Food uptake follows rules defined by feeding behaviour that determines the kind and quantity of food ingested by fish larvae as well as how live prey and food particles are detected, captured and ingested. Feeding success depends on the progressive development of anatomical characteristics and physiological functions and on the availability of suitable food items throughout larval development. The fish larval stages present eco-morpho-physiological features very different from adults and differ from one species to another. The organoleptic properties, dimensions, detectability, movements characteristics and buoyancy of food items are all crucial features that should be considered, but is often ignored, in feeding regimes. Ontogenetic changes in digestive function lead to limitations in the ability to process certain feedstuffs. There is still a lack of knowledge about the digestion and absorption of various nutrients and about the ontogeny of basic physiological mechanisms in fish larvae, including how they are affected by genetic, dietary and environmental factors. The neural and hormonal regulation of the digestive process and of appetite is critical for optimizing digestion. These processes are still poorly described in fish larvae and attempts to develop optimal feeding regimes are often still on a 'trial and error' basis. A holistic understanding of feeding ecology and digestive functions is important for designing diets for fish larvae and the adaptation of rearing conditions to meet requirements for the best presentation of prey and microdiets, and their optimal ingestion, digestion and absorption. More research that targets gaps in our knowledge should advance larval rearing.

Despite considerable progress in recent years, many questions regarding fish larval nutrition remain largely unanswered, and several research avenues remain open. A holistic understanding of the supply line of nutrients is important for developing diets for use in larval culture and for the adaptation of rearing conditions that meet the larval requirements for the optimal presentation of food organisms and/or microdiets. The aim of the present review is to revise the state of the art and to pinpoint the gaps in knowledge regarding larval nutritional requirements, the nutritional value of live feeds and challenges and opportunities in the development of formulated larval diets.

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The current commercial production protocols for Atlantic cod depend on enriched rotifers and Artemia during first-feeding, but development and growth remain inferior to fish fed natural zooplankton. Two experiments were conducted in order to identify the underlying factors for this phenomenon. In the first experiment (Exp-1), groups of cod larvae were fed either (a) natural zooplankton, mainly copepods, increasing the size of prey as the larvae grew or (b) enriched rotifers followed by Artemia (the intensive group). In the second experiment (Exp-2), two groups of larvae were fed as in Exp-1, while a third group was fed copepod nauplii (approximately the size of rotifers) throughout the larval stage. In both experiments, growth was not significantly different between the groups during the first three weeks after hatching, but from the last part of the rotifer feeding period and onwards, the growth of the larvae fed copepods was higher than that of the intensive group. In Exp-2, the growth was similar between the two copepod groups during the expeimental period, indicating that nutrient composition, not prey size caused the better growth on copepods. Analyses of the prey showed that total fatty acid composition and the ratio of phospholipids to total lipids was slightly different in the prey organisms, and that protein, taurine, astaxanthin and zinc were lower on a dry weight basis in rotifers than in copepods. Other measured nutrients as DHA, all analysed vitamins, manganese, copper and selenium were similar or higher in the rotifers. When compared to the present knowledge on nutrient requirements, protein and taurine appeared to be the most likely limiting nutrients for growth in cod larvae fed rotifers and Artemia. Larvae fed rotifers/Artemia had a higher whole body lipid content than larvae fed copepods at the end of the experiment (stage 5) after the fish had been fed the same formulated diet for approximately 2 weeks.

The current best practice intensive culture of larval Atlantic cod includes feeding rotifers from onset of exogenous feeding until 25–30 days after hatching. These larvae grow considerably slower and develop higher frequencies of deformities than larvae reared in semi‐extensive systems, using copepods as feed. The present study compares the micronutrient concentrations in rotifers with those of copepods, with the aim of identifying nutrients that may be limiting for normal growth and development of cod larvae. An additional criterion used is the nutrient requirements given for fish in general, by NRC (1993), as nutrient requirements of cod remains to be determined. Rotifers were fed on four different diets, consisting of baker's yeast with cod liver oil (3.3 : 1 dry weight (DW)/v), baker's yeast with Algamac 2000TM (3.5 : 1 DW), baker's yeast with live algae Chlorella (4.1 : 1 DW), and Culture Selco 3000TM (CS). CS was a complete commercial diet for rotifers while the other diets are considered as based on raw ingredients. Compared with copepod nutrient levels, rotifers grown on yeast‐based diets supplemented with either cod liver oil, Algamac 2000 or Chlorella were apparently sufficient for covering the requirements in cod larvae for all the B‐vitamins, except thiamine. Rotifers cultured on the CS diet also had sufficient amounts of thiamine. Of the minerals, only calcium and magnesium were sufficient, using this criterion while iron was on the borderline. However, with reference to the requirements given for larger fish (NRC 1993), only thiamine, vitamin A, manganese, selenium and perhaps copper, appear too low in the rotifers cultured without extra micronutrient supplementation. The other nutrients were present at levels intermediate between copepod and fish requirement levels. This study suggests that it is necessary to develop enrichment techniques to produce rotifers with sufficient amounts of all micronutrients. Such techniques will also be important tools for determining which nutrients are present at levels below the actual requirements in cod larvae.

The present study reports the complete coding sequences for two paralogues for leptin (sLepA1 and sLepA2) and leptin receptor (sLepR) in Atlantic salmon. The deduced 171-amino acid (aa) sequence of sLepA1 and 175 aa sequence for sLepA2 shows 71.6% identity to each other and clusters phylogenetically with teleost Lep type A, with 22.4% and 24.1% identity to human Lep. Both sLep proteins are predicted to consist of four helixes showing strong conservation of tertiary structure with other vertebrates. The highest mRNA levels for sLepA1 in fed fish (satiation ration=100%) were observed in the brain, white muscle, liver, and ovaries. In most tissues sLepA2 generally had a lower expression than sLepA1 except for the gastrointestinal tract (stomach and mid-gut) and kidney. Only one leptin receptor ortholog was identified and it shares 24.2% aa sequence similarity with human LepR, with stretches of highest sequence similarity corresponding to domains considered important for LepR signaling. The sLepR was abundantly expressed in the ovary, and was also high in the brain, pituitary, eye, gill, skin, visceral adipose tissue, belly flap, red muscle, kidney, and testis. Fish reared on a rationed feeding regime (60% of satiation) for 10 months grew less than control (100%) and tended to have a lower sLepA1 mRNA expression in the fat-depositing tissues visceral adipose tissue (p<0.05) and white muscle (n.s.). sLepA2 mRNA levels was very low in these tissues and feeding regime tended to affect its expression in an opposite manner. Expression in liver differed from that of the other tissues with a higher sLepA2 mRNA in the feed-rationed group (p<0.01). Plasma levels of sLep did not differ between fish fed restricted and full feeding regimes. No difference in brain sLepR mRNA levels was observed between fish fed reduced and full feeding regimes. This study in part supports that sLepA1 is involved in signaling the energy status in fat-depositing tissues in line with the mammalian model, whereas sLepA2 may possibly play important roles in the digestive tract and liver. At present, data on Lep in teleosts are too scarce to allow generalization about how the Lep system is influenced by tissue-specific energy status and, in turn, may regulate functions related to feed intake, growth, and adiposity in fish. In tetraploid species like Atlantic salmon, different Lep paralogues seems to serve different physiological roles.

The simultaneous effect of temperature (5, 7, 10 and 13°C) and light on the rates of oxygen consumption and ammonia excretion of larval and early juvenile Atlantic cod Gadus morhua was examined. Larvae increased their mean dry body mass by 2000 times within 48 d. Instantaneous growth rate exceeded 30% d -1 towards the end of the study period, and proportionality of growth followed a triphasic pattern, during which body water content significantly declined but no inflection could be detected in the metabolic exponents. Data were rigorously tested via Model-I (least squares) and Model-II (geometric mean) regression techniques, and the aerobic metabolic rate was found to scale allometrically with both dry and wet body mass. The metabolic exponent was not affected by increasing temperature, but was significantly decreased by the presence of light (b = 0.88 to 0.89 for light-adapted larvae; b = 0.90 to 0.91 for dark-adapted larvae). The effect of light on small larvae (4 to 7 mm standard length, SL) caused a 30 to 40% increase in metabolic rate, while no effect was observed in larger juveniles (40 to 60 mm SL). Acute temperature acclimation of Atlantic cod of 4 to 60 mm SL (0.04 to 350 mg dry mass) demonstrated normal thermal sensitivity with Q 10 values of 2.4 for dark-adapted larvae and 2.6 for light-adapted larvae. Rates of ammonia excretion also scaled allometrically with wet and dry body mass and showed greater variability in dark-adapted compared to light-adapted larvae. Comparison of the molar rates of ammonia excretion and oxygen consumption revealed that Atlantic cod larvae have a high reliance on amino acids as fuel for energy dissipation. With lipids as the assumed co-substrate, amino acids were estimated to account for 70 to 95% of total substrate oxidation for larvae up to 7 mm SL (first 3 to 4 wk of post-hatch development). Beyond 7 mm SL, the reliance on amino acids as fuel began to decline, but even in juveniles of 40 to 60 mm SL, amino acids still represented the dominant source of fuel. For juveniles of between 10 and 20 mm SL, both the rates of oxygen consumption and ammonia excretion remained unaffected by the presence of food in the gut. For short-term fasted juveniles (35 to 60 mm SL), however, a substantial decline in the rate of ammonia excretion was observed. This indicates that during short-term fasting (8 to 12 h) early juvenile Atlantic cod conserve amino acids, rather than funneling them into the tricarboxylic acid cycle.KEY WORDS: Scaling · Metabolism · Fuel preference · Free amino acid · Q 10 · Temperature · Cod larvae Resale or republication not permitted without written consent of the publisherMar Ecol Prog Ser 243: [217][218][219][220][221][222][223][224][225][226][227][228][229][230][231][232][233][234] 2002 1975). For the juvenile and adult stages of fishes, however, a general exponent of 0.8 has been proposed (Winberg 1956, Wieser 1995, Clarke & Johnston 1999. Giguère et al. (1988) have proposed that the metabolic rates of larval fishes scale isometrically with incre...