The utilisation of dietary carbohydrates and their effects on fish metabolism are reviewed. Details on how dietary carbohydrates affect growth, feed utilisation and deposition of nutrients are discussed. Variations in plasma glucose concentrations emphasizing results from glucose tolerance tests, and the impact of adaptation diets are interpreted in the context of secondary carbohydrate metabolism. Our focus then shifts to selected aspects of hormonal regulation of carbohydrate metabolism and dietary carbohydrates and their variable effects on glycogen and glucose turnover. We analyse the interaction of carbohydrates with other nutrients, especially protein and protein sparing, and de novo synthesis of lipids, and finish by discussing the correlation of dietary carbohydrates with fish health.

Carbohydrates in fish nutrition: effects on growth, glucose metabolism and hepatic enzymes

Teleost fishes represent a highly diverse group consisting of more than 20,000 species living across all aquatic environments. This group has significant economical, societal and environmental impacts, yet research efforts have concentrated primarily on salmonid and cyprinid species. This review examines carbohydrate/glucose metabolism and its regulation in these model species including the role of hormones and diet. Over the past decade, molecular tools have been used to address some of the downstream components of these processes and these are incorporated to better understand the roles played by carbohydrates and their regulatory paths. Glucose metabolism remains a contentious area as many fish species are traditionally considered glucose intolerant and, therefore, one might expect that the use and storage of glucose would be considered of minor importance. However, the actual picture is not so clear since the apparent intolerance of fish to carbohydrates is not evident in herbivorous and omnivorous species and even in carnivorous species, glucose is important for specific tissues and/or for specific activities. Thus, our aim is to up-date carbohydrate metabolism in fish, placing it to the context of these new experimental tools and its relationship to dietary intake. Finally, we suggest that new research directions ultimately will lead to a better understanding of these processes.

Glucose metabolism in fish: a review

This review summarizes information regarding digestion and absorption of carbohydrates in cultivated fish Relevant results of studies of digestive enzymes, eg amylase, chitinase, cellulase and brush border disaccharidases are presented Fish amylases appear to be molecularly closely related and to have characteristics comparable to mammalian amylases Whether chitinases and cellulases are endogenous enzymes of some fish species is still a matter of speculation, although recent molecular evidence, at least for chitinase seems to settle the issue in favour of endogenous sources Feed and intestinal microbes may be the source of polysaccharidases in fish feeding on nutrients-containing non-starch polysaccharides Knowledge regarding monosaccharide transport in fish intestine as interpreted from studies of brush border membrane vesicles, everted sleeves of fish intestinal sections and molecular biology is discussed Glucose transporters of the intestinal brush border show characteristics similar to those found in mammals A tabulatory presentation of experimental details and results reported in the literature regarding starch digestibility is included as a basis for discussion Although numerous investigations on digestion of starch and other carbohydrates in fish have been published, the existing information is highly fragmentary As yet, it is impossible to derive a cohesive picture on the integrated process of carbohydrate hydrolysis and absorption and interaction with diet composition for any of the fish species under cultivation The physiological mechanisms behind the species differences are not known

Carbohydrates in fish nutrition: digestion and absorption in postlarval stages

Farming of Atlantic salmon (Salmo salar) began in Norway in the late 1960s. During the 1980s and 1990s the production spread, mainly to other northwestern European countries and to Chile. In 1980 annual world production was less than 10,000 t and this increased to 850,000 t whole fish produced in 2000. It is projected to approach 2 million t in 2010. The growth rate of the fish in the farms is continuously improving, due to genetic selection, improved diets and feeding and improved management. The improvement in growth ascribed to genetic selection can be as much as 15–20% per generation if the salmon are from a combined family–phenotype selection programme (Gjedrem, 1983). Recent Canadian results have shown that transgenic salmon, which express growth hormone receptor activity in muscle tissues, have a growth rate 2.6–2.9 times that of the control population, mainly because of increased feed intake (Cook et al., 2000a). Selection or transgenic techniques may also influence the composition of the growth, result in improved feed conversion (Thodesen et al., 1999; Cook et al., 2000a) and change the rate of metabolism (Thodesen et al., 1999; Cook et al., 2000b,c). Economical feed conversions in Norwegian salmon farming have changed from more than 2 kg feed dry matter (DM) kg−1 gain in 1980 to approximately 1.15 kg kg−1 at the end of the 1990s. In salmon the biological feed conversion can vary from less than 0.6 to more than 1.3 kg intake kg−1 gain, and the proportion of protein and energy in diet partitioned into gain can vary from 25 to 60%, depending on the formulation of the diet and the size of the fish. Thus, requirement figures must be based on the requirements of the animal, related to growth, nutrient partitioning and the specific physiological processes in which the nutrient is required. Requirements differ throughout the life cycle. This is especially important in salmon, which undergo physiological changes during smoltification and transfer from a life in fresh water to salt water. In addition to developing the ability to maintain the ion balance in the hyperosmotic salt water, the salmon also

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Atlantic Salmon, Salmo Salar

Glucose plays a key role as energy source in the majority of mammals, but its importance in fish appears limited. Until now, the physiological basis for such apparent glucose intolerance in fish has not been fully understood. A distinct regulation of hepatic glucose utilization (glycolysis) and production (gluconeogenesis) may be advanced to explain the relative inability of fish to efficiently utilize dietary glucose. We summarize here information regarding the nutritional regulation of key enzymes involved in glycolysis (hexokinases, 6-phosphofructo-1-kinase and pyruvate kinase) and gluconeogenesis (phosphoenolpyruvate carboxykinase, fructose-1,6-bisphosphatase and glucose-6-phosphatase) pathways as well as that of the bifunctional enzyme 6-phosphofructo-2-kinase/fructose-2,6-bisphosphatase. The effect of dietary carbohydrate level and source on the activities and gene expression of the mentioned key enzymes is also discussed. Overall, data strongly suggest that the liver of most fish species is apparently capable of regulating glucose storage. The persistent high level of endogenous glucose production independent of carbohydrate intake level may lead to a putative competition between exogenous (dietary) glucose and endogenous glucose as the source of energy, which may explain the poor dietary carbohydrate utilization in fish.

Nutritional regulation of hepatic glucose metabolism in fish

Individually tagged Atlantic salmon, Salmo salar L., were fed three different adaptation diets containing low, medium or high levels of extruded starch, before intraperitoneal injection of glucose. The injections were done at winter and at summer temperatures. This was to investigate whether the tolerance for glucose varied according to adaption diet and/or according to season. The results showed a temperature-dependent response. At winter temperatures (2°C) the Atlantic salmon had problems adapting to both medium and high levels of dietary starch. This was indicated by lower growth response in the adaptation period, as well as by delayed glucose regulation. At summer temperatures (12.5°C) the Atlantic salmon could tolerate and utilize both medium and high levels of dietary starch. In summer also all groups responded evenly to the tolerance tests with respect to blood glucose regulation. Incorporation of liver glycogen was higher at summer than at winter temperatures in the high- carbohydrate group. No effect was found on haematocrit values as a response to diet or season.

Glucose tolerance in Atlantic salmon, Salmo salar L., dependence on adaption to dietary starch and water temperature

Two size groups (0.5 and 3 kg) of Atlantic salmon, Salmo salar L., were stressed by netting and transport from sea cages into land-based tanks. Handling lasted for 1 h. Before onset of stress, basal concentrations of haematocrit and haemoglobin, plasma cortisol and glucose, and liver and muscle glycogen were measured. All levels were within ranges reported for salmon. After handling and at chosen time intervals during recovery, samples were taken to judge the impact of handling and whether secondary changes in carbohydrate metabolism were related of fish size. A low cortisol peak indicated a mild stress reaction after handling independent to fish size. Plasma glucose peaked as cortisol declined, and returned to basal levels within 48 h. Liver glycogen seemed to be the main source of plasma glucose. No changes were measured in muscle glycogen concentrations. The results indicate a high tolerance to handling stress in Atlantic salmon independent of fish size.

Å. Krogdahl

Papers

Carbohydrates in fish nutrition: effects on growth, glucose metabolism and hepatic enzymes

Glucose tolerance in Atlantic salmon, Salmo salar L., dependence on adaption to dietary starch and water temperature

Effect of handling and fish size on secondary changes in carbohydrate metabolism in Atlantic salmon, Salmo salar L.