     

Energy allocation strategy modifies growth–survival trade-offs in juvenile fish across ecological and environmental gradients

In young temperate zone fishes, conflicting energy demands lead to variability in growing season and winter survival. Growing season survival is driven by size-dependent predation risk whereas winter survival is constrained by autumn body size, energy storage and winter duration. We developed a model of the seasonality of energetics coupled to empirical measures of resource availability, size-dependent predation and temperature seasonality for rainbow trout (Oncorhynchus mykiss) in two sets of lakes in British Columbia, Canada, representing endpoints of a gradient of temperature, growing season duration and winter duration. This model was used to determine the energy allocation strategy which maximized first-year survival across these gradients. Survival was sensitive to the timing of the switch from somatic to storage strategies in cold, short growing season, low resource environments. A broader range of energy allocation strategies were viable in warmer, longer growing season and higher resource lakes. We used empirical observations of autumn energy storage and our modeled values for size-dependent minimal lipid levels needed to survive winter in each system to estimate winter survival for juvenile rainbow trout. Winter survival estimates were 6% in cold lakes with low resources, 82% in warm, lakes with low resources and 100% in warm lakes with high resources. Fish in warm lakes with ample resources allocated substantially more to storage than the minimum required to survive winter generated from our model, suggesting additional selection pressures for increased storage when there was ample surplus energy. We concluded that growth–survival trade-offs, modified by seasonality of the environment, influenced the growing season energy allocation strategies for young-of-the-year fish, and suggested this may be important for understanding population viability across environmental gradients.     

Distribution and population dynamics of Porphyra (Bangiales, Rhodophyta) in the southern Western Cape, South Africa

Although Porphyra is commercially farmed in many countries, in South Africa only small harvests of wild populations for sale as nori have been carried out. The discovery that Porphyra improves growth of South African abalone (Haliotis midae) farmed inland-based tanks has led to increased pressure to harvest wild populations. This paper reports on a survey of the distribution and seasonality of Porphyra in the southern Western Cape. Porphyrawas present at all sites surveyed, and showed considerable temporal variation. A significant amount of the Porphyra present is in reserves and therefore protected from harvesting. Close rexamination of one site revealed seasonal populations of Porphyra that occupied different niches dependent on season. Recruitment peaked in spring and autumn, leading to dense summer and winter populations. Summer populations generally grew lower in the eulittoral than winter populations. No pattern in the mortality of larger thalli wasde tected, though sporeling mortality was high following recruitment peaks. Although it seems that most sites in the southern Western Cape are suitable for harvesting, the taxonomy of the genus in the region urgently needs revision if populations are to be appropriately managed. This revised version was published online in August 2006 with corrections to the Cover Date.     

A 30% reduction in switchgrass rhizome reserves did not decrease biomass yield

A long-standing question in perennial grass breeding and physiology is whether yield improvement strategies could compromise winter survival. Since perennial grasses rely on stored carbohydrates for winter maintenance and spring regrowth, yield improvement strategies could reduce winter survival if they increase biomass and grain yields at the expense of carbon allocation to storage. Therefore, it is crucial to comprehend the dependence of regrowth on storage reserves. We experimentally depleted switchgrass (Panicum virgatum L.) rhizome reserves by storing rhizomes for 2 weeks at 5°C (control treatment) and 25°C (reserve-depleted treatment). During the storage period rhizome respiration was 5.3× higher at 25°C (0.010 μmol CO₂ g⁻¹ min⁻¹ at 5°C vs. 0.054 μmol CO₂ g⁻¹ min⁻¹ at 25°C; p < 0.0001) and the starch content was depleted by 30% by the end of storage. Surprisingly, reserve-depleted switchgrass had 60% larger leaf area (LA; LA_control = 149 cm² pot⁻¹ vs. LA_depleted = 239 cm² pot⁻¹; p = 0.013) and produced ~40% more aboveground biomass than control plants (9.46 g pot⁻¹ vs. 6.63 g pot⁻¹; p = 0.112). In addition, reserve-depleted switchgrass restored its rhizome starch reserves to pre-storage levels. Switchgrass showed a large plasticity among its source-sink components to buffer the imposed reserve depletion. It increased plant photosynthesis by increasing the photosynthetic leaf area while keeping photosynthesis constant on a leaf area basis and readjusted the timing and activity of sink organs. These results suggest that switchgrass, and potentially other perennial grasses, largely over-invest in storage reserves. Therefore, current breeding strategies in perennial grasses aimed to extend the aboveground growing season should not compromise crop persistence. Our study also has implications on long-term yield dynamics as it highlights sink limitations as potential driver of the yield decline commonly observed in perennial grasses 5+ years after cultivation.     

Biphasic growth in fish II: empirical assessment

In [Quince, et al., 2008. Biphasic growth in fish I: Theoretical foundations. J. Theor. Biol., doi:10.1016/j.jtbi.2008.05.029], we developed a set of biphasic somatic growth models, where maturation is accompanied by a deceleration of growth due to allocation of energy to reproduction. Here, we use growth data from both hatchery-raised and wild populations of a large freshwater fish (lake trout, Salvelinus namaycush) to test these models. We show that a generic biphasic model provides a better fit to these data than the von Bertalanffy model. We show that the observed deceleration of somatic growth in females varies directly with gonad weight at spawning, with observed egg volumes roughly 50% of the egg volumes predicted under the unrealistic assumption of perfectly efficient energy transfer from somatic lipids to egg lipids. We develop a Bayesian procedure to jointly fit a biphasic model to observed growth and maturity data. We show that two variants of the generic biphasic model, both of which assume that annual allocation to reproduction is adjusted to maximise lifetime reproductive output, provide complementary fits to wild population data: maturation time and early adult growth are best described by a model with no constraints on annual reproductive investment, while the growth of older fish is best described by a model that is constrained so that the ratio of gonad size to somatic weight (g) is fixed. This behaviour is consistent with the additional observation that g increases with size and age among younger, smaller breeding females but reaches a plateau among older, larger females. We then fit both of these optimal models to growth and maturation data from nineteen wild populations to generate population-specific estimates of ‘adapted mortality’ rate: the adult mortality consistent with observed growth and maturation schedules, given that both schedules are adapted to maximise lifetime reproductive output. We show that these estimates are strongly correlated with independent estimates of the adult mortality experienced by these populations.     

Seasonal changes in condition, nutrition, gonad maturation and energy content in barbel, Barbus sclateri, inhabiting a fluctuating river

The objective of this study was to investigate the seasonal cycle of condition, nutrition and gonad development, as well as the magnitude of seasonal variations in energy content of somatic and gonad tissues in juveniles, males and females of Sclater's barbel, Barbus sclateri, from the upper Guadalete River (south Spain). The influence of reproductive cycle on somatic changes was also investigated and discussed. Measurements of condition factor (K), somatic index (SI), gonadosomatic index (GSI) and somatic and gonad energy content (J g^-1 dw) were made in individual specimens taken from the Guadalete River monthly for 12 months. This freshwater ecosystem is characterised by strong seasonal fluctuations in water and flow level, temperature and food supply. It was found that in general juveniles, males and females of barbel exhibited a similar condition, nutrition and somatic energy cycle throughout the year, with the highest values in spring and the lowest in summer. Depletion of K, SI and somatic energy storage from spring until mid-summer seems to be associated with high metabolic demands during this period, and in adult fish also with spawning-related activity. The gonad index and energy content of the gonad were the highest in April and the lowest in summer for both sexes. Spawning took place during late spring – early summer, with fish quiescent by mid summer. The energy required for ovarian development (3970 J g^-1 dw) was greater than the one for testes development (2763 J g^-1 dw). Data on gonad energy content indicated a period (March to April) of intense energy accumulation (64% males and 37% females) which was related to the decline in the average somatic energy content in males and females. The somatic energy content was linearly related to K and SI. In the same way, GSI correlated positively with gonad energy. Linear trends were found between somatic parameters (K, SI and energy content) and gonad parameters (GSI and energy content). This revised version was published online in July 2006 with corrections to the Cover Date.