Studies to establish an optimal diet for Carcinus maenas II. Protein and lipid requirements

Carcinus maenas (L.) was maintained on synthetic foods containing various proteins and lipids. Food quality was evaluated in terms of the crustacean's growth, duration of intermoult periods, and time of survival. Casein was found to be an adequate source of protein. Substituting amino acids or hydrolysates for casein results in reduced growth and delayed moults or absence of moulting. Animal proteins like casein are better suited than vegetable proteins. Correspondingly cod liver oil is more adequate than the vegetable lipid maize oil. The amount of salts and the pH-value also influence the quality of food-stuff for C. maenas.     

Variation in condition of coastal Newfoundland 0-group Atlantic cod (Gadus morhua): field and laboratory studies using simple condition indices

Demersal 0-group Atlantic cod (Gadus morhua L.) were collected in Trinity Bay, Newfoundland, Canada, between August 1993 and January 1994. Diet composition and simple indices of condition (hepato-somatic index, condition factors and body ash content) were determined. Laboratory experiments were also conducted to determine whether simple indices of condition reflect differences in energy intake (ration level) and to determine minimum index of condition values, assessed from 0-group cod that died due to exhaustion of their energy reserves. Determination of lower critical indices of condition provided a meaningful interpretation of field data on feeding, condition and survival of 0-group cod. In the wild, indices of condition reflecting variation in liver and muscle energy reserves of cod increased rapidly with the consumption of Calanus finmarchicus, which exhibited a high incidence of oil sacs, and declined abruptly when this lipid-rich prey was no longer consumed. Liver and muscle condition of 0-group cod from the field varied seasonally, but their condition was consistently above that of cod that died in the laboratory. Laboratory studies, using three ration levels, revealed that condition indices differed over time and among ration levels. Indices of condition representing variation in muscle energy reserves exhibited a steady decline once the hepato-somatic index fell below about 2.0. We interpret this as evidence for depletion of lipids first and then proteins, a pattern similar to that of larger juveniles and adults. We conclude that as long as 0-group cod are above a “safe” level of condition their body length can still increase even when condition is declining. Received: 2 September 1997 / Accepted: 8 December 1998     

How does cod (Gadus morhua) cope with variability in feeding conditions during early larval stages?

Response of mesocosm-reared cod (Gadus morhua L.) larvae to different feeding conditions was investigated in 1988 in two mesocosms: a large basin and a smaller bag enclosure within the basin. The basin was filled with seawater, and a community of naturally occurring plankton developed. Plankton concentrations were monitored, and cod larvae stocked in the enclosures were sampled for determination of growth, survival, and gut content. In the bag, insufficient amounts of energetically favourable prey, as copepod nauplii, led to non-selective ingestion of plankton from a broad range of sizes, including considerable amounts of protozoans (tintinnid and oligotrich ciliates). Growth of larvae from the bag was low, with daily specific growth rates (SGR) less than 2.8% the first 3 wk post-hatch. This was followd by rapid increase of SGR to 21.7%, which coincided with a large increase in availability of copepod nauplii. In the basin, high nauplii concentrations led to SGR of 13.7 to 21.7% from onset of feeding to 16 d post-hatch, respectively. Under such conditions, the larvae were highly selective feeders. At 3 wk post-hatch, survival was 36.7 and 38.3% in the basin and bag enclosure, respectively. To cope with variations in the feeding conditions, the cod larvae were shown to be opportunists when nauplii were scarce, and included plankton from several trophic levels in their diet. When nauplii were abundant, cod larvae realized their high potential for growth. Both opportunism and realization of a high growth potential may enhance survival of the larvae.     

Relation of growth and condition with the <Emphasis Type="Italic">Pan</Emphasis> I locus in Atlantic cod (<Emphasis Type="Italic">Gadus morhua</Emphasis> L.) around Iceland

Growth and condition (both somatic and hepatosomatic index) of Atlantic cod spawning at different locations around Iceland was studied in relation to the Pan I locus. South of Iceland cod carrying the Pan I^AB and Pan I^BB genotypes were more frequent while cod carrying the Pan I^AA genotype was more frequent north of Iceland. Differences in growth were detected between cod spawning at different areas around Iceland. Cod spawning south of Iceland grew faster than cod spawning north of Iceland. Differences in growth rate were also observed among cod carrying different Pan I genotypes within a spawning area. The least frequent Pan I genotype expressed the highest growth in both south and north of Iceland. Cod carrying the Pan I^AA grew fastest at spawning locations south of Iceland, while cod carrying the Pan I^BB genotype grew fastest in north of Iceland. A consistent relationship between condition and the different Pan I genotypes was also observed in all the areas. Cod carrying the Pan I^AA expressed the highest somatic condition and the lowest hepatosomatic index. Together, these results indicate that the relation of growth and condition with the Pan I locus is more complicated than earlier thought and is likely to be influenced by other factors, like size-selective fishing and food supply.     

Habitat associations and diet of young-of-the-year Pacific cod (<Emphasis Type="Italic">Gadus macrocephalus</Emphasis>) near Kodiak,Alaska

The influence of environmental variables and habitat on growth and survival of juvenile gadoid species in the Atlantic has been clearly demonstrated; conversely, in the North Pacific little is known about the habitat and ecology of juvenile Pacific cod (Gadus macrocephalus Tilesius). The hypothesis that density of young-of-the-year (YOY) Pacific cod in nearshore habitats is predicted by shallow depth and the presence of eelgrass and macroalgae was tested in a variety of nearshore habitats adjacent to commercial fishing grounds near Kodiak Island, AK. From 10 to 22 August 2002, a beach seine and small-meshed beam trawl were used to capture YOY Pacific cod (n = 254) ranging from 42 to 110 mm fork length. Depth, water temperature, salinity, sediment grain size, and percent cover by emergent structure (i.e. tube-dwelling polychaetes, sea cucumber mounds, macroalgae) were measured prior to fishing. Density of YOY Pacific cod was highest in areas of moderate depth (15–20 m) and positively and linearly related to percent cover by sea cucumber mounds and to salinity. No previous studies have documented fish utilizing sea cucumber mounds as habitat. Furthermore, eelgrass and macroalgae were inconsequential to cod distribution. Diets consisted mainly of small calanoid copepods, mysids, and gammarid amphipods and were significantly related to cod length and depth. This work provides important information on previously undocumented factors that affect distribution and feeding of YOY Pacific cod, which ultimately influence growth and survival in this species.     

Influence of the water-accommodated fraction of no. 2 fuel oil on energetics of Cancer irroratus larvae

The influence of the water-accommodated fraction of No. 2 fuel oil on aspects of larval development of Cancer irroratus (Say) has been investigated. Rates of survival, respiration, ingestion and growth were estimated for both oil-treated and untreated larvae. Larvae reared in oil-treated water exhibited lower survival to megalopa than did larvae reared in untreated water. The presence of oil also altered several aspects of larval energetics. Reductions were seen in food consumption and growth rate while the metabolic cost of maintenance increased. One overall effect that may occur as a result of lowered larval survival and growth rate would be the reduction in larval recruitment in oil-polluted areas.     

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     

Variability in nursery function of tropical seagrass beds during fish ontogeny: timing of ontogenetic habitat shift

Seagrass beds are often considered to be important nurseries for coral reef fish, yet the effectiveness of these nursery functions (refuge and food availability) at different juvenile stages is poorly understood. To understand how the demands of juvenile fish on seagrass nursery functions determines the timing of ontogenetic habitat shifts from seagrass beds to coral reefs, we conducted visual transect survey and field tethering and caging experiments on three different sizes of the coral reef fish Pacific yellowtail emperor (Lethrinus atkinsoni) during its juvenile tenure in seagrass beds at Ishigaki Island, southern Japan. The study showed that although the number of individual L. atkinsoni juveniles decreased by >90 % during their stay in the seagrass nursery, the shelter and/or food availability functions of the nursery, at least for a juvenile size of approximately 5 cm total length (TL), provided the best survival and growth option. The timing of ontogenetic migration to coral reefs of larger fish (>8 cm TL) was attributed to foraging efficiency for larger food items in different habitats. Overall, the function of the seagrass bed nursery changed with juvenile body size, with marginally higher survival and significantly greater growth rates during early juvenile stages in seagrass beds compared to coral reefs. This would contribute to the enhancement in the number of individuals eventually recruited to adult populations.     

Use of kappa-carrageenan microbound diet (C-MBD) as feed for Penaeus monodon larvae

The performance of an artificial practical diet, kappacarrageenan microbound diet (C-MBD) was assessed on Penaeus monodon larvae at the SEAFDEC Broodstock and Maturation Experimental Laboratory in March 1986. Shrimps were reared from zoea₁ to post-larvae₁ using five dietary treatments: (a) natural food — Chaetoceros calicitrans and Artemia salina (b) C-MBD; (c) combination of natural food and C-MBD; (d) commercial diet (microencapsulated, MED); (e) combination of natural food and commercial diet. Results showed slow development with larvae fed the commercial diet. Feeding with C-MBD in combination with natural food resulted in the highest % survival among treatments (69.6), but this was not significantly different (P>0.05) from those obtained with larvae fed natural food alone, C-MBD alone or their combination. While mean values for survival of larvae fed the commercial diet, either alone or in combination, was significantly lower (p<0.05) than all other treatments, their mean growth indices were comparable with larvae fed C-MBD alone or in combination. The low levels of protein, lipid and essentially fatty acids (which are considered important nutrients during larval development) contained in the commercial diet may well justify the results on metamorphosis, survival and growth of the larvae fed this diet. The good performance of C-MBD in this experiment suggests that this kind of diet can be used as partial or total replacement to the traditional algal food.     

Egg survival,embryonic development,and larval characteristics of northern shrimp (<Emphasis Type="Italic">Pandalus borealis</Emphasis>) females subject to different temperature and feeding conditions

Laboratory experiments on ovigerous females of northern shrimp (Pandalus borealis) were used to assess the effects of temperature and food ration on female condition during incubation and examine how combined effects of temperature and female condition influenced egg survival, embryonic development, and larval characteristics. Ovigerous females were maintained at 2°C, 5°C, and 8°C and fed on a low (three times/week; 2–2.7% W/W) or high ration (five times/week at satiation). The increase in temperature accelerated the developmental time of the eggs but their survival at 8°C was reduced. Conversion efficiency of yolk reserves in developing embryos was significantly reduced at elevated temperatures and larvae hatching at 2°C and 5°C were significantly larger and heavier than those hatching at 8°C. The experimental design did not result in any effect of food ration on the energetic condition of females or on egg characteristics and their biochemical composition. However, lower energy reserves were observed for females held at 8°C.     

Decreased lipid storage in juvenile Atlantic cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>) during settlement in cold-water eelgrass habitat

We characterized the prey field and the lipid classes/fatty acids in the flesh of age 0 juvenile cod (Gadus morhua) during their late-summer/fall arrival and settlement into eelgrass (Zostera marina) in coastal Newfoundland. Examination of available prey demonstrated a high abundance of small zooplankton (Acartia, Microsetella and Oithona sp.) with no larger Calanus sp. prey. Breakpoint analysis showed significant changes in the accumulation of relative (mg g⁻¹ wet weight) and absolute (μg fish⁻¹) amounts of lipid with standard length at the time of settlement (~60 mm standard length). Settling juvenile cod showed an alternate lipid utilization strategy where they catabolized phospholipids (PL) to a greater extent than triacylgylcerols (TAG). Polyunsaturated fatty acids (PUFA) content in cod flesh decreased as fish grew indicating that nearshore zooplankton quality was not optimal for PL formation. The dramatic reduction in cod PL was likely due to both catabolism of muscle and a lack of dietary PUFA suitable for PL synthesis. However, juvenile cod continued to grow, leading to decreased lipid stores and suggesting that cod settling into eelgrass are under intense selection pressure for growth prior to the onset of winter, possibly as a means of escaping gape-limited predation. These data contrast better-studied freshwater and estuarine systems in which lipid storage is critical for successful overwintering.     

Some effects of food density on the growth and behaviour of plaice larvae

The effect of food density on the growth, survival, and swimming activity of plaice larvae (Pleuronectes platessa L.) was investigated under controlled laboratory conditions. Suboptimal food concentrations decrease the growth of the height of the body musculature relative to length, and increase the amount of time spent searching for food. Older larvae are able to withstand much longer periods without food than young larvae. On the basis of these experiments, it is suggested that food limitation in a larval plaice population is likely to result in a concave mortality curve.     

Stimulation of root formation in Posidonia oceanica cuttings by application of auxins (NAA and IBA)

At the present, the long period (from 3 up to 12 months) to form roots from rhizome cuttings of Posidonia oceanica is the major cause of transplant failure. To promote earlier rooting, the effects of different concentrations (5 and 10 mg/l) of two auxins, namely indole-3-butyric acid (IBA) and α-naphtalen aetic acid (NAA), and the time of collection on rooting of plagiotropic and orthotropic cuttings of P. oceanica were tested. Rooting, survival and growth of cuttings were assessed 1 month after planting in a mesocosm. Results demonstrated that the use of auxin was essential to achieve root initiation within the observation period. Irrespective of plant source and collection time, IBA and NAA (5 mg/l) treatments increased the rooting capacity in cuttings. The highest rooting success (i.e. percentage of the survived cuttings that rooted) obtained was 33%. Results also revealed that the survival rate and percentages of cuttings with leaf growth and emergence of new leaves on their terminal shoot were influenced by the collection time × plant source interaction, but were unaffected by the auxin treatment. Orthotropic cuttings taken in November and February showed the highest survival rate (100%). No differences in survival among months were detected for plagiotropic cuttings. In May and July, more plagiotropic cuttings survived (80±7 and 75±6%) compared to orthotropic cuttings (58±7 and 51±4%). Overall, more plagiotropic cuttings showed leaf elongation compared to orthotropic ones, but the inverse was observed in November. A higher percentage of cuttings with emergence of new leaves was observed in the plagiotropic type as compared to the orthotropic one, and in general February and November were the best months for leaf production (28±4 and 38±7%). Finally, the percentage of cuttings that had changed leaf growth orientation (from orthotropic to plagiotropic) was significantly higher in November (63±13%) compared to the other months and tended to be higher in cuttings treated with the auxins compared to controls. These preliminary findings indicate that exposure to IBA or NAA (5 mg/l) was effective in stimulating rooting in P. oceanica without inhibiting plant growth. Therefore, this simple pre-planting practice should be beneficial in restoration attempts. Based on the trend observed, rhizomes should be prepared preferably in late autumn–winter and then planted into restoration stands to achieve a prompt rooting response and the best survival rate. Clearly, future work is needed to maximize the rooting success of cuttings.     

Rearing of prawn larvae,Penaeus japonicus,fed soy-cake particles and diatoms

Soy cake^* was homogenized and filtered through an 80 mesh net to produce small, detritus-like particles. The effect of soy-cake particles, an artificial detritus, on the survival and growth of Penaeus japonicus Bate zoea was then tested. The soy cake was considered to be effective for survival and growth of the zoea. The best survival rate, 85.9%, was obtained in a tank receiving soy cake in quantities of 0.16 mg/zoea/day. An experiment was also performed to compare three kinds of food: particles of soy cake alone, diatoms (Chaetoceros rigidus) alone, and a half and half mixture of the two. The best growth, 2.49 mm, and the highest survival rate, 79.1%, were found in larvae fed the mixture of soy-cake particles and diatoms. The results thus seem to demonstrate that particles of soy cake could be usefully applied to the mass-culture of the penaeid prawn.     

Studies on the nutrition of marine flatfish. Growth of the plaice Pleuronectes platessa on diets containing proteins derived from plants and other sources

The growth of 0-group Pleuronectes platessa L. on diets containing plant proteins was examined. Diets were made acceptable to the fish by including in them cod flesh which had been predigested with proteolytic enzymes, as well as a proportion of air-dried cod meal. A leaf-protein concentrate was fed at 6 levels ranging from 0 to 63 % of the total dietary protein; B.P. protein concentrate (Toprina) and soyabean meal were examined at a single level — about 45 % of the dietary protein. The remainder of the protein in these diets was animal protein, cod meal or cod flesh. Analysis of the weight gains after a 12 week feeding period and of the growth curves showed that, as the proportion of leaf protein in the diet increased, the growth rate diminished, the effect becoming more pronounced when leaf protein reached 40 % or more of the dietary protein. Growth on the diet containing B.P. protein concentrate was superior to that on the diet containing soyabean meal and was comparable with diets containing low levels of leaf protein. The protein efficiency ratios (PER= g live weight gain/g protein fed) for the different diets showed similar trends to the growth rates. Diets containing 40 % or less of leaf protein and the B.P. protein diet had PERs of 1.4 to 1.5. The soyabean meal diet and diets containing more than 40 % of the protein as leaf protein had PERs of 1.0 or less. The results are considered in the context of production diets for flatfish.     

Spatial anatomy of species survival: effects of predation and climate-driven environmental variability

The majority of survival analyses focus on temporal scales. Consequently, there is a limited understanding of how species survival varies over space and, ultimately, how spatial variability in the environment affects the temporal dynamics of species abundance. Using data from the Barents Sea, we study the spatiotemporal variability of the juvenile Atlantic cod (Gadus morhua) survival. We develop an index of spatial survival based on changes of juvenile cod distribution through their first winter of life (from age-0 to age-1) and study its variability in relation to biotic and abiotic factors. Over the 25 years analyzed (1980-2004), we found that, once the effect of passive drift due to dominant currents is accounted for, the area where age-0 cod survival was lowest coincided with the area of highest abundance of older cod. Within this critical region, the survival of age-0 cod was negatively affected by its own abundance, by that of older cod, and by bottom depth. Furthermore, during cold years, age-0 cod survival increased in the eastern and coldest portion of the examined area, which was typically avoided by older conspecifics. Based on these results we propose that within the examined area top-down mechanisms and predation-driven density dependence can strongly affect the spatial pattern of age-0 cod survival. Climate-related variables can also influence the spatial survival of age-0 cod by affecting their distribution and that of their predators. Results from these and similar studies, focusing on the spatial variability of survival rates, can be used to characterize species habitat quality of marine renewable resources.     

Conservative maternal care in an iteroparous mammal: a resource allocation experiment

When resources are limited, life history theory predicts a trade-off between growth, reproduction and survival. In summer, lactating females of temperate large herbivores such as the white-tailed deer (Odocoileus virginianus) normally have access to abundant forage but also face the high energetic needs of lactation and recovery from winter mass loss. At high population density, however, females may face a trade-off between allocating resources for maintenance and for reproduction. To simulate the effects of increased intra-specific competition at high density, we measured for 2 years how an experimental food restriction of approximately 20% affected current reproduction and body mass changes of adult females and their fawns during the fawning and lactation periods. Fawn survival decreased 35%, and fawn growth decreased 26% in the food-restricted treatment. There was no effect of food restriction on female mass. Irrespective of treatment, however, lactating females gained 30 g/day less than non-lactating females, and females that had weaned a fawn the previous year gained 20 g/day less than females that had not. We conclude that when resources were scarce, females adopted a conservative strategy favouring their own survival, mass recovery and future reproductive potential over their current reproduction, probably to maximise their lifetime reproductive success.     

Foraging behaviour of larval cod (Gadus morhua) influenced by prey density and hunger

Fish larvae meet diverse environmental conditions at sea, and larval growth and chance of survival depend on a flexible response to environmental variability. The present study focuses on the flexibility of the foraging behaviour of larval cod in a series of laboratory experiments on larval search activity, prey selectivity, and hunger in a variable prey environment. Gadus morhua eggs were collected in March 1992 and 1993 from the Kattegat area, Denmark, fertilised and incubated in the laboratory. After hatching, the larvae were transferred to rearing tanks of 172 litres. The behaviour of larvae (6 to 7 mm long) was observed visually, and prey attacks, swimming activity and gut contents were registered across a range of 1 to 120 copepod nauplii l^-1. When prey density decreased, larvae increased their swimming activity, increased their responsiveness to prey (distance of reaction) and decreased their prey size selectivity. Behavioural response was to a large degree determined by the level of hunger, represented by the number of newly ingested prey in the gut. The findings show that cod larvae have a flexible response to changes in feeding conditions and imply that larvae can grow and survive even in the lower range of (mean) prey densities measured at sea.     

Continued decline of an Atlantic cod population: how important is gray seal predation?

The continental shelf ecosystem on the Eastern Scotian Shelf (ESS) has experienced drastic changes. Once common top predators are a small fraction of their historical abundance, and much of the current community structure is now dominated by pelagic fishes and invertebrates. Embedded within this food web, Atlantic cod and gray seal populations have recently exhibited nearly opposite trends. Since 1984, cod populations have decreased exponentially at a rate averaging 17% per year, whereas gray seals have continued to increase exponentially at a rate of 12%. We reexamined the impact of gray seals on Atlantic cod dynamics using more than 30 years of data on the population trends of cod and gray seals while incorporating new information on seal diet and seasonal distribution. The closure of the cod fishery over 10 years ago allowed for a better estimation of natural mortality rates. We quantified the impact of seals on ESS cod by (1) estimating trends in seal and cod abundance, (2) estimating the total energy needed for seal growth and maintenance from an energetics model, (3) using estimates of the percentage of cod in the total diet derived from quantitative fatty acid signature analysis (QFASA) and of the size-specific selectivity of cod consumed (derived from otoliths collected from fecal samples), and (4) assuming a gray seal functional response. Uncertainties of the model estimates were calculated using the Hessian approximation of the variance-covariance matrix. Between 1993 and 2000, cod comprised, on average, < 5% of a gray seal's diet. Our model shows that, since the closure of the fishery, gray seals have imposed a significant level of instantaneous mortality (0.21), and along with other unknown sources of natural mortality (0.62), are contributing to the failure of this cod stock to recover.     

Predation on euphausiids by cod,Gadus morhua,in winter in Icelandic subarctic waters

The euphausiids consumed by cod (Gadus morhua Linnaeus, 1758) taken in November 1983 and February–March 1984 on feeding grounds in the subarctic shelf area north-west, north-east, and south-east of Iceland have been examined. A total of 2 714 euphausiids occurred in the 1 029 stomachs with food which were analysed; of these 1 640 (60%) could be identified to species. Thysanoëssa inermis (Krøyer, 1846) and Meganyctiphanes norvegica (M. Sars, 1857) were most numerous, constituting 58 and 40%, respectively, of the euphausiids which could be identified. The other euphausiids were, in declining numbers of abundance, T. raschii (M. Sars, 1864), T. longicaudata (Krøyer, 1846), and Nematoscelis megalops G. O. Sars, 1883. The mean number of euphausiids occurring per stomach showed considerable variation between both sampling areas and time of sampling. In each area, the mean number of M. norvegica occurring per stomach was similar at both sampling times, while for T. inermis it was greater in February–March than in November. The mean number of euphausiids occurring per stomach increased with increasing size of cod, being 1 to 2 and 4 to 6 in 10 to 30 cm and 50 to 70 cm cod, respectively. An examination of the diel variation in the occurrence of the euphausiids in the stomachs indicated two peaks, one in the morning before sunrise and another in the evening around and after sunset.