The role of the generally unrecognised microprey source as food for larval fish in the Irish Sea

Few studies have studied the food supply to larval fish in the Irish Sea; thus, we have assessed the full prey-field available to larval fish, ranging from protozoa to copepods. Specifically we assessed if fish larvae feed on protozoa, as suggested by others, and if densities of the protozoa and the appropriate size of metazoan prey were previously underestimated. By examining the gutcontents of fish larvae, the prey available to them, and the potential accessibility of prey to fish, we develop a simple food web, presented as a box-model. By doing so, we indicate that the lack of focus on small metazoa and protozoa has underestimated the food available to fish larvae; without these, we might have concluded that prey levels were too low to support the growth of the larval fish assemblage. Our methods were as follows. Sampling was at two sites, off the Isle of Man, with distinct physical and biological structures, soon after fish spawning: the southwest coast, where many species occur in spring-summer (23 April; 6, 19 May; 1, 22 June; 12 July) and the east coast, where only herring larvae occur in September–November (12, 28 October). Microplankton (15–200 μm), mesozooplankton, and larval fish were collected at 1, 15, and 25 m: microplankton with 1.5 L bottles and a 64 μm-mesh net; mesozooplankton and larval fish with a Gulf VII high-speed sampler (280 μm mesh). The 64 μm mesh net, mounted on the Gulf VII, provided simultaneous hauls. Fixed samples were evaluated to determine species composition, abundance, and biomass. Larval fish diet was determined from fish collected by short net hauls: fixed guts were examined and prey, including protozoa, analysed. Using physical data as a guide, plankton data were integrated through the water column to determine standing stocks. Size-based food availability to larval fish was estimated from the gut contents. The role of protozoa was examined, assuming that they are digested at the same rate as metazoan and if they are digested 2.5–10 times faster; increased digestion rates indicated that they contributed substantially to the larval fish diet.     

Foraging behavior in early Atlantic cod larvae (Gadus morhua) feeding on a protozoan (Balanionsp.) and a copepod nauplius (Pseudodiaptomussp.)

Food limitation is likely to be a source of mortality for fish larvae in the first few weeks after hatching. In the laboratory, we analyzed all aspects of foraging in cod larvae (Gadus morhua Linnaeus) from 5 to 20 d post-hatching using protozoa (Balanion sp.) and copepod nauplii (Pseudodiaptomus sp.) as prey. A camera acquisition system with two orthogonal cameras and a digital image analysis program was used to observe patterns of foraging. Digitization provided three-dimensional speeds, distances, and angles for each foraging event, and determined prey and fish larval head and tail positions. Larval cod swimming speeds, perception distances, angles, and volumes increased with larval fish size. Larval cod swam in a series of short intense bursts interspersed with slower gliding sequences. In 94% of all foraging events prey items were perceived during glides. Larval cod foraging has three possible outcomes: unsuccessful attacks, aborted attacks, and successful attacks. The percentage of successful attacks increased with fish size. In all larval fish size classes, successful attacks had smaller attack distances and faster attack speeds than unsuccessful attacks. Among prey items slowly swimming protozoans were the preferred food of first-feeding cod larvae; larger larvae had higher swimming speeds and captured larger, faster copepod nauplii. Protozoans may be an important prey item for first-feeding larvae providing essential resources for growth to a size at which copepod nauplii are captured. Received: 20 April 1999 / Accepted: 12 January 2000     

Laboratory study of predation by Aurelia aurita on larvae of cod,flounder, plaice and herring: development and vulnerability to capture

Capture success of the medusa Aurelia aurita preying on various developmental stages of fish larvae was measured together with larval reactivity and escape speed after being stung. These experiments were conducted in the spring of 1983 with A. aurita medusae collected from Loch Etive, Scotland and laboratory-reared larvae of Gadus morhua L., Platichthys flesus L., Pleuronectes platessa L. and Clupea harengus L. Capture success of the medusae increased with medusa size, but decreased with advancing larval development. Smaller species of larvae were more vulnerable to capture. Larval reactivity to encounters with medusae increased with advancing development, and larger species of larvae were more reactive to encounters. Larval escape swimming speeds also increased with advancing larval development and size. These results indicate that earlier stages of larvae within a species and smaller species of larvae at a given stage are more vulnerable to predation by medusae since they are less reactive to encounters. Apparently they are more susceptible to the effects of neurotoxins. Predation rates on different developmental stages of herring larvae are documented and compared with rates predicted by a predation model. Predictions fell within the range of observed predation rates, but tended to overestimate rates by larger medusae feeding on larger herring larvae. This indicates the possibility of predator satiation and/or behavioural avoidance.     

Fatty acid profiles of juvenile salmon indicate prey selection strategies in coastal marine waters

Juvenile salmon exhibit high growth rates upon their arrival into the marine environment. Dietary changes from freshwater and estuarine habitats to those derived from the marine environment may play an important role in ultimate adult survival. We measured the total lipid and fatty acid (FA) composition of juvenile Chinook salmon (Oncorhynchus tshawytscha), coho salmon (O. kisutch), and 18 of their potential prey items sampled from coastal waters during their first few months at sea. Coho salmon had significant reductions in their lipid content (% wet weight) between May and June, likely due to early marine growth. We did not find a significant drop between May and June Chinook salmon lipid content, which may indicate an earlier ontogenetic selection to marine prey that are higher in lipids and essential fatty acids (EFAs). Juvenile salmon ate prey of both high and low lipids. Significant FA compositional changes occurred for both coho and Chinook salmon between May and June. In May, the FA profile of juvenile salmon, especially coho salmon, did not resemble their prey items; however, in June, there was a strong correlation between salmon and their common fish prey as determined by gut content analysis. Significant increases in the level of EFAs, especially docosahexaenoic acid (DHA, 22:6n-3) accounted for the majority of the monthly differences in salmon tissue FA composition. In order for juvenile salmon to adequately meet their physiological requirements, they may have adapted to select advantageous prey with higher levels of EFAs, especially DHA, in order to rapidly increase their growth and ultimate survival.     

Temporal and spatial variation in the growth rates of Baltic herring (<Emphasis Type="Italic">Clupea harengus membras</Emphasis> L.) larvae during summer

It has been suggested that larval survival determines the year-class strength in most marine fish species. During their growth and development, the ability of the larvae to catch prey and avoid predation will increase. However, the factors affecting short-term changes in the growth of Baltic Sea herring have been little studied in the field. We collected Baltic herring (Clupea harengus membras L.) larvae from five different towing areas in the Archipelago Sea (SW Finland) during May and June 1989, right after the main spawning season. Twenty thousand two hundred and ten larvae were analysed and the area-specific growth rate (i.e. increase in standard length) was estimated by tracing the larval cohorts from the length-frequency data. This represents the first Baltic herring study with daily sampling during a long study period. The growth rate was related to environmental factors, such as temperature, number of zooplankters, and wind speed and direction. Large variation in larval growth rate occurred between areas: lowest and highest growth rates were 0.18 and 0.52 mm·day^-1. Temperature was an important variable controlling larval-fish growth rate. An increase of one 1°C in average water temperature corresponded to an increase in growth rate of 0.043 mm·day^-1. This may have been caused either by a direct temperature effect (changes in metabolic rate) or by the indirect effect of changes in food availability. We also found the densest herring populations in the areas with highest average water temperature. However, temperature and larval growth rate both increased towards the inner archipelago.     

Male characteristics, sperm traits, and reproductive success in winter-spawning Celtic Sea Atlantic herring, Clupea harengus

The relationship between sperm characteristics and reproductive success was examined in male herring, Clupea harengus L. Males were categorised as being first-time or repeat spawners on the basis of their age; they were also grouped according to whether their sperm were immediately active and exhibited forward motion on contact with seawater (FM) or had little or only vibratory motion (VM). Unlike the Pacific herring C. pallasii Valencienes, Atlantic herring sperm is usually motile on contact with seawater. The age, weight and gonadosomatic index (testes mass as a percentage of somatic mass = GSI) were measured and used as characteristics for individual fish. Sperm traits measured were (1) adenosine triphosphate (ATP) concentration, (2) sperm count, (3) duration of sperm motility. Reproductive success for each male was estimated from the fertilisation rate and from the length of larvae at hatching. Fertilisation rates for all fish were generally >80%. The ATP concentration of non-activated spermatozoa was negatively correlated with fertilisation rate. Among repeat spawners, fish with higher GSIs produced larvae that were larger at hatching. Although VM sperm fertilised eggs at rates equivalent to fertilisation by FM sperm, the larvae produced by VM sperm were significantly smaller at hatching. Larval length tended to increase in parallel with the duration of sperm motility, but the relationship was not significant in these tests. The results did not indicate any age or size pattern to spawning readiness in male herring. Sperm that are not yet ready to be shed are not fully motile on contact with seawater, but are still capable of fertilising eggs that hatch successfully. There is likely to be a progression of males which come into spawning readiness within a spawning shoal; therefore it is possible that paternal influences would result in a progressive decrease in larval size over the spawning period in winter-spawning Celtic Sea herring. Received: 22 November 1997 / Accepted: 8 June 1998     

Lipid class and fatty acid composition of brain lipids from Atlantic herring (Clupea harengus) at different stages of development

Little is known about the changes in composition of brain lipids and fatty acids at different stages of development in fish. Wild Atlantic herring (Clupea harengus L.) were collected from Loch Linnhe and the Firth of Clyde, Scotland, from August 1990 to March 1991. Lipid class and fatty acid compositions of brain lipids were studied at four different stages of development: larvae at the end of the yolk sac stage, two juvenile stages and sexually mature adults. The total lipid content in brains increased during development, and larval brains contained higher proportions of neutral lipids and lower proportions of polar lipids than the brains of juvenile or adult herring. Increased proportions of polar lipids in juvenile and adult herring brains were mainly due to increased percentages of phosphatidylcholine (PC), phosphatidylethanolamine (PE), cerebrosides and sulphatides. The increase in the proportions of the glycolipid classes suggested increasing levels of myelination with development. In total lipids, saturated fatty acids generally decreased and monounsaturated fatty acids and dimethyl acetals (derived from PE-plasmalogen) increased from larvae to adults. However, the proportions of polyunsaturated fatty acids in individual phosphoglycerides were generally highest in juvenile stages, due mainly to increased 22:6n-3, and were lowest in adult fish. Relatively high percentages of 24:1 isomers were found in all the phosphoglycerides, but primarily PC, and these increased during development from larvae to adult. Fatty acids were distributed between individual phosphoglycerides with a characteristic pattern that did not change with development, although the relative amounts of individual fatty acids were altered. The variations and roles of the different lipid components of herring brain are discussed with respect to lipid compositions and functions in brains of other fishes and vertebrates.     

A laboratory study of predation by Aurelia aurita on larval herring (Clupea harengus): Experimental observations compared with model predictions

Predation by the medusa Aurelia aurita L. on early first-feeding stage larvae of the herring clupea harengus L. was studied in the laboratory. The medusae were captured in Loch Etive, Scotland. Herring larvae were reared from the extificially fertilized eggs of spawning Clyde herring caught in March, 1982. Swimming speeds, volume searched”, capture efficiency and predation rates increased as medusa size increased. Predation rates on fish larvae increased with prey density, but appeared to approach a maximum at high prey densities; in 1 h experiments, a maximum rate of predation of 6.64 larvae h^-1 was estimated by fitting an Ivlev function. A model to predict predation rates was constructed from swimming speeds, sizes and densities of medusae and larvae, and capture efficiency. The rates of predation predicted from the model fell within the range of experimental data, but tended to underestimate rates and did not account for saturation of medusae. Swimming patterns of medusae changed after prey capture: (a) before capture, encounter rates were low and medusae were relatively less active; (b) after capture of 1 larva, encounter rates doubled, with the stimulated medusae exhibiting increased activity and an aftered “searching” path; and (c) after capture of many larvae, swimming speeds and encounter rates of medusae decreased.     

Nitrogen balance in marine fish larvae: influence of developmental stage and prey density

The utilization and fate of nitrogen in larvae of plaice (Pleuronectes platessa), blenny (Blennius pavo) and herring (Clupea harengus), from the stage of first-feeding to metamorphosis, was examined under laboratory conditions. Rates of ammonia excretion, primary amine defaecation, and growth in terms of protein-nitrogen were monitored throughout larval life. Data were used to calculate daily ration, the coefficient of nitrogen utilization (absorption efficiency), and gross and net growth efficiencies. The developmental pattern of nitrogen balance was similar for plaice and blenny larvae. These species showed increasing growth efficiency (k₁: 55 to 80%) with decreasing weight-specific waste nitrogen losses with age. Absorption efficiencies. were high (83 to 98%) in plaice and blenny larvae, and tended to increase with development in the former species. Ration relative to body weight decreased with growth in both species. Herring larval development, although at a slower rate than blenny and plaice, appeared normal up to 33 d, after which high mortality occurred. Absorption efficiency in this species tended to decline (83 to 43%) with age, until metabolic costs exceeded the absorbed ration and growth ceased. Artemia sp. nauplii proved a suitable food source for the rearing of plaice and blenny larvae, but this diet may have long-term toxicity or deficiency effects on herring. Availability and density of food affected nitrogen balance in the larvae of all three species. Feeding stimulated the output of wastes in excretion and defaecation by a factor of up to ten times the 12-h non-feeding basal rates. Waste nitrogen output reached a peak some 2 to 3 h after commencement of feeding and returned slowly to the baseline in 5 to 10 h after cessation of feeding. There was an asymptotic increase in ration, ammonia output and growth of larvae as prey density increased. Ration saturated at a higher prey density (>4 prey ml^-1) than either growth or excretion rate (1 prey ml^-1). Thus the efficiency with which food is absorbed and utilized for growth must eventually decline in response to high prey density. The idea that larval fish are adapted to maximize ingestion and growth rate, rather than optimize growth efficiency and thus to respond to prey occurring in either low density or in occasional patches, is supported by these results.     

Feeding ecology of herring (Clupea harengus) larvae in the turbid Blackwater Estuary

Most studies on feeding by herring larvae (Clupea harengus) have taken place in clear, open waters, but several herring stocks around the world spawn in inshore and estuarine regions. An example is the spring-spawning Blackwater Estuary (Essex, England) stock. Samples were collected in this estuary to examine prey selectivity and feeding levels in relation to biological and environmental conditions. Herring larvae negatively selected copepod nauplii, but positively selected the copepodite and adult stages of Acartia spp. Gastropod larvae were also positively selected. Particles >150 μm width were preferred, whilst particles smaller than this value were preferentially rejected. Concentrations of potential prey items in the water were in the range of 6.0 to 49.7 organisms l⁻¹ with a median concentration of 15.0 organisms l⁻¹ (n = 26). These values are towards the low end of prey concentrations quoted in the literature as being required to sustain herring larval growth and survival. However, theoretical considerations suggest that, in this environment, levels of tidally-induced turbulence enhance encounter rates between larval herring and their prey. On the other hand, turbidity is also related to tidal current speed and might reduce feeding success by decreasing underwater light levels. Measurements at two sites in the estuary confirmed that tidally-induced turbidity reduced the effective water depth in which herring larvae could visually feed by up to 50% at times of peak current speed. However, with the gut-content data available in the present study, it was not possible to discern any clear relationships between feeding success and the state of the tide. Feeding success appeared to be more strongly influenced by surface light-levels. Received: 24 June 1998 / Accepted: 17 February 1999     

Amino acid metabolism and protein turnover in larval turbot (Scophthalmus maximus) fed natural zooplankton or Artemia

The present paper studied the influence of different food regimes on the free amino acid (FAA) pool, the rate of protein turnover, the flux of amino acids, and their relation to growth of larval turbot (Scophthalmus maximus L.) from first feeding until metamorphosis. The amino acid profile of protein was stable during the larval period although some small, but significant, differences were found. Turbot larvae had proteins which were rich in leucine and aspartate, and poor in glutamate, suggesting a high leucine requirement. The profile of the FAA pool was highly variable and quite different from the amino acid profile in protein. The proportion of essential FAA decreased with development. High contents of free tyrosine and phenylalanine were found on Day 3, while free taurine was present at high levels throughout the experimental period. Larval growth rates were positively correlated with taurine levels, suggesting a dietary dependency for taurine and/or sulphur amino acids. Reduced growth rates in Artemia-fed larvae were associated with lower levels of free methionine, indicating that this diet is deficient in methionine for turbot larvae. Leucine might also be limiting turbot growth as the different diet organisms had lower levels of this amino acid in the free pool than was found in the larval protein. A previously presented model was used to describe the flux of amino acids in growing turbot larvae. The FAA pool was found to be small and variable. It was estimated that the daily dietary amino acid intake might be up to ten times the larval FAA pool. In addition, protein synthesis and protein degradation might daily remove and return, respectively, the equivalent of up to 20 and 10 times the size of the FAA pool. In an early phase (Day 11) high growth rates were associated with a relatively low protein turnover, while at a later stage (Day 17), a much higher turnover was observed. Received: 19 March 1997 / Accepted: 14 April 1997     

Comparison of laboratory rates of predation of five species of marine fish larvae by three planktonic invertebrates: effects of larval size on vulnerability

Rates of predation by the invertebrates Aurelia aurita, Thysanoessa raschi and Euchaeta norvegica on larval stages of cod (Gadus morhua L.), flunder (Platichthys flesus L.), plaice (Pleuronectes platessa L.), herring (Clupea harengus L.), and turbot (Scophthalmus maximus L.) were determined. Experiments were conducted in late winter and early spring 1982 with predators collected in Loch Etive, Scotland and prey obtained from several locations in Great Britain. Early stages of the smallest species, cod, flounder and turbot, tended to be most vulnerable to all three predators, while the early stages of the larger species, plaice and herring, and older stages of all species, were less vulnerable. For all stages and species of larvae, predation rates by the three predators were most closely related to larval length and escape swimming speed. Larval length itself was closely correlated to indices of larval escape ability. Low predation rates on large larvae by E. norvegica could be due to handling difficulties, whereas for A. aurita and T. raschi these low rates were due to escape abilities of the larger larvae. Prey movement is an important stimulus eliciting predation in E. norvegica but not in A. aurita or T. raschi.     

Growth and mortality in young larval herring (Clupea harengus); effects of repetitive changes in food availability

Following yolk resorption, laboratory-reared larval Baltic herring (Clupea harengus L.) were exposed to two sequences of food restriction for 5 d and re-alimentation for 10 d. Comparisons regarding larval growth (standard length and content of water-soluble protein), mortality and content of the sum of trypsin and trypsinogen were made with larvae at a continuous high ration. Larvae exposed to varying prey abundance grew less in length than the control, and during the second high-ration period (Day 22 to 32) growth in length ceased. From the first low-ration period onwards, the content of water-soluble protein in these larvae was lower than that of the control larvae, and the survival rate of the low-high ration group was 59% compared to 77% in the larvae at a continuous high ration. In contrast, the effects of varying food availability were minor on larval content of trypsin and trypsinogen. Results are compared with previous findings in larval Clyde herring, and the effects of larval stock and timing and duration of food restriction on larval growth performance are discussed.     

Larval growth and settlement of the European oyster (Ostrea edulis) as a function of food quality measured as fatty acid composition

Larval growth rate and settlement of the European flat oyster Ostrea edulis were experimentally studied as a function of the composition of dietary fatty acids. Diets differing in fatty acid composition were composed by mixtures of the microalgae Isochrysis galbana, Pavlova lutheri and Chaetoceros calcitrans. Fatty acid content in the tissue of the feeding larvae, analyzed by gas chromatography and mass spectrometry, reflected the composition in the diet. Larval growth rate was significantly correlated to the three omega-3 polyunsaturated fatty acids (PUFA) C18:3, C18:4 and C22:6, with minor differences for neutral and polar lipids. No relation between growth rate and the omega-3 PUFA C20:5 was detected, a PUFA often implied as essential for bivalves. It is suggested that naturally occurring variability in fatty acid composition may constrain larval growth. In settlement experiments in both still water and flume flow little substrate selectivity was found for some contrasting substrates. It is concluded that differences in dietary fatty acids may explain as much of settlement success as the variability of substrates. Received: 12 October 1998 / Accepted: 6 April 1999     

Individual variations in fatty acid composition and concentration as indicators of the nutritional condition of wild pointhead flounder larvae

We investigated the fatty acid compositions and concentrations of wild marine fish larvae with a highly accurate method because our knowledge of them has been seriously limited compared with cultured larvae. This study presents estimates of the fatty-acid-based nutritional condition of individual larvae in the field. Because the pointhead flounder Cleisthenes pinetorum displays relatively high stock size fluctuations, we investigated the developmental change in the fatty acid compositions of the body trunk, head, and eye and the annual fluctuations in the fatty acid concentrations in the trunk. We show that the process of fatty acid accumulation is not uniform across body parts and that the trunk is a better indicator of larval nutritional status than other parts because there is less time lag. Starved larvae with simultaneously high docosahexaenomic acid ratios and low total fatty acid concentrations, as observed in laboratory experiments, are rare in the wild. Thus, starved larvae must be removed rapidly by predators before they can experience a relatively long period of starvation in the wild. Fatty acid accumulation was greater in the larvae of the 2005 year class than in those of the 2006 year class in their first feeding stage, according to the optimal model derived with generalized linear model. A previous study indicated that the 2005 year class showed stronger recruitment than the 2006 year class. We conclude that the fatty acid analysis of wild larvae is a useful index of their nutritional status and mortality, especially in the first feeding stage.     

Morphological and histological changes during the growth and starvation of herring and plaice larvae

Reared herring (Clupea harengus L.) and plaice (Pleuronectes platessa L.) were examined for morphological and histological changes during growth and starvation. The growth rate of herring larvae of 0.22 mm/day was less than that reported for wild stock, but this difference was attributed to survival of runts in laboratory. Larval plaice had a growth rate of 0.16 mm/day. The relative condition factor (antilogarithm of intercept of length-weight line) was used to assess condition throughout the larval stages. Starvation resulted in a progressive collapse of the larval body, especially of the ventral body surface around the pectoral girdle of both species (assessed by the pectoral angle) and of the spacing between the organs of the head in herring. There was a breakdown of the herring gut with decreases in epithelial cell height and catabolism of the connective tissue coat and a marked reduction in the transverse sectional area of the plaice liver. The changes in the pectoral angle in both herring and plaice and the eye height to head height ratio in herring should be useful to fishery biologists for assessing nutritional condition, even on board ship.     

Rearing<Emphasis Type="Italic"> Pandalus borealis</Emphasis> (Krøyer) larvae in the laboratory

Northern shrimp Pandalus borealis (Krøyer) larvae hatch in the northern Gulf of St. Lawrence from early May to the end of June, and larval development occurs over a range of relatively cold water temperatures. Because of the long duration of the pelagic phase and the difficulty of sampling all successive larval stages at sea, we used laboratory experiments to assess the effects of water temperature on larval development and growth. In spring 2000, P. borealis larvae were reared from hatching to the first juvenile stages (i.e., stage VI and VII) at three temperatures (3, 5, and 8°C) representing conditions similar to those in spring in the northern Gulf of St. Lawrence. Larval development and growth were dependent on temperature, with longer duration and smaller size (cephalothorax length, CL, and dry mass, DM) at 3°C relative to the 5 and 8°C treatments. There were no significant differences in the morphological characters of the different stages among treatments, indicating that regular moults occurred at each temperature. The results suggest a negative impact of cold temperatures (lower intra-moult growth rates and smaller size) and, possibly, higher cumulative mortality due to longer development time that could affect the success of cohorts at sea. However, CL and DM for stage III and later larvae were smaller than those of larvae identified at the same developmental stage in field locations. It is possible that the diet offered to larvae in this experiment (Artemia nauplii, either newly hatched nauplii or live adults, depending on the developmental stage) was not optimal for growth, even though it is known to support successful P. borealis larval development. In the field, there is the possibility that phytoplankton contributes to the larval diet during the first stages and stimulates development of the digestive glands. Furthermore, the nutritional quality of the natural plankton diet (e.g., high protein content, fatty acid composition) might be superior and favourable to higher growth rates even at lower temperatures.Communicated by R.J. Thompson, St. Johns     

Diet of 0-group stages of capelin (<Emphasis Type="Italic">Mallotus villosus</Emphasis>), herring (<Emphasis Type="Italic">Clupea harengus</Emphasis>) and cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>) during spring and summer in the Barents Sea

Recruitment of capelin in the Barents Sea fail when juvenile herring and cod are abundant and the potential for feeding competition of wild sympatric capelin and herring larvae and small cod juveniles were investigated. The frequency of gut evacuation after capture of capelin larvae were also studied in mesocosms. Small capelin larvae (<35 mm length) fed on small prey including phytoplankton, invertebrate eggs and nauplii, bivalves, other invertebrate larvae and small copepods. Calanus copepodites were only observed in large capelin larvae (>26 mm length). Calanus copepodites were the major food sources for contemporary herring larvae (25–35 mm length) and Calanus and euphausiids were the major prey for small juvenile herring (37–60 mm length) and cod (18–40 mm length). Capelin larvae reared in mesocosms evacuated the guts shortly after capture. Capelin larvae had a smaller mouth and fed on smaller prey than herring and cod of the same length. This implies that the small capelin larvae, in contrast to sympatric small herring and cod, are not tightly linked to the food chain involving Calanus and euphausiids. Thus, exploitative competition between capelin larvae and planktivorous fish that rely on Calanus and euphausiids in the Barents Sea may be relaxed.     

Osmoacclimation in Enteromorpha intestinalis: long-term effects of osmotic stress on organic solute accumulation

The fate of the protease trypsin in intestines of individual herring larvae Clupea harengus L. was studied following digestion of the copepod Acartia tonsa. Trypsin was retained in the intestine during two consecutive pulses of feeding and defaecation of copepods. Quantification of herring trypsin in digested, defaecated copepods showed that ca. 1% of larval intestinal enzyme was defaecated along with 1 to 3 copepods. Following ingestion of a single meal, the level of intestinal trypsin post-ingestion declined to pre-ingestion levels within 1 to 2 d of starvation. All enzyme data thus indicated that trypsin, released in response to ingestion of a meal, was retained. In addition, analysis of fed subgroups of starved larvae clearly indicated that release of trypsin from the pancreas stopped after 6 to 8 d of starvation. As the fish still contained substantial amounts of trypsinogen, the underlying cause might be defective release mechanisms. Daily secretion of trypsin and processes responsible for enzyme retention in the gut are discussed. Assimilation efficiency in herring larvae was estimated for copepodite prey. Average carbon assimilation was 90%.     

Developmental changes in the composition of myofibrillar proteins in the swimming muscles of Atlantic herring,Clupea harengus

Changes in myofibrillar protein composition during development have been investigated in the swimming muscles of the Atlantic herring Clupea harengus L. using a range of electrophoretic techniques. The main muscle-fibre type of larvae, and the fast- and slow-muscle fibres of adult fish were found to contain distinct isoforms of myosin heavy chain (MHC) and myosin light chain 2 (LC2). Larval LC2 was present as a minor component of adult fast-muscle myosin. In contrast, larval and adult fast-muscle myosin appeared to contain identical alkali light chains. Tropomyosin and troponin C were also identical in larval and in adult fast-muscle. All three muscle-fibre types contained unique isoforms of troponin T (TNT) and troponin I (TNI). Larval muscle had multiple isoforms of TNT, some of which may correspond to embryonic forms. It was concluded that although the main muscle-fibre type in larvae shares some myofibrillar proteins with adult fast muscle, it also contains characteristic isoforms of MHC, TNI, TNT and LC2 and therefore represents a distinct fibre type. The particular combination of myofibrillar proteins present at any developmental stage was found to be dependent on the rearing temperature. For example, a higher proportion of embryonic TNT isoforms were present at hatching in larvae reared at 5°C than at either 10 or 15°C. Over a period of 7 d, there was a gradual reduction in the number of TNT isoforms, but the pattern in 5°C larvae after 7 d still did not resemble that in 1 d-old larvae reared at 15°C.