Organismal stoichiometry at the temporal scale: Seasonal variability shapes interspecific differences in fish

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Changes in Lipid Composition in Amur Sleeper <Emphasis Type="Italic">Percottus glenii</Emphasis> (Dybowski, 1877) Yearlings Depending on Body Length in the Floodplain Lake Krugloye (Saratov Reservoir)

Changes in the total content and composition of lipids in yearlings of the Lake Krugloye Amur sleeper depending on their body length during the active feeding period have been analyzed. It is found that fatness of the juvenile Amur sleeper does not change with an increasing body length. The seasonal dynamics of fatness is expressed to a greater degree in fish of the same size. The lipid composition of juvenile specimens of the Amur sleeper varies considerably depending on the length of the body of the fry and the time of observation. The character of the fractional composition of lipids in yearlings of the Amur sleeper with body lengths >25–29 mm in September and October indicates a decline in food supply of the largest specimens of the generation by the end of the active feeding period and their lower readiness for wintering.     

Temporal changes in the relationship between condition indices and proximate composition of juvenile Coregonus artedi

The Fulton‐condition factor ( K ) and per cent whole‐body water content were examined to determine whether these indices can estimate the proximate composition of juvenile lake herring Coregonus artedi exposed to a simulated Lake Superior winter over a 225 day laboratory experiment. The K was positively correlated to whole‐body crude lipid, crude protein, and gross energy content and negatively correlated to whole‐body water content for each sampling period of the experiment (days 75, 150 and 225). In contrast, there was only a weak positive correlation between K and whole‐body ash content. While per cent water content was negatively correlated with crude lipid, crude protein and gross energy content for each of the three sampling periods, the correlation between this predictor and ash content was only weakly negative. The indices can be used to accurately estimate temporal changes in proximate composition of juvenile lake herring during winter periods.     

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.     

Non-selective predator – the versatile diet of Amur sleeper (Perccottus glenii Dybowski, 1877) in the Vistula River (Poland), a newly invaded ecosystem

The Amur sleeper diet spectrum was investigated in addition to its spatial, seasonal and size-related changes, in order to predict which groups of native prey would be most affected by this exotic predator that has rapidly invaded many Eastern and Central European inland waters in recent decades. In total, 527 Perccottus glenii individuals were collected in four sites in the Włocławski Reservoir (the Vistula River, Baltic basin, Poland). Altogether 50 food categories were identified, including crustaceans, insects, molluscs, annelids, araneids, fishes and amphibians. Most of them were associated with aquatic vegetation, indicating that the Amur sleeper collects its prey from the submerged plants rather than directly from the bottom sediments. Amphipods and chironomid larvae, supplemented by zygopteran larvae and molluscs, composed the main forage base in all studied sites, although their importance was varied. The diet composition differed among P. glenii size-groups. Significant variation was found in numeric abundance of amphipods, chironomid larvae, fish and zygopteran larvae (Kruskal–Wallis P < 0.01). Seasonal changes in the diet were also observed whereby in spring and summer, dipteran larvae and amphipods were prevalent food items; the importance of molluscs and zygopteran larvae increased considerably in September; and fish showed the highest occurrence in June, July and August. The broad diet spectrum of the Amur sleeper indicates that it is a non-selective, opportunistic predator and that several taxonomic groups of native hydrofauna as well as macroinvetebrates and fish may be affected by its presence. The highly flexible feeding strategy undoubtedly favours Amur sleeper expansion in invaded watersheds.     

Protein, lipid, water and caloric contents of immature rainbow trout, Salmo gairdneri Richardson, growing at different rates

Protein, lipid, water and caloric contents of immature rainbow trout, relative to size of whole fish, growing at different rates were examined by use of allometric analysis (y = ax^b, Huxley, 1932). Fish grew at different rates as a result of differences in ration size (satiation, or 4–5% of dry body weight), temperature (7 and 12°C) and bGH (bovine growth hormone) administration. In fingerlings, protein, lipid and caloric contents tended to increase (v. body weight) as a percentage of body composition, whereas above fingerling size, protein decreased while lipid and caloric contents still increased. These trends occurred regardless of growth rate differences. The correlations between protein, lipid, caloric contents and body weight were high so reliable estimates of body components can be made from body weight for all experimental treatments. At satiation rations (7 and 12°C), there were no significant differences in protein content, but lipid and caloric contents were significantly higher in control fish. At low rations, protein and caloric contents were lower than those at satiation rations and lipid was lower than in the control group. On a dry weight basis, in uninjected fish at 12°C, ration size did not influence the percentage composition (protein and lipid) but the low ration group had lower energy values per unit of body dry weight. At low temperature (7°C satiation), fish had lower lipid and higher protein content (dry weight basis) than their controls (12°C satiation), thereby resembling bGH injected fish at satiation rations. Values of caloric content estimated from protein and lipid values by use of standard conversion factors differed sufficiently from caloric values directly determined by bomb calorimetry to suggest that caloric values of lipids may change during growth. The increase in body caloric content during growth apparently results mainly from an increase in the percentage of lipid. The similarity of body composition between different growth rate groups implied a tendency for conservation of relative proportions of components such that body composition can be approximated from body weight. Equations were also given to describe the relationship between water content and body constituents, and the relationship between condition factor (K) and both the body dry weight and lipid content.     

Lipid acquisition and retention in tissues of spawning adult paddlefish Polyodon spathula (Walbaum, 1792) in relation to extended and compressed life history patterns in two river‐reservoir systems

Data from recreationally caught adult paddlefish from North Dakota and Oklahoma, USA, was used in an initial attempt to explain variation in lipid content in adult wild paddlefish Polyodon spathula (Walbaum, 1792) by stock and by sex. The objectives of this study were to describe the relative importance of three distinct paddlefish tissues [white muscle, red muscle, and gonadal fat body (GFB)] for lipid storage, and compare lipid content of these tissues between stock and sex. It was hypothesized that lipid content of the three tissue types would be related to each other within individual fish; and that, consistent with previously documented life history differences, lipid concentrations for all tissues would be sex‐ and stock‐specific. Paddlefish were sampled in April and May 2012 from the Grand Lake, Oklahoma, USA (GL) and Yellowstone‐Sakakawea, North Dakota (SAK) stocks. Three tissue types were collected from SAK fish, including white muscle (N = 137; 83M, 54F), red muscle (N = 74; 36M, 38F), and GFB (N = 164; 105M, 59F). Samples from GL fish included white muscle (N = 60; 36M, 24F), and GFB (N = 42; 41M, 1F). Proximate analysis was used to quantify the percentage of lipid, water, ash, and protein in tissue samples. The lipid component of proximate analysis was completed using the ANKOM method of lipid extraction which uses heated petroleum ether to extract lipid. Also investigated were lipid accumulation and storage in adult paddlefish in relation to tissue type: white muscle, red muscle, gonadal fat bodies (GFB) within individual fish, between the sexes, and between the two different stocks. White muscle was significantly correlated with both red muscle and GFB lipid, but red muscle lipid was not significantly correlated with GFB lipid. White muscle lipid content differed by stock and sex, with the more northerly stock having higher lipid content than the more southerly stock. Females had higher white muscle lipid content (SAK F = 17.18, M = 9.09; GL F = 6.74, M = 4.99) and red muscle (SAK F = 47.72, M = 30.93) than males. In contrast, GFB lipid differed by sex but not stock, with females having lower lipid content (SAK = 79.79, GL = 89.08) than males (SAK = 89.08; GL = 91.72). Life history differences, growing season, and the role of metabolism may help explain the differences in concentrations of lipids observed between stocks and between the sexes for all three tissues. It was hypothesized that different tissues may be used for different metabolic processes and that although metabolism likely strongly influences muscle tissue lipid, GFB lipid is probably linked more closely to reproductive demands associated with gonad development.     

The suitability of the free-living nematode Panagrellus redivivus as live food for first-feeding fish larvae

The free‐living nematode Panagrellus redivivus has been recommended as a suitable food source for first‐feeding fish. A new technology for mass production of P. redivivus enables fish hatchery operators to rely on an inexpensive, standardized and permanently available live food for first‐feeding fish larvae. The proximate composition, and the fatty acid and amino acid profiles of nematodes mass produced on oat‐based and purified ingredient media were determined. The quality of nematodes was significantly influenced by the culture medium used. The lipid content and fatty acid composition of nematodes could be modified by using lipid‐enriched media. Mass‐produced nematodes were tested on first‐feeding common carp (Cyprinus carpio L.) and whitefish (Coregonus lavaretus) larvae. Carp larvae, grown on nematodes cultured on oat medium enriched with sunflower oil, showed a higher survival rate (87.1%) than the control group fed frozen zooplankton (82.9%) at the end of the 1‐week feeding experiment. Differences in larval mass between the treatments disappeared after subsequent feeding of a dry diet for 2 weeks. Whitefish larvae can be reared exclusively on a dry diet; here, the initial feeding of nematodes had no effect on final biomass and survival of larvae.     

Critical thermal maximum of Amur sleeper <Emphasis Type="Italic">Perccottus glenii</Emphasis> in different seasons of the year

Seasonal differences in the critical thermal maximum are revealed in Amur sleeper Perccottus glenii at the age of 1+ and 2 years. Maximum values of the critical thermal maximum determined by two criteria (rotation of the body and immobile gill covers) are registered in the summer, and minimum values of this parameter are observed in the winter. Maximum thermal resistance is registered at a low water heating rate (0.2–0.4°C/h), and minimum resistance is reported for a heating rate 5.0–10.5°C/h. The difference between the values of the critical thermal maximum assessed by two criteria reaches 1–2°C. Features of behavior of Amur sleeper are described at different heating rates. Based on the characteristics of thermal adaptation (critical thermal maximum and optimal temperature for growth), Amur sleeper is similar to warm water eurythermal species (goldfish Carassius auratus and carp Cyprinus carpio).     

Effects of diet and temperature on condition, proximate composition and three major macro elements, Ca, P and Mg, in barbel Barbus barbus juveniles

Six feeding groups of 60 early juveniles (0.6?g) of a rheophilic cyprinid barbel Barbus barbus were reared in triplicate in 18?glass aquaria. Fish fed a commercial formulated dry diet Aller Futura were compared with those on natural food??commercially available frozen Chironomidae larvae at 17, 21 and 25°C. Daily food rations were adjusted according to fish biomass, differences in hydration between the two diets, and rearing temperature. No mortality occurred during experiment. Temperature, and especially diet, both affected whole body proximate and mineral composition of B. barbus juveniles. Under conditions of this experiment the formulated diet Aller Futura seems to satisfy the demands for calcium and magnesium, while an elevated lipid content in this diet was found and marginally deficient phosphorus content can be concluded. These resulted in a depressed content of phosphorus and total minerals in the body, and in elevated lipid content and condition coefficient in the Futura-fed fish, but no visible body deformities occurred. Advantages and limitations of the condition coefficient K were discussed. This is a non-destructive, express indicator that can be used as a supplementary tool for estimation of changes in body proximate composition in fish juveniles of similar size within a population.     

Energy allocation in juvenile sablefish: effects of temperature, ration and body size

The lipid deposition of juvenile sablefish Anoplopoma fimbria was examined, in particular, the changes in allocation over time. Growth rates of early juveniles (initial size 36–50 mm total length, L_T) were manipulated using two temperatures (10 and 20° C) and two rations (ad libitum and 3–4% body mass day^?1). Fish L_T, mass and lipid content were measured every 3 weeks for 15 weeks. Irrespective of treatment, the relationship of total lipid content with body size was clearly hyperallometric; small juveniles allocated relatively more energy to growth and less to lipid storage than large juveniles. After adjusting for the influence of body size, temperature and ration significantly influenced body composition but these effects varied over the course of the experiment. In the first 3 week time period, fish on the high ration, high temperature treatment had reduced lipid storage relative to other treatments, but in all subsequent time periods their lipid concentrations were similar to or higher than those of fish on other treatments. In contrast, fish held at low rations and low temperatures initially had average levels of lipid concentration, but after 6 weeks their levels were lower than other treatments. Estimation of allocation to lipid storage over time (proportion of dry mass increase comprised of lipid) suggested that fish in all of the treatments were approaching an asymptotic level of lipid concentration (c. 50–60% of dry mass) but with different rates of lipid increase. Within a treatment, it was predicted that individual differences in allocation would result in trade‐offs between somatic growth and storage. This trade‐off was evident only for fish held on low rations at low temperatures. In contrast, fish held on high rations at high temperatures exhibited the opposite pattern of a positive correlation between somatic growth and storage. These results suggest that lipostatic regulation of appetite is unlikely in juvenile sablefish. When resources are unlimited, this species appears to adopt a maximizing strategy for both somatic growth and lipid accumulation.     

Metabolic fractionation of stable carbon isotopes: implications of different proximate compositions for studies of the aquatic food webs using δ13C data

In two laboratory experiments using tilapia (Oreochromis niloticus) and common carp (Cyprinus carpio), we investigated the effect of lipid content in the fish carcass on the stable carbon isotope ratio (δ¹³C). In both experiments, a significant negative correlation was found between lipid content in the carcass dry matter and δ¹³C of total dry matter, but there was no influence on the δ¹³C of fat-free dry matter or lipids. As the lipid content of fish is known to vary with reproductive stage, season or nutritional state, separate analysis of fractions of the proximate composition of dry matter will lead to more reliable results than analysing the whole body. The differences in δ¹³C between diet and fish carcass (trophic shift) were different for the two species, calling for feeding trials under controlled conditions using the species and the feeds in question before applying the stable isotope tracer technique in the analysis of food webs. Received: 1 December 1997 / Accepted: 1 March 1998     

Life-history traits of Amur sleeper, <Emphasis Type="Italic">Perccottus glenii</Emphasis>, in the invaded Vistula River: early investment in reproduction but reduced growth rate

In recent decades, Amur sleeper is one of the most invasive fish species in Eastern and Central Europe. Generally, it is assumed that success of an invasive species can largely depend on the plasticity of its life-history traits, e.g., indicated by higher investment in reproduction in the initial stage of its invasion. On the other hand, such energy allocation to production of gonads should negatively impact somatic growth rate. The aim of this article was to explore this phenomenon in a non-native population of Amur sleeper inhabiting artificial reservoir on a large lowland river in Central Europe. Through analysis of the population age structure, sex-dependent growth rate and life-history traits we assessed investment in reproduction and its possible relationship with growth pattern as well as compared the results with literature data from native and introduced range. Samples collected monthly from April 2005 to March 2006 were used to estimate sex ratio, GSI, fecundity, eggs diameter frequency over the year, duration of spawning season, and mode of spawning. Age structure was calculated from scales and the same data were also used to back-calculate standard length (SL) at age. The population in the Włocławski Reservoir was represented by eight age groups, and its life span was one of the longest recorded both in its natural range and introduced areas. The weight–length relationship showed that the Amur sleeper grew isometrically and there was no difference between females and males. Although the Amur sleeper growth rate was relatively slow in the Włocławski Reservoir, its increment in the first year of life was comparable to that in other introduced areas and higher than in its natural range. The female age of maturation (1+) found in the reservoir was earlier than observed in its natural range as well as in most introduced areas (2+, 3+). Female length at maturation was similar to that reported elsewhere. The back-calculated standard length (SL) of females and males showed that in the first two age classes (1 and 2) males achieved larger SL than females. Meanwhile, at age 3, 4 and 5, females were larger than males but these differences were insignificant, except for class 5. For both sexes the average annual increment of SL decreased with age but in females the increment was always higher than in males (P < 0.05). The growth rate differences between females and males resulted possibly from their unequal investment in reproduction. Multiple spawning in the Włocławski Reservoir lasted from April to the end of August and was almost 3 months longer than in other regions. Thus, this higher investment in reproduction displayed by, e.g., earlier maturation of females and longer spawning season in the Włocławski Reservoir, may contribute to invasive success of Amur sleeper in newly colonized areas; however, the costs of this strategy result in slower growth in older age classes.     

Parasites of the introduced Amur sleeper, Perccottus glenii (Osteichthyes): Alpha-diversity of parasites and age of the host

Three age groups of the Amur sleeper have been studied: underyearlings (0+), two-year-olds (1+), and three- and four-year-olds (2+ and 3+). The long-cycle parasites prevalent in all three age groups are the merocercoids of Ophiotaenia europaea, third-stage juveniles of Spiroxys contortus, and metacercariae of Isthmiophora sp. and Prohemistomidae gen. sp. Different age groups of the Amur sleeper have different roles in the life cycle of O. Europaea. Underyearlings are second intermediate hosts of O. europaea, and Amur sleepers aged 1+ and older are parathenic hosts. The alpha-diversity of long-cycle parasites of the Amur sleeper increases with the age of the host.     

Body macronutrient composition is predicted by lipid and not protein content of the diet

Diet is an important determinant of fitness‐related traits including growth, reproduction, and survival. Recent work has suggested that variation in protein:lipid ratio and particularly the amount of protein in the diet is a key nutritional parameter. However, the traits that mediate the link between dietary macronutrient ratio and fitness‐related traits are less well understood. An obvious candidate is body composition, given its well‐known link to health. Here, we investigate the relationship between dietary and body macronutrient composition using a first‐generation laboratory population of a freshwater fish, the three‐spine stickleback (Gasterosteus aculeatus). Carbohydrate is relatively unimportant in the diet of predatory fish, facilitating the exploration of how dietary protein‐to‐lipid ratio affects their relative deposition in the body. We find a significant effect of lipid intake, rather than protein, on body protein:lipid ratio. Importantly, this was not a result of absorbing macronutrients in relation to their relative abundance in the diet, as the carcass protein:lipid ratios differed from those of the diets, with ratios usually lower in the body than in the diet. This indicates that individuals can moderate their utilization, or uptake, of ingested macronutrients to reach a target balance within the body. We found no effect of diet on swimming endurance, activity, or testes size. However, there was an effect of weight on testes size, with larger males having larger testes. Our results provide evidence for the adjustment of body protein:lipid ratio away from that of the diet. As dietary lipid intake was the key determinant of body composition, we suggest this occurs via metabolism of excess protein, which conflicts with the predictions of the protein leverage hypothesis. These results could imply that the conversion and excretion of protein is one of the causes of the survival costs associated with high‐protein diets.     

Effects of contrasting omnivorous fish on submerged macrophyte biomass in temperate lakes: a mesocosm experiment

1. Freshwater fish can affect aquatic vegetation directly by consuming macrophytes or indirectly by changing water quality. However, most fish in the temperate climate zone have an omnivorous diet. The impact of fish as aquatic herbivores in temperate climates therefore remains unclear and depends on their dietary flexibility. 2. We tested the effects of a flexible omnivore and an herbivore on aquatic vegetation by comparing the effects of rudd (Scardinius erythrophthalmus, the most herbivorous fish in temperate climates) with grass carp (Ctenopharyngodon idella) in a mesocosm pond study. Exclosures distinguished herbivorous effects of fish on submerged macrophytes from indirect effects through changes in water quality, whereas stable isotope food‐web analysis provided information on fish diets. 3. We hypothesised that rudd, with its flexible diet and preference for animal food items, would only indirectly affect macrophytes, whereas grass carp, with its inflexible herbivorous diet, would directly affect macrophyte biomass. 4. Only grass carp significantly reduced macrophyte biomass through consumption. Rudd had no effect. Food‐web analysis indicated that rudd predominantly consumed animal prey, whereas grass carp included more plants in their diet, although they also consumed animal prey. Grass carp significantly affected water quality, resulting in lowered pH and increased N‐NH₄ concentrations, whereas more periphyton growth was observed in the presence of rudd. However, the indirect non‐herbivorous effects of both fish species had no effect on macrophyte biomass. 5. Both fish species should be considered as omnivores. Despite the fact that rudd is the most herbivorous fish in the western European climate zone, its effect on submerged macrophyte biomass is not substantial at natural densities and current temperatures.     

Proximate composition,energetic value,and relative abundance of prey fish from the inshore eastern Bering Sea: implications for piscivorous predators

Changing ocean conditions and subsequent shifts in forage fish communities have been linked to numerical declines of some piscivorous marine birds and mammals in the North Pacific. However, limited information about fish communities is available for some regions, including nearshore waters of the eastern Bering Sea, where many piscivores reside. We determined proximate composition and energetic value of a suite of potential forage fish collected from an estuary on the Yukon-Kuskokwim Delta, Alaska, during 2002 and 2003. Across species, energy density ranged from 14.5 to 20.7 kJ g⁻¹ dry mass and varied primarily as a function of lipid content. Total energy content was strongly influenced by body length and we provide species-specific predictive models of total energy based on this relationship; some models may be improved further by incorporating year and date effects. Based on observed energetic differences, we conclude that variation in fish size, quantity, and species composition of the prey community could have important consequences for piscivorous predators.     

Amur sleeper (Perccottus glenii Dubowski, 1877) spreading in the Danube River Basin

Whereas earlier Amur sleeper (Perccottus glenii) findings in the Danube River were only single specimens, data reported in this study represent the first record on establishment of a local population. An overview of the spread of the Amur sleeper in the Danube River basin is also presented.     

Fatty acid composition,cholesterol and fat‐soluble vitamins of wild‐caught freshwater spiny eel,Mastacembelus simack (Walbaum, 1792)

Because of its local commercial value, the proximate nutritional composition, fatty acid composition, cholesterol and fat‐soluble vitamins of spiny eel (Mastacembelus simack Walbaum, 1792) were determined in 20 specimens caught in Keban Dam Lake, Elaz??, Turkey in September 2009. Moisture, protein, lipid, ash and cholesterol contents of the spiny eel were 75.9, 17.1, 10.9, 1.0% and 52.6 mg per 100 g, respectively. Fatty acid composition showed that total polyunsaturated fatty acids (PUFA) were the highest (37.1%), followed by monounsaturated fatty acids (MUFA, 35.5%) and saturated fatty acids (SFA, 27.4%). The relation between PUFA and SFA (1.35) was comparable to that of some fish species, and PUFA : MUFA : SFA ratio (1.35 : 1.29 : 1) was very close to that recommended by nutritionists. Among the vitamins (A, D, E and K) analyzed, the vitamin E content was highest followed by D, K and A. In conclusion, M. simack is rich in proteins, unsaturated fatty acids and vitamins and low in cholesterol, with a reasonable PUFA : SFA and PUFA : MUFA : SFA ratio as recommended by nutritionists. M. simack can therefore be recommended for human consumption as a good source of nutrition.     

Common relationships among proximate composition components in fishes

Relationships between the various body proximate components and dry matter content were examined for five species of fishes, representing anadromous, marine and freshwater species: chum salmon Oncorhynchus keta, Chinook salmon Oncorhynchus tshawytscha, brook trout Salvelinus fontinalis, bluefish Pomatomus saltatrix and striped bass Morone saxatilis. The dry matter content or per cent dry mass of these fishes can be used to reliably predict the per cent composition of the other components. Therefore, with validation it is possible to estimate fat, protein and ash content of fishes from per cent dry mass information, reducing the need for costly and time‐consuming laboratory proximate analysis. This approach coupled with new methods of non‐lethal estimation of per cent dry mass, such as from bioelectrical impedance analysis, can provide non‐destructive measurements of proximate composition of fishes.