Article

Interannual variations of lipids and fatty acids during larval development of Crangon spp. In the German Bight, North Sea

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Abstract

Eggs and larval stages of the caridean shrimps Crangon crangon and C. allmanni were collected to determine their lipid and fatty acid compositions. Specimens were sampled after the extremely mild winter of 1989/1990 and during the following year, 1991. The fatty acid composition of the zoeal stages of both C. crangon and C. allmanni from 1991 were predominated by the polyunsaturated fatty acids which are typical for marine organisms. In contrast, unsaturated fatty acids were absent in the early larval stages of C. crangon collected in 1990; however, advanced larval stages showed a similar fatty acid composition compared to larvae from 1991. The major phospholipid of eggs and larvae was phosphatidylethanolamine. Eggs contained also phosphatidylcholine and phosphatidylserine, which are probably utilized during embryonic development until hatching because in the early larval stages, only phosphatidylethanolamine was found. In stage VII, again phosphatidylcholine and phosphatidylserine occurred. The lipid and fatty acid composition seems to be an indicator for the survival chances of the planktonic stages. The absence of unsaturated fatty acids may be interpreted as a consequence of unfavorable nutritional conditions during the early developmental phase of the shrimps.

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... Kerfoot 1974, Boddeke 1982, between years (e.g. Kattner et al. 1994, Urzúa et al. 2012, and perhaps over longer time scales (see Reed et al. 2012). Such POI variation at the intraspecific level can provide insights into the evolutionary processes that shaped the macroecology of development modes, which firstly drove local adaptation, speciation, and eventually, the diverse multitude of reproductive and development modes present in the aquatic environment. ...
... Variations in POI are known within populations on temporal scales and have been demonstrated for decapod crustaceans inter-annually (e.g. Kattner et al., 1994), between seasons (e.g. Boddeke, 1982, Sheader 1983, Oh and Hartnoll, 2004, Urzua et al., 2012, and within a single breeding season (e.g. ...
... Such plasticity is known at a range of scales, e.g. inter-annually, between seasons, and within a single breeding season (Boddeke 1982, Sheader 1983, Kattner et al. 1994, Sheader 1996, Oh & Hartnoll 2004, Urzúa et al. 2012, Urzúa & Anger 2013. POI plasticity is influenced by maternal size and age, food quantity and quality, population density, and temperature. ...
Thesis
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Within the marine environment, the diverse development modes among marine invertebrate taxa follow a macro-ecological pattern across latitude. Generally, across the latitudinal gradient from the tropics to the poles, larval development becomes increasingly independent of external food resources. Low latitude species are fecund and produce relatively poorly provisioned eggs, which develop into swimming and feeding larvae. This mode of development becomes increasingly scarce with increasing latitude, whilst development from large eggs (produced in low numbers) into larvae, which are non-feeding, becomes increasingly prevalent. From the tropics to the poles, there is an increasing mismatch between the longer periods required for larval development (resulting from increasing cold) and the shorter periods of food availability (resulting from increasing seasonality). This macro-ecological trend in development modes results from the convergent evolution of diverse taxa, which have adapted to high latitude environments in much the same way: through producing larvae which develop independently of external food resources. Developmental plasticity during the larval phase is a mechanism by which larvae are able to cope better with unfavourable conditions or variations in their environment. Critically, the interaction between per offspring investment (POI; the quantity and quality of resources allocated to offspring) and developmental plasticity, and the role of this interaction in the evolution of larval development modes, is little considered or studied. Experiments comprising this thesis assess the potential role of this interaction in the evolution of abbreviated and lecithotrophic development within decapod crustaceans, and the establishing of the macro-ecological trend in development outlined above. Experiments used the palaemonine shrimp, Palaemonetes varians, which inhabits temperate, salt marsh, and peripheral brackish water, as a study species. Palaemonine Abstract ii shrimp originated from a tropical marine clade and extant species are found in marine, brackish, and fresh water environments. The evolutionary transition from marine to fresh water has involved life history adaptations in development mode within this group. As species occupy differing habitats along this environmental gradient, this group has been used to study evolutionary adaptations along the environmental gradient from marine to fresh water. Sampling of a wild population of P. varians from Lymington salt marsh (Hampshire, UK) revealed the highly variable environment that this species inhabits. POI within this population varied inter- and intra-annually, though these variations could not be correlated with variations in environmental temperatures. Larvae hatch with significant yolk reserves and can be considered facultative lecithotrophic in the first and second larval instar, and planktotrophic from the third larval instar. Larval development was successful at temperatures between 15 and 30 °C and temperature-mediated developmental plasticity was observed; at higher temperatures, larvae increasingly developed through fewer larval instars. Development through fewer larval instars resulted in more rapid development, but development to a lesser juvenile dry weight at settlement. Consequently, this developmental plasticity may have ecological implications. Developmental plasticity was also influenced by the energy content of larvae at hatching (as proxy for POI). Larvae with greater energy content developed through fewer larval instars at all temperatures, indicating that higher POI buffers larvae against poor conditions during development. Greater energy content also enabled larvae to tolerate starvation for longer and to develop to more advanced larval stages in the absence of food. Developmental plasticity within decapod crustaceans enables larvae to tolerate unfavourable conditions during development. Interestingly, it maximises the potential fitness benefits provided by POI by enabling larvae to settle as juveniles earlier, though at a smaller size. The interaction between POI and developmental plasticity forms a ‘pre-adaptation’ for the evolution of abbreviated development. The results presented in this thesis indicate that, indeed, the evolutionary transition to abbreviated development and lecithotrophy is based on selection for increasing POI. The abbreviation of development associated with increasing POI arises through developmental plasticity in larval instar number.
... From earlier studies (Wehrtmann 1991;Kattner et al. 1994;Paschke et al. 2004), it seems essential that first feeding must take place within the first 24 h after hatching to guarantee high survival and development rate. In most of the larval rearing experiments with C. crangon, Artemia nauplii were used successfully without prior enrichment (Table 1). ...
... Pandian (1967) followed the chemical composition of C. crangon during embryogenesis and observed a relative increase in proteins from 58.7% to 69.3% and a decrease in fat from 32.6% to 15.6% per unit of dry weight, indicating that lipids are the main energy source during egg development. Kattner et al. (1994) investigated the fatty acid composition in eggs and several larval stages of C. crangon and observed that the major fatty acids in the eggs were palmitic acid (16:0) and stearic acid (18:0), palmitoleic acid (16:1n-7), vaccenic acid (18:1n-7), oleic acid (18:1n-9), eicosapentaenoic acid (20:5n-3, EPA) and docosahexaenoic acid (22:6n-3, DHA). Other highly unsaturated fatty acids (HUFAs) detected were linoleic acid (18:2n-6) and arachidonic acid (20:4n-6, ARA), although at lower percentages. ...
... Data on the lipid and fatty acid profiles in eggs and larvae of C. crangon are very rare (Table 4). The main n-3HUFAs in the eggs were EPA and DHA, with a EPA/DHA ratio of 2 (Kattner et al. 1994), which seems to be a normal ratio for boreal seashore species (Narciso & Calado 2001). This medium EPA/DHA ratio indicates that C. crangon can be placed at a medium trophic level, as DHA is highly conserved across the food chain. ...
Article
The European brown shrimp, Crangon crangon, is a highly valued commercial species fished in the north-eastern Atlantic, especially the North Sea. The shrimp fisheries are mainly coastal and exert high pressures on the local ecosystems, including estuaries. The culture of the species provides an alternative to supply a niche market (large live/fresh shrimps) in a sustainable manner. However, after more than a century of biological research on this species, there is still little knowledge on its optimal rearing conditions. C. crangon remains a difficult species to keep alive and healthy for an extended period of time in captivity. This review is based on a comprehensive literature search and reflects on the current status of experimental rearing techniques used for this species, identifies the problems that compromise the closing of the life cycle in captivity and provides examples on how these problem issues were solved in the culture of commercial shrimp species or other crustaceans. The ability to consistently produce high-quality offspring could initiate the commercial production of this valuable shrimp. A further advantage of the ability to consistently produce high-quality offspring of this species would facilitate research on the development of new bioassays with this ecologically and economically important species in a wide variety of biochemical and physiological studies.
... Lipids are equally important during critical non-feeding developmental periods such as embryogenesis and metamorphosis (e.g. Anger et al., 1989;Kattner et al., 1994;Petersen and Anger, 1997). In subpolar regions, where productivity occurs in short seasonal pulses (see Knox, 1994), some lithodid crab species have evolved a non-feeding mode of larval development as a special adaptation to strong seasonality of planktonic food availability (Anger, 1996;Shirley and Zhou, 1997;Lovrich et al., 2003;Calcagno et al., in press). ...
... Although lipid deposits are generally the major energy source during non-feeding developmental periods in decapods (e.g. Herring, 1974;Holland, 1978;Amsler and George, 1984;Kattner et al., 1994;Wehrtmann and Kattner, 1998), practically nothing is known about the lipid content and composition of lecithotrophic lithodid crab larvae. Inferring from elemental and protein data, Anger (1996) estimated that lipids comprised about 30% of the dry mass of freshly hatched larvae of L. maja, and that about one half of these lipid reserves were catabolized during non-feeding development to metamorphosis. ...
... In the latter, phospholipids also decreased logarithmically during larval development. A high proportion of triacylglycerols, which is the most important energy source, may thus be typical of lecithotrophic larvae, whereas the lipids in the eggs and larvae of other decapod species are often dominated by phospholipids (Clarke, 1977(Clarke, , 1979(Clarke, , 1993Kattner et al., 1994;Wehrtmann and Graeve, 1998;Graeve and Wehrtmann, 2003). ...
Article
Full-text available
During the larval development of the subantarctic king crab, Lithodes santolla, and stone crab, Paralomis granulosa, we compared changes in the carbon, fatty acid and protein contents of larvae reared under constant conditions from hatching to metamorphosis, either in presence or absence of food (Artemia spp. nauplii). In both species the feeding condition had no influence on any of the chemical parameters studied, indicating a fully lecithotrophic (i.e. non-feeding) mode of development from hatching of the first zoea to metamorphosis of the late megalopa. Dry mass and carbon contents at hatching were similar in the larvae of both species, but L. santolla contained initially higher total amounts of fatty acids and protein than P. granulosa. Both species utilized considerable portions of their total fatty acid pool which decreased logarithmically throughout the time of development. At metamorphosis, it was almost exhausted in P. granulosa, while L. santolla had consumed only about 60%. Protein utilization, in contrast, was higher in L. santolla (40%) than in P. granulosa (20%). Triacylglycerol was the principal storage lipid in both species, accounting initially for about 75% of the lipid fraction; it was strongly utilized during larval development. Phospholipid constituted the second largest lipid class; it also decreased in P. granulosa, but to a lesser extent in L. santolla. The major fatty acids of both species were 18:1(n−9), 20:5(n−3) and 16:0 as well as, in lower proportions, 18:1(n−7), 22:6(n−3), 16:1(n−7) and 18:0. Monounsaturated fatty acids represented the dominant group in L. santolla, whereas P. granulosa contained similar amounts of mono- and polyunsaturated fatty acids. In L. santolla, monounsaturated fatty acids, especially 16:1(n−7), were preferentially utilized as compared to polyunsaturates. Due to a particularly strong lipid utilization in P. granulosa, all individual fatty acids were largely depleted at metamorphosis, showing similar extents of consumption. L. santolla had higher initial lipid and protein stores that seem to be used more economically as compared to P. granulosa.
... Until now, few studies have examined the lipid composition of crustacean eggs (e.g. Dawson & Barnes 1966, Pandian 1970a, Achitiv & Barnes 1976, Amsler & George 1984, Biesiot & Perry 1995 and even fewer have looked at the FA depletion during embryonic development (Clarke et al. 1990, Kattner et al. 1994, Wehrtmann & Graeve 1998, Wehrtmann & Kattner 1998, Nates & McKenney 2000, Narciso & Morais 2001. A lot of research has been conducted to determine the essential FA requirements of crustacean species with aquaculture potential; a great deal of this research has focused on the quantitative and qualitative lipid requirements of commercially important species and relatively few studies have examined other crustacean species (D'Abramo & Sheen 1993, Fox et al. 1994. ...
... Palaemon elegans ovigerous females were captured with dip-nets in the intertidal zone of Cape Raso, Cascais, Portugal. The egg mass was removed from the females and eggs were classified according to the following criteria (modified from Kattner et al. 1994): (1) Stage 1 -uniform yolk and no embryonic development visible; (2) Stage 2eyes clearly visible with 1/2 yolk consumed; and (3) Stage 3 -almost no yolk present and embryo fully developed. ...
... Consequently, P. martia martia and P. serratus eggs have higher DHA:EPA ratios (1.1 and 0.6 at the start of the embryonic development, respectively) than P. elegans (0.2). In fact, with increasing depth (i.e. increase in pressure and decrease in temperature), there appears to be an increase in the DHA:EPA ratio of the eggs, as a result of the rise in the DHA level and a reduction in the EPA content (Kattner et al. 1994). ...
Article
The present study examines the changes in volume and lipid biochemistry during the embryonic development of 3 temperate caridean species - Plesionika martia martia, Palaemon serratus and P. elegans-with similar reproductive strategies but occupying different ecological niches. Egg volume, water content and lipid embryonic metabolism are analysed and discussed in relation to early life history and environmental conditions. An increase in egg volume and water content during embryogenesis was noted in all species, although it was larger in P. serratus. F serratus also had the largest eggs, followed by P. elegans and P. martia martia. The quantitatively most important fatty acids (FA) in the eggs are the saturates (SFA) 14:0, 16:0 and 18:0, the monounsaturates (MUFA) 16:1(n-7), 18:1(n-9) and 18:1(n-7), and the polyunsaturates (PUFA) 18:2(n-6), 18:3(n-3), 20:4(n-6), 20:5(n-3) (eicosapentaenoic acid, EPA) and 22:6(n-3) (docosahexaenoic acid, DHA). Looking at the predominant FA, there appears to be a higher similarity between P. martia martia and P. serratus. The eggs of these species present similar levels of SFA, highly unsaturated (HUFA) and (n-3) FA but P. martia martia eggs have an extremely elevated MUFA and a low PUFA content. P. elegans has high levels of SFA, PUFA, HUFA and (n-3) FA. R serratus and particularly R martia martia eggs are characterised by a higher DHA:EPA ratio than P. elegans. The similarity between the FA profile of P. martia martia and P. serratus eggs may suggest that these 2 species are exposed to more comparable environmental conditions than P. elegans. As for the utilisation of FA classes during embryonic development, all species showed the same trend - MUFA were found to be the major energetic fuel during embryonic development while SFA and HUFA seemed to be conserved. A steady decrease in total lipids, particularly tri- (TAG) and diacylglycerol (DAG), and FA contents was noted in the 3 species. The utilisation of these lipid classes during the incubation period was comparatively low in P, martia martia and P. elegans in relation to P. serratus eggs. This may suggest a higher dependence of the newly hatched larvae of P. martia martia and P, elegans on their lipid reserves and has been interpreted as an adaptation to the early life history of these species, during which there might be a reduced availability of food.
... cellular membranes, nervous system) of vital importance during ontogeny (Kayama et al., 1980;Bell & Dick, 1990;Beltz et al., 2007). Also, during the initial stages of the life cycle, individuals are often exposed to periods of famine and/or planktonic food shortages, and lipid reserves accumulated internally are crucial for survival and successful growth (Kattner et al., 1994Rosa et al., 2007;Urzúa & Anger, 2013). ...
... Some types of structural fatty acids may help crustacean larvae to present a greater degree of flexibility of their jaw that would allow them to capture a wide spectrum of prey in marine environments with strong seasonality of temperature and food availability (see Anger, 2001;Liddy et al., 2005;Pandian, 2016). Similar responses have been described for crustacean species inhabiting coastal areas with marked seasonality of environmental conditions (Kattner et al., 1994(Kattner et al., , 2007Calado et al., 2013). Crustaceans, in general, require a large amount of longchain polyunsaturated fatty acids (lcPUFA; FA with more than 20 carbons) for various important physiological processes (e.g. ...
Article
Pleuroncodes monodon , an important fishery resource and key species in the Humboldt Current Large Marine ecosystem, has a prolonged reproductive period from winter until end of summer, and during this time females incubating their embryos are exposed to seasonal variation in food availability and in temperature. Additionally, in order to ensure successful reproduction and survival of embryos, changes occur in the main internal reserves and/or sources of energy of P. monodon . The aim of this study was to determine the extent of seasonal variation (winter vs summer) in the lipid content and fatty acid composition of ovigerous females and their embryos. The results show that a higher percentage of saturated and polyunsaturated fatty acids are found in females in winter. Similarly, the composition of fatty acids in embryos found here indicates that winter embryos have more saturated fatty acids and essential fatty acids (C18:2n6cis, C18:3n6 and C22:6n3) than do summer embryos. According to PCA analysis of fatty acid profile, samples from summer may be distinguished into two isolated groups with conspicuous variations in fatty acids profile of embryo and hepatopancreas. While in winter, the opposite pattern occurs in the fatty acid profile of embryo and hepatopancreas. These variations may be related to relevant physiological processes (reproduction and growth) and of their ontogeny (development and survival of offspring). Seasonal variation in the lipid content and composition of fatty acids of P. monodon could directly impact this species’ reproduction and survival and subsequently could have consequences on the food web and fishery exploitation.
... Phyllosoma have been reared on Artemia and mussel gonad with varying success (Illingworth et al. 1997, Kittaka 1988, however, appropriate nutrition requires the identification of the essential elements within the diet (Kanazawa and Koshio 1994). Lipid has been found to be of prime importance in crustacean larval stages (Kattner et al. 1994, Sasaki et al. 1986), and appears to be the main storage product in late stage phyllosoma, which is then used as an energy source during the nonfeeding puerulus stage (Jeffs et al. 1999. ...
... However, in starved stage II larvae, MUFA significantly decreased whereas PUFA significantly increased, possibly indicating a preferential use of MUFA during starvation and a sparing or retention of PUFA. Kattner et al. (1994) found the FA profile of larval caridean shrimp also showed major differences dependent on stage and larval sample. The variation in results in the present study suggest a response for P. cygnus which is dependent on the stage of the larvae. ...
Article
Full-text available
Total lipid, lipid class, and fatty acid analyses were conducted on fed and starved stage I and II phyllosoma of the western rock lobster Panulirus cygnus. In both stages, the decrease in dry mass of starved larvae and increase in dry mass of Artemia-fed larvae were accompanied by a decrease and increase in lipid content, respectively. Lipid accounted for 6.7% of the decrease in dry mass in starved stage I larvae, which increased to 35.0% in stage II larvae. Also, lipid accounted for 6.2% of the increase in dry mass of fed stage I larvae, increasing to 19.2% in stage II larvae. The major lipid classes in all phyllosoma samples were polar lipids (84.1-94.3%) followed by sterols (6.6-12.1%; mainly cholesterol). Gravimetrically, fed larvae increased predominantly in polar lipid whereas in starved larvae, polar lipid was the major lipid class catabolized, with the sterol content not changing significantly. Hydrocarbons, wax esters, diacylglyceryl ether, triacylglycerols, and free fatty acids were all minor lipid classes (<5% of total lipid). Fatty acid analysis showed six major components present; 16:0, 18:1n-9, 18:0, 20:4n-6 (arachidonic acid; AA), 20:5n-3 (eicosapentaenoic acid; EPA) and 22:6n-3 (docosahexaenoic acid; DHA). These fatty acids all increased gravimetrically in fed larvae and decreased in starved larvae. In starved larvae, small decreases were seen in the relative contribution of EPA, DHA, 16:1n-7, and 18:1n-9, with AA increasing. In fed larvae, most of the major fatty acids remained at a similar relative level, and larvae were able to accumulate AA and EPA, but not DHA, above the relative level (%) in Artemia. The results are useful in the identification of nutrients required during development and as such with the design of diets used in phyllosoma culture.
... As larvas dos crustáceos decápodes dispõem de requisitos nutricionais que variam intra e inter-especificamente, assim como sazonal e anualmente . Nos primeiros estádios de desenvolvimento, estes requisitos estão dependentes dos aspectos qualitativos e 6 quantitativos das reservas armazenadas no embrião (Kattner et al.,1994). verificou que na maioria das espécies de decápodes as reservas vitelinas existentes nos embriões, provenientes do investimento parental, são catabolizadas quase na sua totalidade durante o desenvolvimento embrionário. ...
... Newly hatched decapod crustacean larvae are known to display dietary requirements varying intra and interspecifically, seasonally and annually . Such dietary requirements are dependent on the qualitative and quantitative aspects of energetic reserves stored in the egg yolk (Kattner et al., 1994). In general, since yolk reserves are rapidly catabolized, decapod larvae must start feeding immediately after hatching and avoid the deleterious effects induced by starvation. ...
... The food pathway in the pelagic marine ecosystem originates in the phytoplankton whose organic constituents are transformed by animals through food webs and different trophic levels (Nielsen and Hansen, 1995; Kiørboe, 2001). Lipid classes and fatty acid compositions generally characterise marine species or groups of organisms and they can be used to assess nutritional condition, survival expectations and trophic relationships at the different developmental stages (Lee et al., 1971a; Sargent and Whittle, 1981; Sargent et al., 1987; Sargent and Falk-Petersen, 1988; Kattner et al., 1994; Anger, 1998; Harding and Fraser, 1999; Reuss and Poulsen, 2002). Lipid biomarkers have been found to be useful in the analyses of predator-prey relationships in food web studies (Graeve et al., 1994a,b; Kharlamenko et al., 1995; St. John and Lund, 1996; Graeve et al., 1997; Desvilettes et al., 1997; Falk-Petersen et al., 2000; Hamm et al., 2001; Arts et al., 2001). ...
... In tiger prawn (Penaeus esculentus), Dall et al. (1992) found no clear evidence that 20:5n-3 and 22:6n-3 are essential and the essential 20:4n-6 could be synthesized from 18:2n-6. However, for larval shrimp (Crangon spp.) low amounts of PUFA may be interpreted as a consequence of unfavourable nutritional conditions (Kattner et al., 1994). ZI larvae from a coastal and a fjord station in May 2000 both had high larval abundance, low TG condition, lower proportions of 16:0, 16:1n-7, 20:5n-3, 22:6n-3, and higher proportions of 18:0, compared to offshore collected ZI larvae. ...
Article
Full-text available
Lipid class and fatty acid compositions were determined in shrimp larvae (Pandalus borealis and P. montagui) collected along transects across banks on the West Greenland shelf in June 1999, May and July 2000. The lipid class contents were investigated as indices of larval shrimp lipid condition and food type. Fatty acid compositions were investigated for lipid biomarkers to establish trophic relationships between larval shrimp and potential prey. Phospholipids were the dominant lipid class in all six pelagic development stages of larval P. borealis and P. montagui, accounting for 80 to 92% of the total lipid. In all six stages the contents of free fatty acids were more abundant than triacylglycerol. With increasing larval size from hatching to a carapace length of ~2.3 mm, there was a decreasing trend in the contents of hydrocarbons attributed to reduced amount of phytoplankton in the diet. Triacylglycerol (TG) content to wet weight ratio was investigated as index of nutritional larval condition and survival potential. Proportions of larvae with TG indices >0.2 were relatively high in May 2000 and June 1999, but generally low in July 2000. The fatty acids 16:0, 16:1n-7, 18:0, 18:1n-9, 18:1n-7, 20:5n-3, and 22:6n-3 were major lipid components of the larvae and their mean proportions varied within relatively small ranges between larval stages. The dominant fatty acids were 16:0, 20:5n-3, and 22:6n-3 with average proportions of about 17%, 20%, and 13%, as percentage of total fatty acids. Biomarkers for algae (16:0, 16:1n-7, 18:1n-7, 20:5n-3, diatoms: elevated 16:1n-7/16:0 ratios), flagellates (18:0, 18:4n-3, 22:6n-3), and zooplankton (18:1n-9, 20:1n-9, 22:1n-11) indicate that larval P. borealis and P. montagui are omnivorous. ZI larvae in high concentrations at coastal and fjord stations in May 2000 and most larvae (ZIV–ZVI) from July 2000 were low in lipid. Larvae from June 1999 had relatively high lipid contents probably due to better feeding and growth history. Our results suggest variability in growth and survival expectations of larvae related to variations in larval food availability between years.
... The food pathway in the pelagic marine ecosystem originates in the phytoplankton whose organic constituents are transformed by animals through food webs and different trophic levels (Nielsen and Hansen, 1995; Kiørboe, 2001). Lipid classes and fatty acid compositions generally characterise marine species or groups of organisms and they can be used to assess nutritional condition, survival expectations and trophic relationships at the different developmental stages (Lee et al., 1971a; Sargent and Whittle, 1981; Sargent et al., 1987; Sargent and Falk-Petersen, 1988; Kattner et al., 1994; Anger, 1998; Harding and Fraser, 1999; Reuss and Poulsen, 2002). Lipid biomarkers have been found to be useful in the analyses of predator-prey relationships in food web studies (Graeve et al., 1994a,b; Kharlamenko et al., 1995; St. John and Lund, 1996; Graeve et al., 1997; Desvilettes et al., 1997; Falk-Petersen et al., 2000; Hamm et al., 2001; Arts et al., 2001). ...
... In tiger prawn (Penaeus esculentus), Dall et al. (1992) found no clear evidence that 20:5n-3 and 22:6n-3 are essential and the essential 20:4n-6 could be synthesized from 18:2n-6. However, for larval shrimp (Crangon spp.) low amounts of PUFA may be interpreted as a consequence of unfavourable nutritional conditions (Kattner et al., 1994). ZI larvae from a coastal and a fjord station in May 2000 both had high larval abundance, low TG condition, lower proportions of 16:0, 16:1n-7, 20:5n-3, 22:6n-3, and higher proportions of 18:0, compared to offshore collected ZI larvae. ...
Data
Full-text available
Lipid class and fatty acid compositions were determined in shrimp larvae (Pandalus borealis and P. montagui) collected along transects across banks on the West Greenland shelf in June 1999, May and July 2000. The lipid class contents were investigated as indices of larval shrimp lipid condition and food type. Fatty acid compositions were investigated for lipid biomarkers to establish trophic relationships between larval shrimp and potential prey. Phospholipids were the dominant lipid class in all six pelagic development stages of larval P. borealis and P. montagui, accounting for 80 to 92% of the total lipid. In all six stages the contents of free fatty acids were more abundant than triacylglycerol. With increasing larval size from hatching to a carapace length of ~2.3 mm, there was a decreasing trend in the contents of hydrocarbons attributed to reduced amount of phytoplankton in the diet. Triacylglycerol (TG) content to wet weight ratio was investigated as index of nutritional larval condition and survival potential. Proportions of larvae with TG indices >0.2 were relatively high in May 2000 and June 1999, but generally low in July 2000. The fatty acids 16:0, 16:1n-7, 18:0, 18:1n-9, 18:1n-7, 20:5n-3, and 22:6n-3 were major lipid components of the larvae and their mean proportions varied within relatively small ranges between larval stages. The dominant fatty acids were 16:0, 20:5n-3, and 22:6n-3 with average proportions of about 17%, 20%, and 13%, as percentage of total fatty acids. Biomarkers for algae (16:0, 16:1n-7, 18:1n-7, 20:5n-3, diatoms: elevated 16:1n-7/16:0 ratios), flagellates (18:0, 18:4n-3, 22:6n-3), and zooplankton (18:1n-9, 20:1n-9, 22:1n-11) indicate that larval P. borealis and P. montagui are omnivorous. ZI larvae in high concentrations at coastal and fjord stations in May 2000 and most larvae (ZIV–ZVI) from July 2000 were low in lipid. Larvae from June 1999 had relatively high lipid contents probably due to better feeding and growth history. Our results suggest variability in growth and survival expectations of larvae related to variations in larval food availability between years.
... The relationship between clutch size and egg size may reflect the developmental mode and serves as an indicator of the energy content of the eggs (Herring 1974; Jaeckle 1995). Triacylglycerols dominate the larger eggs and phospholipids the smaller ones (Herring 1974; Clarke 1977a, b; Kattner et al. 1994). Wehrtmann and Graeve (1998) found a substantially higher amount of lipids in eggs of tropical benthic shrimps compared to benthic caridean shrimps of temperate waters. ...
... The fatty acid composition of C. antarcticus eggs is in good agreement with data reported by Clarke (1977b), although no such data exist for Notocrangon antarcticus and Nematocarcinus lanceopes. The same major fatty acids occurred in all species and resembled closely those of other decapod eggs (Clarke 1977aClarke , b, 1979Clarke , 1993 Hopkins et al. 1993; Kattner et al. 1994; Graeve et al. 1997; Wehrtmann and Graeve 1998; Wehrtmann and Kattner 1998). However, the proportions of the fatty acids were different in the three species. ...
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Eggs of the decapod shrimps, Chorismus antarcticus, Nematocarcinus lanceopes and Notocrangon antarcticus were taken to analyse their morphometric, lipid and fatty acid composition. Almost all females carried undifferentiated eggs (stage I). The average number of recently extruded eggs was lowest in C. antarcticus with 163 eggs, and highest in Nematocarcinus lanceopes with 1,220 eggs, while Notocrangon antarcticus produced on average 350 eggs. The lipid content (% of dry mass) of the eggs was 18.8% for C. antarcticus, 14.3% for Notocrangon antarcticus and 18.1% for Nematocarcinus lanceopes. Dominant lipid classes in eggs of all species were phospholipids and triacylglycerols. The storage lipid, triacylglycerol, was slightly elevated in the eggs of Nematocarcinus lanceopes (mean of 52.7%) compared to the other two species. The fatty acid compositions of the decapod eggs were similar in the three species. Dominant fatty acids were 20:5(n-3), 18:1(n-9), 16:1(n-7), 16:0 and 18:1(n-7), comprising 75.8-78.4% of total fatty acids. Among the species, the eggs of C. antarcticus had the highest proportions of polyunsaturated fatty acids (39.6%), dominated by 20:5(n-3), and the lowest percentage of monounsaturates (41.8%). The eggs of Nematocarcinus lanceopes and Notocrangon antarcticus contained almost the same proportion of polyunsaturated fatty acids (28.0% and 28.4%, respectively), whereas Nematocarcinus lanceopes had the highest amount of monounsaturates due to the dominance of 18:1(n-9). Based on our findings, we assume that eggs produced by polar decapod crustaceans do not contain substantially more lipids than related species from temperate or tropical regions. However, additional studies are necessary to substantiate any general conclusion about the relationship of egg lipid content and composition with climatic zones.
... Embryo size increases with latitude and decreases with average water temperature (Thatje et al. 2004). Initial embryo size also varies intra-speciWcally between latitudinally or geographically separated populations (Lardies and Wehrtmann 2001), between diVerent reproductive seasons (Boddeke 1982), and on a year-to-year basis in the same locality (Kattner et al. 1994). In the present study, we compare embryo size, water content and FA dynamics during embryogenesis of three deep-sea pandalid shrimps (Decapoda, Caridea, Pandalidae) and two portunid swimming crabs (Decapoda , Brachyura, Portunidae). ...
... 1) with shrimp traps (at 600–700 m depth) on board a small trap-Wshing vessel of the " Museu Municipal do Funchal—História Natural. " The embryo mass was removed from the females and embryos were classiWed according to the following criteria (modiWed from Kattner et al. 1994 ): stage 1—uniform yolk and no embryonic development visible; stage 2—eyes clearly visible with ½ yolk consumed; stage 3—almost no yolk present 123 and embryo fully developed. To determine embryo volume , 30 embryos were taken from each female (nine females, three per embryonic stage) and length and width were measured under a stereomicroscope (Olympus ® , model SZ6045TR) with a calibrated micrometer eyepiece. ...
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Variations in embryo size and fatty acid (FA) dynamics during embryogenesis were evaluated in deep-sea pandalids and portunid swimming crabs from the Portuguese continental margin and Madeira Island slope and compared with previous data on neritic and deep-sea lobsters and shrimps (collected between February 2001 and March 2004). Inter-specific variations in embryo size seem to be dictated primarily by phylogeny rather than by differences in reproductive or early life history traits. FA reserves were significantly correlated with embryo size (P<0.001). Principal component analysis revealed differences among three groups (1—neritic caridean shrimps, 2—deep-sea pandalids of the genus Plesionika, and lobsters, 3—portunid crabs and the deep-sea pandalid Chlorotocus crassicornis, Costa 1871). Group 1 was clearly separated by PC1 mainly due to the higher percentage of essential C18 (linoleic and linolenic acids) and C20 (namely eicosapentaenoic) polyunsaturated FA (specific markers of primary producers). PC2 separated Group 2 from Group 3 due to differences in the percentage of several saturated FA (including odd-numbered FA—bacterial markers) and C18 monounsaturated FA (namely 18:1n−9, a general marker of carnivory). Therefore, these differences among groups seem to result from distinctions in diet and ecological niche. Intra-specific differences in FA composition between western and southern Plesionika martia martia (A. Milne-Edwards, 1883) populations may reflect higher water temperatures on the south sub-tropical coast. Lobster embryonic development was more demanding of lipid energy than that of the other decapod species, which may reflect an evolutionary trend in decapod taxa related to an increasing degree of lecithotrophy. However, a lower FA catabolism can be interpreted as an enhanced independence of the newly hatched larvae from external energy sources. Higher FA content at hatching and, as a consequence, a greater independence from the external environment should increase the chances of larval survival.
... This seasonal difference in energy allocation per offspring, provides winter larvae with greater resistance to feeding stress (Paschke et al., 2004). It is possible that the composition of Zoea 1 is more variable between breeding years than between laying seasons (Kattner et al., 1994b), reflecting conditions (e.g. temperature, food availability, phytoplankton bloom) during vitellogenesis and egg incubation (Viegas et al., 2012;Urzúa and Anger, 2013;Mika et al., 2014) or genetic effects (Wu et al., 2010;Tropea and Greco, 2015;Marciano et al., 2018). ...
Article
Seasonal variations in environmental conditions determine the success of decapod larval development, and females transmit more energy in sub-optimal conditions to maximise the fitness of their offspring. The objective of this study was to focus on the combined effects of temperature (14, 18 and 22 °C) and food quality on the performance of larvae produced by 5 young (0+) and 5 old (I+) Palaemon serratus females. We prepared 3 diets based on Artemia, in decreasing order of total fatty acid content: freshly hatched nauplii (N), unenriched met-anauplii (M) and metanauplii enriched with a mixture of microalgae (ME). At hatching, the larvae produced by I+ females had a higher biomass but a similar fatty acid concentration to those produced by 0+ females. Larvae survived better and developed relatively faster as temperature increased, and the longer they waited to meta-morphose, the greater their weight at metamorphosis. These performances were diet-dependent, with more survival and more growth in less time with diet N than with the other two. Larvae from I+ females performed better than those from 0+ females, especially under the most stressful conditions. The greater biomass of the larvae of I+ females seems to have enabled them to follow a shorter, and therefore faster, development path than those of 0+ females. The larvae's diet also had an impact on post-metamorphic composition: larvae eating a diet richer in fatty acids produced richer juveniles and those eating a poorer diet produced juveniles with slightly more essential fatty acids. This study supports the high plasticity of caridean shrimp larval development and the importance of maternal effects on the fitness of offspring.
... The dietary requirements of the first larval stage may vary inter-and intraspecifically, as well as seasonally or interannually. This depends on energy reserves and essential nutrients stored in the egg yolk, and on larval abilities to synthesize macromolecular nutrients from precursors (Kattner et al., 1994;Staton & Sulkin, 1991). Given the rapid depletion of yolk reserves, the larvae of most marine invertebrates and fish species need to start feeding within a limited time after hatching to avoid starvation. ...
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Macrobrachium macrobrachion is an African native brackish river prawn with a high commercial value. Currently, there is little information on the post-larval production of this species. Two experiments were conducted in the laboratory to develop production techniques for this species. The first experiment analyzed the duration of yolk resorption and the second described the larval stages. Yolk resorption was studied in 240 newly hatched larvae for 24 h based on the reduction in yolk area over time. For larval development stages, six breeding tanks containing 100 L with a density of 50 larvae/L were used. Larvae were fed a combination of Artemia nauplii, Brachionus plicatilis, and pelleted food (Larviva ProStart, Biomar Efico). The results have shown that the area of yolk reserves varied significantly in the hours after hatching. At 14 h after hatching, each larva resorbed approximately 85% of its yolk reserve, and at 18 h after hatching, each of them still had approximately 6.1%. Twelve larval stages were identified and described in three critical stages. M. macrobrachion larvae are lecithotrophic and need to start exogenous feeding for 14 h at the earliest and 18 h at the latest after hatching. These results are the first to highlight the potential for mass production of brackish river prawn prawns.
... These early larvae apparently did not develop into late stages and juveniles because late stages were lacking in the plankton samples until June 1990. Kattner et al. (1994) suggested malnutrition of larvae, because the early stages sampled in 1990 showed deficiencies of unsaturated fatty acids and a lack of triacylglycerols, which are important energy stores of crustaceans. ...
Article
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Crangon crangon is a key species in the southern North Sea and the most valuable target of coastal fisheries. Recruitment and stock development are highly variable. As recruitment is based on larval production, we determined the timing of larval release in relation to the annual temperature course and the winter water temperature. The annual temperature courses over 50 years (1961–2010) showed a strong inter-annual variation but also a continuous increase in winter water temperatures. The share of ovigerous C. crangon females starts to increase in October/November, reaches a maximum in May, and decreases continuously towards September/October. During cold winters (0°C), egg development is strongly retarded and hatching of larvae culminates in a sharp peak in early June. Warm winters (6°C) facilitate embryogenesis and the earliest larvae hatch in January and February. The temperature sums from November to April correlated significantly with the lowest winter water temperatures. Consequently, the lowest winter water temperatures provide a good indication for the appearance of C. crangon larvae and an estimate for the match or mismatch with the spring bloom and the production of microzooplankton, the preferred food of C. crangon larvae.
... In marine invertebrates, proteins and lipids play an important role as the main energy reserves or as cell membrane components (Kattner et al. 1994, García et al. 2002, Lee et al. 2006, Jimeń ez & Kinsey 2015. During stressful environmental conditions (e.g. ...
Article
The brown shrimp Crangon crangon is a key component of the North Atlantic coastal food web and an important target species for the fishery economy. As the brown shrimp contains large amounts of protein and essential fatty acids, its consumption makes it a beneficial choice for humans. Commercially harvested crustaceans like C. crangon are frequently affected by bacterial shell disease, with necrotizing erosions and ulcerations of the cuticle. To determine whether shell disease influences the nutritional value of C. crangon, total protein and lipid contents, as well as fatty acid compositions of muscle tissue and hepatopancreas, together with the hepatosomatic index, were examined in healthy and affected individuals. The biochemical composition of the tissues did not differ significantly between the 2 groups. Also, the hepatosomatic index, as an indicator of energy reserves in shrimps, was similar between healthy and affected animals. Our results indicate that the nutritional value of C. crangon is not affected by shell disease, as long as it remains superficial as in the present study.
... For example, the distribution of the European anchovy over a wide latitudinal range (northern Europe to South Africa) strongly correlates with mtDNA variation (i.e., clade frequency), suggesting strong local adaptation and high correlation with the temperature difference over the whole species' range (Silva et al. 2014). For several marine invertebrates, temporal variation in the size of offspring has been demonstrated to occur between years and seasons, as well as within a single breeding season (Kattner et al. 1994;Oh and Hartnoll 2004;Urzúa et al. 2012). ...
Article
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Maternal investment (MI), the energy allocated by mothers to offspring, has important effects on the life-history traits of marine organisms. Variation in such traits shows strong correlation with latitude for several marine taxa (Thorson’s rule). Large-scale latitudinal variation in MI within a single species suggests population genetic divergence, while temporal changes in MI, rather, reflect plasticity. At higher latitudes (i.e., colder waters), traits associated with MI (brood weight, fecundity, egg volume, and energy content) increase. To identify phenotypic plasticity along a latitudinal gradient in MI traits (brood weight, egg volume, density number, and egg lipid composition), five populations of the kelp crab Taliepus dentatus along the coast of Chile (30°S–42°S) were investigated during the summer (December–February) and winter months (June–August) of 2015–2016. Despite this wide latitudinal range, the sea surface temperature (SST) difference between the northernmost and the southernmost sites was only approximately 2.0 °C in winter and 5.5 °C in summer. In summer, when latitudinal variation in SST was highest, brood weight, egg density, fecundity, and egg lipids increased with latitude, while egg volume decreased. No trends in MI were observed in winter when the SST gradient was almost non-existent. These results suggest that the relationship between MI and latitude is shaped by temperature rather than being site-specific. The seasonality of latitudinal MI traits also suggests a trade-off between the costs of female maintenance and/or brooding behaviours and MI. When investigating latitudinal and temporal variation in marine brooder MI, the effect of temperature on life-history traits and the associated costs of female brooding should be quantified.
... The capacity of crustacean larvae to tolerate temporary periods of starvation is therefore believed to be essential for their survival in the pelagic environment, particularly in nutritionally unstable habitats (Anger 2001). In newly hatched larvae, tolerance to starvation as well as dietary requirements are determined by energy reserves and essential nutrients stored in the egg yolk (Kattner et al. 1994;Urzúa & Anger 2011). The availability of egg yolk reserves may, however, vary intra and interspecifically depending on seasonal criteria (Paschke et al. 2004;Gebauer et al. 2010;Urzúa & Anger 2013), maternal factors (Marshall et al. 2008;Moland et al. 2010) and the environmental conditions prevailing during the reproductive period (Giménez 2010;Urzúa & Anger 2013). ...
Article
One of the key factors affecting larval survival is food availability. This study investigated the influence of starvation on the nutritional condition of zoea I larvae of Pleuroncodes monodon. Experimental treatments with differential periods of starvation and subsequent feeding (point of no return: PNR) in addition to treatments with differential periods of feeding and subsequent starvation (point of reserve saturation: PRS) were used to quantify larval survival and the occurrence of lipid droplets in the hepatopancreas. Larval survival differed significantly depending on the starvation and feeding treatment administered. A high percentage of survival was found for the starvation treatment until day 1 (S1-PNR), for the feeding treatment until day 4 (F4-PRS), and for the continuously fed control groups (FC). Survival was minimal for the starvation treatment lasting until day 7 (S7-PNR) and for the continuously starved control groups (SC). In turn, similar tendencies were observed in the utilization of energy reserves; the lipid droplets significantly decreased throughout the PNR treatment, while the presence of lipid droplets gradually increased during the PRS treatment. All these larval condition parameters can be used in fishery models of population dynamics, which estimate the nutritional status of the offspring and their effects on survival.
... The lipidomic approach employed in the present study allowed an unprecedented insight on the lipid dynamics during embryonic development in decapod crustaceans. TLC analysis revealed the biochemical The important role that PC plays during embryogenesis was reflected in our results, with their decrease in relative abundance from stage 1 to 3 embryos agreeing with data from other marine species 14,15 . PC is recognized as the major component of biological membranes 16 , therefore of paramount importance on cells biochemistry and physiology 17,18 . ...
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Embryogenesis is an important stage of marine invertebrates with bi-phasic life cycles, as it conditions their larval and adult life. Throughout embryogenesis, phospholipids (PL) play a key role as an energy source, as well as constituents of biological membranes. However, the dynamics of PL during embryogenesis in marine invertebrates is still poorly studied. The present work used a lipidomic approach to determine how polar lipid profiles shift during embryogenesis in two sympatric estuarine crabs, Carcinus maenas and Necora puber. The combination of thin layer chromatography, liquid chromatography-mass spectrometry and gas chromatography-mass spectrometry allowed us to achieve an unprecedented resolution on PL classes and molecular species present on newly extruded embryos (stage 1) and those near hatching (stage 3). Embryogenesis proved to be a dynamic process, with four PL classes being recorded in stage 1 embryos (68 molecular species in total) and seven PL classes at stage 3 embryos (98 molecular species in total). The low interspecific difference recorded in the lipidomic profiles of stage 1 embryos appears to indicate the existence of similar maternal investment. The same pattern was recorded for stage 3 embryos revealing a similar catabolism of embryonic resources during incubation for both crab species.
... Similarly, the relatively low levels of SE−WE in all species and seasons is likely related to the fact that relatively few Calanus spp. (which are known to be high in SE−WE; Kattner et al. 1994) are found in the GoL (Costalago et al. 2011). In contrast, the higher indication of Oithona and Oncaea spp. in the diets of forage fishes is not surprising as they re present the dominant copepods in the GoL throughout the year (Costalago et al. 2011, Espinasse et al. 2013. ...
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We describe the total lipid content, lipid class composition and fatty acid profiles of adult forage fishes (anchovy, sardine and sprat) sampled in the NW Mediterranean Sea in 2010 and 2011. Inter- and intra-species differences were mostly related to sampling period with limited effect of gender or total length. As an assemblage, total lipid content and relative levels of triacylglycerols and fatty acids 16:1n7, 20:5n3 and 14:0 in forage fish were highest in summer and autumn, indicating better feeding conditions and a more pronounced diatom-supported food web. In contrast, total lipid content was lowest at the end of winter and spring, and coincided with high levels of 22:6n3, indicating a more herbivorous diet based on dinoflagellates. Resource partitioning and niche separation, as inferred from fatty acid profiles, were apparent between species. Sardine showed a more diverse, temporally separated feeding strategy than anchovy, and dietary overlap was higher in winter than summer with sardine having higher markers of copepods, 22:1n11 and 20:1n9. Sprat collected in winter occupied a separate niche area to both sardine and anchovy with higher total lipid content and carnivory biomarker 18:1n9. Our results show that the lipid dynamics of forage fishes can be used to gain quantitative insights into sub-system level changes in species interactions, including prey and predator productivity.
... Our study provided further insights on this topic by revealing that larval biochemical profiles can differ significantly even between consecutive years. Interannual variability in lipid and TAG larval content has already been documented to occur over consecutive years in other commercially important decapods (Kattner et al. 1993;Wickins et al. 1995;Palacios et al. 1998Palacios et al. , 1999Ouellet & Allard 2002;Pedersen & Storm 2002;Ouellet & Plante 2004). The diet of M. brachydactyla in the wild is known to include about 100 different items (e.g. ...
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The spider crab Maja brachydactyla is an important commercial species in Europe and supports intensive fisheries in the NE Atlantic. A field survey was performed to assess long-term and consecutive interannual (2005–2010) variation of the biochemical composition of newly hatched larvae of M. brachydactyla. Larval biochemical profiles differed significantly among years, with pronounced differences being recorded in 2010. Differences among batches of newly hatched larvae were mainly explained by the contribution of triacylglycerols and, to a lesser degree, by protein and lipid content. The use of different nutrition indices is discussed. The biochemical composition of newly hatched larvae from M. brachydactyla was highly variable, even though surveyed broodstock was obtained from the same local population and was always sampled during the same season. The unpredictable biochemical profiles of newly hatched larvae may condition their survival and recruitment. This largely overlooked aspect of larval variability should be considered in future fisheries management strategies and captive production of marine organisms that still rely on wild seeds.
... Such plasti - city is known at a range of scales, e.g. inter-annually, between seasons, and within a single breeding sea- son (Boddeke 1982, Sheader 1983, 1996, Kattner et al. 1994, Oh & Hartnoll 2004). POI plasticity is influenced by maternal size and age, food quantity and quality, population density, and temperature. ...
Article
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At the inter-specific level, per offspring investment (POI) is associated with larval development mode and follows a macro-ecological trend within the marine environment; higher POI and its associated greater degree of endotrophy and abbreviated development is found in cooler, high latitude regions. Here, the implications of between-brood-variation in hatchling energy content (measured as carbon mass) on larval starvation resistance and developmental plasticity within the caridean shrimp Palaemonetes varians were assessed. Results demonstrate that greater POI provides increased endotrophy and the potential for abbreviated development. In the absence of food, P. varians larvae from broods of higher hatchling energy content developed to more advanced larval stages and survived for longer before succumbing to starvation. In the presence of food, P. varians larvae from broods of higher hatchling energy content developed through fewer larval instars, showed higher growth rates, and had shorter development times. Also, for larvae developing through the same number of larval instars, larvae from broods of higher hatchling energy content developed to greater juvenile dry weight. These data support the hypothesis that macro-ecological trends in development mode are driven by inter-specific variations in POI. At the intra-specific level, phenotypic plasticity allows for environmentally mediated variations in POI. Under differential selection pressures, this flexibility may, thus, eventually permit the evolution of the diverse and complex life cycles observed in the marine environment.
... These FA are common in caridean shrimps with abbreviated larval development (Thatje et al. 2004; Calado et al. 2010). The high and largely stable content of stearic acid is explained by its predominance in membrane phospholipids (Kattner et al. 1994; Wehrtmann & Graeve 1998 ). High initial proportions of OA, LA, and EPA, which are essential FA in crustaceans (i.e. ...
Article
Palaemonetes zariquieyi, an endemic palaemonid species of shrimp that lives in freshwater and brackish coastal habitats in eastern Spain, shows an abbreviated, non-feeding larval development comprising only three zoeal stages. To identify the endogenous bioenergetic fuel that allows for food-independent development from hatching to metamorphosis, larvae were reared under controlled laboratory conditions, and ontogenetic changes in dry weight (W), elemental (CHN) and lipid composition (total lipids, principal lipid classes, fatty acids [FA]) were quantified at the onset of each zoeal stage and in the first juvenile. Values of W, C and H per larva and per mass unit of W decreased throughout the time of larval development, while the N content showed only a weak decline (suggesting strong lipid but only little protein degradation). Correspondingly, directly measured values of total lipids (both in µg/larva and in % of W) decreased gradually, with neutral lipids consistently remaining the predominant and most strongly used fraction; sterol esters and waxes were not detected. In contrast to the neutral lipids, the fraction of polar lipids per larva remained stable and, as a consequence, tended to increase as a percentage of total lipids. Likewise, other important lipid fractions such as free fatty acids and cholesterol remained stable throughout the time of larval development. Among the FA, palmitic (16:0), oleic (18:1n-9), linoleic (18:2n-6) and eicosapentaenoic (20:5n-3) acid were predominant, showing a significant decrease during larval development; stearic (18:0), vaccenic (18:1n-7) and arachidonic acid (20:4n-6) were found only in small amounts. Our results indicate that the lecithotrophic development of P. zariquieyi is primarily fuelled by the utilization of lipids (especially triacylglycerides and other neutral lipids), which is reflected by a decreasing carbon content. Proteins and polar lipids, by contrast, are preserved as structurally indispensable components (nerve and muscle tissues, cell membranes). The abbreviated and non-feeding mode of larval development of P. zariquieyi may have an adaptive value in land-locked freshwater habitats, where planktonic food limitation is likely to occur. The patterns of reserve utilization are similar to those previously observed in other palaemonid shrimps and various other groups of decapod crustaceans with lecithotrophic larvae. This suggests a multiple convergent evolution of bioenergetic traits allowing for reproduction in food-limited aquatic environments.
... These FA are common in caridean shrimps with abbreviated larval development (Thatje et al. 2004;Calado et al. 2010). The high and largely stable content of stearic acid is explained by its predominance in membrane phospholipids (Kattner et al. 1994;Wehrtmann & Graeve 1998). High initial proportions of OA, LA, and EPA, which are essential FA in crustaceans (i.e. ...
Article
Full-text available
Palaemonetes zariquieyi, an endemic palaemonid species of shrimp that lives in freshwater and brackish coastal habitats in eastern Spain, shows an abbreviated, non-feeding larval development comprising only three zoeal stages. To identify the endogenous bioenergetic fuel that allows for food-independent development from hatching to metamorphosis, larvae were reared under controlled laboratory conditions, and ontogenetic changes in dry weight (W), elemental (CHN) and lipid composition (total lipids, principal lipid classes, fatty acids [FA]) were quantified at the onset of each zoeal stage and in the first juvenile. Values of W, C and H per larva and per mass unit of W decreased throughout the time of larval development, while the N content showed only a weak decline (suggesting strong lipid but only little protein degradation). Correspondingly, directly measured values of total lipids (both in µg/larva and in % of W) decreased gradually, with neutral lipids consistently remaining the predominant and most strongly used fraction; sterol esters and waxes were not detected. In contrast to the neutral lipids, the fraction of polar lipids per larva remained stable and, as a consequence, tended to increase as a percentage of total lipids. Likewise, other important lipid fractions such as free fatty acids and cholesterol remained stable throughout the time of larval development. Among the FA, palmitic (16:0), oleic (18:1n-9), linoleic (18:2n-6) and eicosapentaenoic (20:5n-3) acid were predominant, showing a significant decrease during larval development; stearic (18:0), vaccenic (18:1n-7) and arachidonic acid (20:4n-6) were found only in small amounts. Our results indicate that the lecithotrophic development of P. zariquieyi is primarily fuelled by the utilization of lipids (especially triacylglycerides and other neutral lipids), which is reflected by a decreasing carbon content. Proteins and polar lipids, by contrast, are preserved as structurally indispensable components (nerve and muscle tissues, cell membranes). The abbreviated and non-feeding mode of larval development of P. zariquieyi may have an adaptive value in land-locked freshwater habitats, where planktonic food limitation is likely to occur. The patterns of reserve utilization are similar to those previously observed in other palaemonid shrimps and various other groups of decapod crustaceans with lecithotrophic larvae. This suggests a multiple convergent evolution of bioenergetic traits allowing for reproduction in food-limited aquatic environments
... Lipids are essential nutrients (Kattner et al., 1994); they provide essential fatty acids, cholesterol and a large amount of energy. Lipids are the primary energy reserve accumulated during larval development of crustaceans (Heras et al., 2000) and are metabolised by the non-feeding puerulus stage in spiny lobsters . ...
Article
Stocking density is an important factor affecting the competency of later stage lecithotrophic spiny lobster larvae, yet its influence on biochemical composition has rarely been considered. Biochemical analysis of phyllosoma during ontogeny provides information on the energy storage requirements of late instar phyllosoma and their ability to survive metamorphosis and achieve the energetic demands of the puerulus stage. The current study is the first to examine biochemical composition of spiny lobsters through the entire larval phase. Survival, growth, development, and biochemical composition were measured in Sagmariasus verreauxi phyllosoma that were cultured at High Density (HD) and Low Density (LD) from hatch to puerulus. Protein measured directly by Lowry was considerably lower than crude protein as calculated from nitrogen (N) content using N × 6.25, suggesting that a conversion factor of 6.25 was too high. Survival of phyllosoma was significantly higher in the HD treatment after instar 9 due to high mortalities of LD phyllosoma caused by high ozonation during instar 9. However, HD phyllosoma were less susceptible to the high ozonation event possibly due to the larger biomass in HD tanks. Phyllosoma growth and development were more advanced in LD phyllosoma after 108 d. Instar 17 LD phyllosoma were also significantly larger than instar 17 HD phyllosoma. The C:N ratio confirmed proportionally more lipid than protein was accumulated during larval development before a significant decrease in lipid reserves between instar 17 and the puerulus stage by over 21% to fuel the process of metamorphosis and the non-feeding puerulus stage. The study demonstrated the larval phase of S. verreauxi is important for accumulating lipid reserves to fuel metamorphosis and the puerulus stage and provides a more complete picture of the culture requirements of spiny lobsters during ontogeny, particularly for the rarely studied late phyllosoma instars.
... Within the Crustacea, seasonal (Boddeke 1982, Amsler & George 1984, Sheader 1996, Paschke 1998, inter-annual (Kattner et al. 1994) and latitudinal (Crisp 1959, Clarke et al. 1991, Wehrtmann & Kattner 1998, Lardies & Castilla 2001, Lardies & Wehrtmann 2001, Brante et al. 2003 intraspecific differences in egg traits have been reported. Furthermore, in Crustacea with consecutive ovipositions, egg quality may vary with spawning order, a feature of considerable importance in shrimp aquaculture (e.g. ...
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Previous study on Cancer setosus Molina, 1782 showed that latitudinal changes in temperature control the number of annual egg masses. This study focused on the effects of pre-oviposition temperature and female size on egg traits in C. setosus from northern (Antofagasta, 23° S) and central-southern (Puerto Montt, 41°S) Chile. Blastula eggs produced in nature ranged in dry mass (DM) from 9.1 to 15.1 μg, in carbon (C) from 4.8 to 8.4 μg, in nitrogen (N) from 1.0 to 1.6 μg, in C:N ratio between 4.7 and 5.4, and in volume (V) between 152 and 276 mm3 × 10-4 per female. Blastula eggs from females caught early in the reproductive season in Puerto Montt (September 2006) were significantly higher in DM, C, N, and V than those of females caught 2 mo later, reflecting a seasonal increase in water temperature. In Puerto Montt 'early' and 'late' season blastula eggs were higher in DM, C, N, and V than eggs from Antofagasta by about 32 and 20%, respectively. Subsequent egg masses produced in captivity in Puerto Montt followed this pattern of smaller eggs with lower DM, C, and N content at higher pre-oviposition temperatures. In Antofagasta no significant difference in DM, C, N, and V between eggs produced in nature and subsequent eggs produced in captivity was found and all egg traits were significantly positively affected by maternal size. Reproductive plasticity in C. setosus helps to explain the species wide latitudinal distribution range.
... The dietary requirements of the first larval stage may vary inter-and intraspecifically as well as seasonally or interannually, depending on energy reserves and essential nutrients stored in the egg yolk, and on larval abilities to synthesize macromolecular nutrients from precursors (Staton & Sulkin 1991, Kattner et al. 1994, Paschke 1998. As a consequence of a rapid depletion of yolk reserves, the larvae of most marine invertebrate and fish species must start feeding within a limited time after hatching to avoid starvation effects. ...
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The North Sea shrimp Crangon crangon (Linnaeus, 1758) has seasonal dimorphism in egg size, with larger winter eggs and smaller summer eggs. In the laboratory, we compared the tolerance of nutritional stress in Zoea I larvae hatching from the different types of eggs (referred to as 'winter larvae', WL, and 'summer larvae', SL, respectively). Starvation tolerance was quantified as median point-of-no-return (PNR50) and point-of-reserve-saturation (PRS50)-PNR50 is defined as the time when 50% of starved larvae have lost the capability to recover (after subsequent feeding); PRS50 is the time when 50% of fed larvae attain the capability to develop through the rest of the moulting cycle using internally stored energy reserves. These critical points in the moulting cycle were estimated by fitting sigmoidal dose-response curves of cumulative mortality to the time of initial starvation or feeding, respectively. Significant seasonal variation was observed in the initial biomass at hatching (16.2 vs 14.7 mug dry mass in WL and SL, respectively) as well as in the development duration of continuously fed larvae (fed controls, FC; average Zoea I stage durations: 4.4 vs 5.0 d). Likewise, WL showed a consistently shorter development duration after 1 to 4 d initial starvation and subsequent feeding (PNR treatments). In treatments with 3 to 5 d initial starvation, mortality was also significantly lower in WL than in SL. Both larval groups showed an increasingly delayed moult to the Zoea 11 stage with increasing time of initial starvation, but this effect was significantly weaker in WL than in SL. As a consequence, the mean PNR50 value was higher in WL than in SL (4.8 vs 3.5). When zoeae were continuously starved from hatching onwards (starved controls, SC), WL were able to survive significantly longer than SL (8.8 vs 6.4 d). In experiments with differential periods of initial feeding and subsequent starvation (PRS experiments), 50% of the WL exceeded their PRS after only 1 d of food availability, while SL required at least 2 d of feeding to become independent of further food supply. PSR50 values of WL and SL differed significantly (1.0 vs 1.6 d). Our results indicate a shorter development and stronger starvation resistance in WL compared to SL. Seasonal variation in egg size and initial biomass and physiological condition of early larvae allow for an extended period of reproduction, including larval hatching under conditions of low or unpredictable planktonic food availability in winter and early spring.
... Seasonal variations in reproductive traits have also been detected in the elemental composition and energy content of eggs and first-stage larvae of various other decapod crustaceans such as shrimp (Crangon crangon; Boddeke 1982; Paschke et al. 2004;Urzúa et al. 2012;Crangon spp.;Kattner et al. 1994) and crab (Neohelice granulata; Bas et al. 2007). Heavier embryos and larvae with higher energy and C:N values were also in these species observed at the beginning of the seasons of egg laying and hatching, indicating an enhanced lipid content. ...
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In the Seno de Reloncaví, southern Chile, seasonal changes in dry weight (DW) and elemental composition (CHN) were studied in embryo (initial embryonic stage), newly hatched zoeae, and newly settled megalopae of a porcelain crab, Petrolisthes laevigatus. Samples were taken throughout the seasons of egg laying (March-December), hatching (August-February), and settlement (October-February). Values of DW and CHN per embryo or larva, respectively, were consistently minimum in the middle of each season and maximum near its beginning and end. Patterns of seasonal variation in early embryonic biomass may thus be carried over to larvae at hatching and, possibly, to the settlement stage. Such carry-over effects may be selectively advantageous, as zoeae released at the beginning or near the end of the hatching season face conditions of poor planktonic food availability in combination with low winter temperatures or decreasing temperatures at the end of summer (enforcing long development duration). Hence, an enhanced female energy allocation into egg production may subsequently translate to enhanced yolk reserves remaining at hatching, allowing for a larval development under unfavourable winter conditions. In summer, by contrast, plankton productivity and temperatures are generally high, allowing for fast larval growth and development. This coincides with minimal biomass and energy contents both at hatching and settlement. In conclusion, our data suggest that seasonal patterns in the biomass of early developmental stages of P. laevigatus may reflect phenotypic variability as an adaptive response to predictable variations in environmental conditions, allowing this species to reproduce in temperate regions with marked seasonality in water temperature and plankton productivity.
... Seasonal effects have been identified as an important source of intraspecific variation in the reproduction of temperate marine invertebrates, in general (Ghiselin 1987). Among the Crustacea, intraspecific variability in reproductive traits such as egg size has been observed not only as a presumable response to seasonal variations in environmental conditions (Boddeke 1982;Bas et al. 2007), but also between years (Kattner et al. 1994;Ouellet and Plante 2004), and in latitudinal as well as bathymetric gradients (e.g. Wehrtmann and Kattner 1998;Lardies and Castilla 2001;Thatje et al. 2004;Brante et al. 2004;Laptikhovsky 2006;Fischer and Thatje 2008). ...
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In the shrimp Crangon crangon, an important fishery resource and key species in the southern North Sea, we studied temporal variations in size, biomass (dry weight, W) and chemical composition (C, N, protein and lipid) of eggs in an initial embryonic stage. Data from 2 years, 1996 and 2009, consistently revealed that egg size and biomass varied seasonally, with maxima at the beginning of the reproductive season (January), decreasing values throughout spring, minima in June–July, and a slight increase thereafter. This cyclic pattern explains why “Winter eggs” are on average larger and heavier than “summer eggs”. Using a modelling approach, we estimated the duration of oogenesis in relation to seasonally changing seawater temperatures. According to an additive model of multiple explanatory variables, the C content per newly laid egg showed in both years a highly significant negative relationship with day length (r² = 0.38 and 0.40, respectively; P < 0.0001), a weak positive relationship with temperature (r² = 0.08 and 0.09; P < 0.05), and a weak negative relationship with phytoplankton biomass (r² = 0.11 and 0.12; P < 0.05) at the estimated time of beginning oogenesis. Phenotypic plasticity in initial egg size and biomass is interpreted as an adaptive reproductive trait that has evolved in regions with strong seasonality in plankton production and periods of larval food limitation. In contrast to biomass per egg, the percentage chemical composition remained similar throughout the reproductive period. Both the absolute and percentage values also showed significant interannual variations, which caution against generalizations based on short-term studies of reproductive traits of C. crangon and other species of shrimp.
... The dietary requirements of the first larval stage may vary inter-and intraspecifically as well as seasonally or interannually, depending on energy reserves and essential nutrients stored in the egg yolk, and on larval abilities to synthesize macromolecular nutrients from precursors (Staton & Sulkin 1991, Kattner et al. 1994, Paschke 1998. As a consequence of a rapid depletion of yolk reserves, the larvae of most marine invertebrate and fish species must start feeding within a limited time after hatching to avoid starvation effects. ...
... From other comparative studies of decapod crustaceans we know that reproductive traits may vary significantly with seasonal changes in temperature andor daylength. For instance, there are larger winter and smaller summer eggs in the North Sea shrimp, Crangon crangon (Boddeke, 1982); in the same species, interannual differences have also been documented (Kattner et al., 1994). In some euryhaline species, there are physiological and genetic differences between partially isolated freshwater and estuarine populations. ...
Article
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In the evolution of decapod crustaceans, interspecific variation in egg size is considered as an important life-history trait that is linked with the duration of embryonic and larval development, the number and type of larval stages, and with juvenile size. Aiming to provide a quantitative characterization of reproductive traits in related decapod taxa differing in lifestyle (freshwater, estuarine, marine) and geographic-climatic distribution (tropical-temperate), we compared size, biomass, and elemental composition of eggs of caridean shrimps from three families: seven species of Palaemonidae (three congeners of Macrobrachium: M. olfersii, M. carcinus, M. acanthurus; four species of Palaemon: P. northropi, P. pandaliformis, P. elegans, P. adspersus), two Atyidae (Potimirim potimirim, Atya scabra), and one Pandalid (Pandalus montagui). Egg size was measured as larger and smaller diameter (D1 D2), volume was calculated from D1 and D2, and biomass was measured as dry mass (W), carbon (C), nitrogen (N), hydrogen (H), and energy (E, estimated from C) contents. The smallest size and lowest biomass were found in the eggs of two freshwater atyids (both originating from Brazil); the largest size occurred in a marine species, P. montagui (from the North Sea); and intermediate values in freshwater, estuarine, and marine palaemonid species (from Brazil and the Baltic Sea, respectively). Among the Palaemon species, the most limnic (P. pandaliformis) showed a significantly larger egg size and volume (P
... The importance of egg lipid droplets is likely related to the amount of energy required by the early developmental stages. Eggs of the shrimp species Penaeus setiferus, Crangon crangon and C. allemanni lack or have only trace amounts of storage lipids (Kattner et al. 1994b, Lee & Broudy unpubl. data). ...
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Zooplankton storage lipids play an important role during reproduction, food scarcity, ontogeny and diapause, as shown by studies in various oceanic regions. While triacylglycerols, the primary storage lipid of terrestrial animals, are found in almost all zooplankton species, wax esters are the dominant storage lipid in many deep-living and polar zooplankton taxa. Phospholipids and diacylglycerol ethers are the unique storage lipids used by polar euphausiids and pteropods, respectively. In zooplankton with large stores of wax esters, triacylglycerols are more rapidly turned over and used for short-term energy needs, while wax esters serve as long-term energy deposits. Zooplankton groups found in polar, westerlies, upwelling and coastal biomes are characterized by accumulation of large lipid stores. In contrast, zooplankton from the trades/tropical biomes is mainly composed of omnivorous species with only small lipid reserves. Diapausing copepods, which enter deep water after feeding on phytoplankton during spring/summer blooms or at the end of upwelling periods, are characterized by large oil sacs filled with wax esters. The thermal expansion and compressibility of wax esters may allow diapausing copepods and other deep-water zooplankton to be neutrally buoyant in cold deep waters, and they can thus avoid spending energy to remain at these depths. Lipid droplets are often noted in zooplankton ovaries, and a portion of these droplets can be transferred to developing oocytes. In addition to lipid droplets, zooplankton eggs have yolks with lipovitellin, a lipoprotein with approximately equal amounts of protein and lipid. The lipovitellin lipid is predominantly phosphatidylcholine, so during reproduction females must convert a portion of their storage lipid into this phospholipid. Developing embryos use their lipovitellin and lipid droplets for energy and materials until feeding begins. The various functions storage lipids serve during the different life history stages of zooplankton are very complex and still not fully understood and hence offer a multitude of fascinating research perspectives.
... Stevens et al. 2004b, Lischka & Hagen 2007). In contrast, interannual variability in dietary quality remains poorly documented, but has been suggested to play a role in controlling population dynamics of marine organisms on long time scales (Kattner et al. 1994, Litzow et al. 2006 ). Here, we use fatty acid trophic markers to characterize interannual variability in the diet of an important calanoid copepod from a productive and highly variable coastal ecosystem over a period of 6 yr. ...
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Although food quality is thought to play an important role in the survival of marine organisms, the extent of natural variability in food quality over long time scales remains poorly char- acterized. We present a 6 yr time series of fatty acid data from the calanoid copepod Neocalanus plumchrus, an important contributor to mesozooplankton biomass in the Strait of Georgia and the northeast Pacific Ocean. Fatty acid profiles indicate significant spatiotemporal differences in the diet of this copepod. Spatially, oceanic specimens display fatty acid signatures characteristic of omnivo- rous copepods while coastal animals display primarily herbivorous, diatom-based signatures. Tempo- rally, the fatty acid profiles of coastal N. plumchrus shifted from an omnivorous oceanic diet to an her- bivorous, diatom-based diet between 2001 and 2006. The ratio of diatom to flagellate fatty acid markers increased over time, peaking from 2005 to 2006. The composition of flagellate markers also changed from primarily dinoflagellate markers (rich in docosohexaeonic acid) to green algal markers (poor in this essential fatty acid). The diet of N. plumchrus as deduced from fatty acids correlates with phytoplankton community composition. The abundance of coastal N. plumchrus in the Strait of Geor- gia was strongly correlated with the ratio of docosahexaeonic acid to eicosapentaeonic acid in the lipids of these copepods. We also discuss the potential for an imbalance of essential fatty acids sup- plied by a diatom-rich diet to limit the growth and survival of N. plumchrus in the Strait of Georgia.
... Egg-bearing females of N. norvegicus were collected from February to June 2001 after being landed in Cascais (Portugal) by commercial fishing vessels. The egg mass was removed from the females and eggs were classified according to the following criteria (modified from Kattner et al., 1994): stage I, uniform yolk and no embryonic development visible; stage II, eyes clearly visible with 1/2 yolk consumed; stage III, almost no yolk present and embryo fully developed. Thirty eggs were separated from each female (9 females, 3 per each embryonic stage) and length and width were measured under a stereo microscope (OlympusR, model SZ6045TR) with a calibrated micrometer eyepiece. ...
Article
The objectives of the present study were to investigate the total and free amino acid profiles and lipid dynamics (lipid classes and fatty acids) during embryogenesis of Nephrops norvegicus, in order to understand the early larval protein and lipid requirements. There was a significant increase in total essential (EAA) and nonessential amino acid (NEAA) contents during embryonic development (P<0.05). The major EAA were arginine, histidine and leucine, while the most important NEAA were glutamic acid, aspartic acid, glycine and proline. The higher percent increase occurred in respect to NEAA (19.9%), mainly due to the significant increase of glycine (51.8%) and alanine (35.1%). The free amino acid (FAA) content, especially the free nonessential amino acids (FNEAA), also increased significantly, being the quantitatively most important glycine, proline, taurine and glutamic acid. The free essential amino acids (FEAA) also revealed significant variations and the most important were arginine, lysine and leucine. A higher percent increase was attained with FNEAA (66.8%) in comparison to FEAA (49.6%), mainly due to the significant increase of homocystine (87.7%), valine (83.5%), glutamine (82.5%) and glutamic acid (76.1%). The absorption of dissolved organic compounds from seawater can explain the increase in free and total amino acid (FAA and TAA) contents, because a significant increase in the water content during development was observed (P<0.05). A substantial decrease in all neutral lipid classes (P<0.05) was observed during embryonic development, namely diacylglycerols (DAG) (97.9% of utilization), triacylglycerols (TAG) (93.2%), sterol esters (StE) (91.3%) and monoacylglycerols (MAG) (90.4%). The quantitatively most important fatty acids were the saturates (SFA) 14:0, 16:0 and 18:0, the monounsaturates (MUFA) 16:1n-7, 18:1n-9 and 18:1n-7, and the polyunsaturates (PUFA) 18:2n-6, 18:3n-3, 20:4n-6, 20:5n-3 and 22:6n-3. The unsaturated fatty acids (UFA) are used up at a higher rate (54.5% of utilization) than SFA (42.8%); within the UFA, MUFA are more consumed than PUFA (59.3% and 52.7%, respectively). In terms of the utilization of individual fatty acids, there was a preferential consumption of 20:4n-3, 22:6n-3, 18:1n-9, 22:5n-3 and 16:1n-7. It is evident that N. norvegicus depends primarily on lipid reserves of the egg during early ontogeny. On the contrary, this species tends to conserve EAA and increase NEAA and FNEAA contents during embryonic development.
... PUFA have been largely demonstrated to be essential for marine animals as they generally can not be synthesised de novo (Kanazawa et al. 1979;Albentosa et al. 1996). There is a wide variation in the essential fatty acids (EFA) requirement of aquatic animals (Takeuchi 1997) to supply fatty acids necessary for reproduction, physiology, and other biological activities (Pillsbury 1985;Frolov & Pankov 1992;Kattner et al. 1994). Some of these fatty acids have also been shown to be of special significance for gonad maturation and broodstock quality (Soudant et al. 1996). ...
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Mature specimens of Argopecten purpuratus (Lamarck, 1819), obtained from the Culture Center of Tongoy Bay, northern Chile were conditioned at 20°C and fed three different diets (mixture of pure microalgae, microalgae supplemented with a lipid emulsion, and microalgae supplemented with carbohydrates). Fatty acids were analysed at 0, 18, 38, 58, and 72 days of gonad conditioning. Both muscle and gonads showed a significant decrease in the relative distribution (weight % of total fatty acids) of saturated, monounsaturated, and n‐6 polyunsaturated fatty acids (mg/g dry weight). Generally, an increase of the n‐3 polyunsaturated group was observed for gonads until Day 58, thereafter a decrease was observed, probably as a result of environmental stress. A similar trend was observed for eicosapentaenoic and docosahexaenoic acids. n‐3 polyunsaturated fatty acids were the most preserved fatty acids, suggesting their importance as essential fatty acids for gonad maturation. The lipid‐supplemented diet afforded the best gonadal maturation and spawning occurred after 72 days.
... The number of eggs produced increases with female size (available cephalothroracic space) and decreases with increasing average egg volume (Hines, 1982Hines, , 1986). Intraspecifically, decapod initial egg size varies latitudinally and within geographically isolated populations (Wehrtmann and Kattner, 1998; Kyomo, 2000; Rosa et al., 2007), between different reproductive seasons (Amsler and George, 1984; Pond et al., 1996; Calado et al., 2005), and on a year-to-year basis in the same locality (Kattner et al., 1994). In brachyurans, egg size is not generally correlated with adult size (Hines, 1986; Oh and Hartnoll, 1999). ...
Article
We examined potential fertility, egg volume, and water, lipid, and fatty acid content through embryogenesis in a population of female U. rapax from Sebastian Inlet, Florida. Carapace width (CW) ranged from 10.80 to 20.09 mm (N = 184), and each female carried 5000 to 30,000 eggs in the last stage of development. Female CW was found to be a good predictor of the number of eggs in the later stage of development (potential fertility = 7.908 CW2.7655, R2 = 0.749). Egg volume increases (from 0.0079 to 0.0134 mm3) was mildly correlated (r = 0.79) with an increase in egg water content (from 60 to 69%). Egg lipid and fatty acid content decreased through embryogenesis, due to its importance as energy source. The most consumed fatty acids were the monounsaturated (97.81 μg . mg dw−1) followed by the saturated (64.34 μg · mg dw−1) and polyunsaturated (38.69 μg · mg dw−1). Fatty acids 16:0, 18:2n-6, 16:1n-7, and 18:2n-6 are consumed preferentially (39.91, 38.45, 29.4 and 23.93 μg · mg dw−1, respectively), while essential polyunsaturated fatty acids, particularly docosahexaenoic acid (22:6n-3) and eicosapentaenoic acid (20:5n-3), are conserved. Egg fatty acid profile also reflects diet and habitat of adults. A medium-low EPA/DHA ratio suggests U. rapax occupies a medium trophic level. The low ratio (18:1n-7/18:1n-9) and high percentages of 18:1n-9 fatty acid (18%) and essential C18 and C20 polyunsaturated fatty acids (19-23%) suggests adults are omnivores consuming primary producers (like algae and mangrove leaves) and small invertebrates. The high percentage of odd-numbered fatty acids (above 3-3.5%) also suggests scavenger/detritivore behaviour.
... The number of eggs produced increases with female size (available cephalothroracic space) and decreases with increasing average egg volume (Hines, 1982Hines, , 1986). Intraspecifically, decapod initial egg size varies latitudinally and within geographically isolated populations (Wehrtmann and Kattner, 1998; Kyomo, 2000; Rosa et al., 2007), between different reproductive seasons (Amsler and George, 1984; Pond et al., 1996; Calado et al., 2005), and on a year-to-year basis in the same locality (Kattner et al., 1994). In brachyurans, egg size is not generally correlated with adult size (Hines, 1986; Oh and Hartnoll, 1999). ...
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Fatty-acid and energy content of Maja brachydactyla eggs at different developmental stages (recently spawned, half-developed and ready to hatch) were analysed in order to understand what is being consumed and produced during the embryonic development. Egg volume increased during development (34%, 0.187 to 0.285 mm3, N = 270) and was negatively correlated with egg energy and fatty-acid content (r = −0.80 and r = −0.46, respectively), which decreased through embryogenesis. The most consumed fatty acids were the PUFA (21.2 μg · mg dw−1), followed by the SFA (18.8 μg · mg dw−1) and MUFA (14.9 μg · mg dw−1). Palmitic (16:0), oleic (18:1n-9) and eicosapentaenoic (EPA, 20:5n-3) acids were preferentially consumed (13.14, 9.21 and 8.67 μg · mg dw−1, respectively). The fatty acid composition found in M. brachydactyla eggs reflected the habitat and omnivorous and detritivorous scanvenger diet of the adults, although the consumption of algae was more important than previously thought, at least in the area where these adults were captured. Pre-hatching eggs have a high PUFA content (64.5 μg · mg dw−1 or 46.3% of the egg fatty-acid content). We conclude that larvae of this species might need a diet rich in PUFA, particularly EPA and DHA, for successful development. From the culture perspective, live preys commonly used in aquaculture will likely require to be enriched with DHA.
... Newly hatched decapod crustacean larvae are known to display dietary requirements varying intra and interspecifically, seasonally and annually (Anger, 2001). Such dietary requirements are dependent on the qualitative and quantitative aspects of energetic reserves stored in the egg yolk (Kattner et al., 1994). In general, since yolk reserves are rapidly catabolized, decapod larvae must start feeding immediately after hatching and avoid the deleterious effects induced by starvation. ...
Article
The genus Lysmata is present worldwide in a variety of habitats and newly hatched larvae from different species may display variable tolerance to starvation. This work evaluates the effect of starvation on survival of newly hatched and zoea II larvae of Lysmata amboinensis, Lysmata boggessi, L. debelius and L. seticaudata. Survival of newly hatched larvae was influenced by the period of starvation and by parental species. L. boggessi and L. seticaudata newly hatched larvae displayed facultative primary lecithotrophy (FPL), with all continuously starved larvae molting to zoea II. In general, when starved for similar periods, L. debelius larvae displayed significantly higher survival to zoea II than L. amboinensis. However, no L. debelius or L. amboinensis larvae endured starvation for more than 24 h. Survival from zoea II to zoea III was affected by the duration of starvation period, by parental species and by starvation of the first zoeal stage (for L. boggessi and L. seticaudata). Zoea II from all tested species, produced from starved or fed zoea I, displayed similar survival when larval preys were always available. However, when starved for only 24 h, zoea II of L. boggessi and L. seticaudata originating from continuously starved zoea I displayed significantly lower survival than those produced from fed larvae. Only zoea II originating from fed zoea I displayed facultative secondary lecithotrophy (FSL) and larvae from all tested species were able to successfully molt to the third zoeal stage in the total absence of food. This feature has never been previously recorded among caridean shrimps. Larval preys ingested in zoea I contribute to the build up of energetic reserves that latter are catabolized during starvation in zoea II, allowing some larvae to molt to zoea III. Since differences were recorded among the survival displayed by starved zoea II of Lysmata with FPL (L. boggessi and L. seticaudata) and those without such feature (L. aniboinensis and L. debelius), it seems that egg yolk reserves are totally depleted in starved larvae and that energy accumulated through larval preys ingestion plays the crucial role. Future studies may help to confirm the current informal division of genus Lysmata: "crowd" species (L. boggessi and L. seticaudata) displaying FPL vs "pairs" species (L. amboinensis and L. debelius) without FPL. Additionally, it may also be confirmed that "pairs" species are ancestral to "crowd" species, since it is unlikely that FPL would have regressed to full planktotrophy. (C) 2007 Elsevier B.V. All rights reserved.
... Triacylglycerol, a short-term energy reserve molecule, is another common lipid reserve utilised by crustaceans (Teshima et al. 1977;Clarke et al. 1989;Virtue et al. 1993;Kattner et al. 1994;Hagen et al. 1996) and many other marine organisms (Benson and Lee 1975). TAG was found only in trace amounts in recently moulted and starved postpuerulus juveniles, with slightly greater proportions in the fed control animals. ...
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Post-settlement survival success in rock (spiny) lobsters is likely to be influenced by the nutritional condition of the non-feeding pueruli at settlement. Consequently, we predict that survival shortly after settlement depends on food resources being replenished rapidly. In the present study the length of the starvation period on survival in post-puerulus juveniles of the western rock lobster Panulirus cygnus was investigated using laboratory experiments. All experimental pueruli were collected from Seven Mile Beach, Western Australia, using sandwich puerulus collectors. Total lipid, lipid class, fatty acid, total protein and total glycogen analyses were conducted on fed and starved post-puerulus juvenile lobsters, from moult into the first juvenile stage to 30days post-moult. From a total of 165 samples used in this study, 15 post-puerulus juveniles were sampled initially and five post-puerulus juveniles sampled every 5 days thereafter for biochemical analysis. Post-puerulus juveniles survived up to 34days of starvation. The maximal starvation time for post-puerulus juveniles from which recovery was observed is estimated to be in excess of 22days. Lipids, in particular phospholipids, are shown to be the most important source of energy catabolised during the starvation period, with greater than 62% reduction in total lipid at day 30. Proteins were also catabolised during starvation, but to a much lesser extent than lipid, decreasing by 34% in total content at day 25. Carbohydrates were present always as a minor constituent, however their content did not change significantly during starvation. Our results suggest that the pueruli collected in December 2004 were generally in sufficiently good condition to survive further starvation after settlement; nutritional levels of the post-puerulus juveniles were enhanced compared to those observed in previous studies. We suggest that the high levels of energy reserves in rock lobster post-pueruli may provide an adaptive advantage to aid the transition from pelagic to benthic living by minimising stress and the consequences of failing to locate immediately suitable food resources.
... The biochemical composition of crustacean eggs has been fairly well investigated (see Pandian, 1994 for a review) and several studies report on the constituents of decapod eggs during embryogenesis (e.g. Pandian, 1970;Herring, 1973;Amsler and George, 1984;Clarke et al., 1990;Clarke, 1993;Biesiot and Perry, 1995;Lardies and Wehrtmann, 1996) although few have looked at the FA depletion during embryonic development (Clarke et al., 1990;Kattner et al., 1994;Wehrtmann and Graeve, 1998;Wehrtmann and Kattner, 1998;Narciso and Morais, 2001). Much research has been conducted to determine the essential FA requirements of crustacean species with aquaculture potential; a great part of this research has focused on the quantitative and qualitative lipid requirements of commercially important species and relatively few studies have examined other crustacean species (e.g. ...
Article
Variations in egg volume and fatty acid (FA) content through embryogenesis were evaluated in Uca species from Inhaca island, Mozambique. Egg volume increased 96.1%, 93.3%, 84.2%, 92.9%, 96.3%, respectively, in Uca annulipes, Uca inversa, Uca urvillei, Uca chlorophthalmus and Uca vocans (p < 0.05). Fatty acid content decreased through embryogenesis, showing its importance as fuel during embryonic development. Major fatty acids were 16:0, 18:0, 16:1n-7, 18:1n-9, 18:1n-7, 18:2n-6, 20:5n-3 and 20:4n-3. Unsaturated fatty acids (UFA) and saturated fatty acids (SFA) were used up at a similar rate for U. annulipes and U. inversa contrarily to the other three species. Within the UFA, MUFA were more consumed than PUFA for all species except U. chlorophthalmus. The high values detected for fatty acid trophic markers (essential C18 and C20 PUFAs) and odd-numbered fatty acid suggest that Uca species occupy medium trophic level, primarily omnivores and scavengers/detritivores consuming algae common in the intertidal habitats. The fatty acid consumption pattern during embryonic development was essentially similar between species with some variation as expected, as FA content varies within species mainly due to female feeding ecology, nutritional and physiological conditions, differential demands on resource allocation and geographic and seasonal variations in embryonic development.
... DHA and EPA were detected in relatively low proportion in eggs of M. rosenbergii. In general, C22:6n-3 has been considered as one of the important fatty acids in decapod eggs, accounting for roughly 10–20% of the total fatty acids (Kattner et al., 1994; Wehrtmann and Kattner, 1998). We found that the contents of both EPA and DHA decreased during the embryonic development, and DHA that was more depleted than EPA. ...
Article
Biochemical composition and digestive enzyme activities of eggs during embryonic development were studied in the freshwater prawn, Macrobrachium rosenbergii. Proteins, lipids and carbohydrates were the main components in the embryos of M. rosenbergii. The proteins in yolk were used mainly as the structural substance, whereas the lipids and carbohydrates were used mainly as the energy sources. Protein content generally increased while lipid and carbohydrate contents decreased during the embryonic development. Seventeen amino acids, including eight essential amino acids, were found in every stage of embryonic development. The ratio of the contents of each essential amino acid (EAA) to total essential amino acid (TAA) remained unchanged during the different stages of embryonic development. The proportional content of glutamic acid was the highest among all the amino acids, and leucine content was the highest among the EAAs. The predominant fatty acids, in terms of relative proportion, were C16:0, C18:1n-9, C18:2n-6, C18:0 and C16:1 in each embryonic development stage. The monounsaturates (MUFA) were the preferentially utilized components of the unsaturates (UFA). C18:1n-9c was mainly used as an energy source during embryonic development, whereas C18:3n-3 and ARA mainly acted as the structural substances in embryos. SFA acted as the main energy source during early stages, from fertilized egg to gastrula stage, and MUFA acted as the main energy source from egg nauplius to egg metanauplius stage. HUFA were used mainly as energy sources during late stages. Of the five digestive enzymes assayed, activities of pepsin, trypsin and amylase were relatively high. Activities of pepsin, trypsin, amylase and cellulase increased during both the early and later embryonic stages, but decreased during the middle stages. The activity of lipase decreased after the gastrula stage. The gastrula stage was a special stage of embryonic development where organ anlage came into being. Activities of pepsin, trypsin, amylase and cellulase reached the highest level during the zoea stage. Variations of biochemical compositions and digestive enzyme activities were closely related to events in morphogenesis during the embryonic development of M. rosenbergii.
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In order to find out the effects of salinity on the reproductive characteristics and embryo quality of female E. sinensis broodstock, the mating and spawning experiments of broodstock under different salinities (3, 6, 9, 12, 15, 18, and 21 ppt) were conducted, and each treatment had three replicate tanks with five males and 10 females stocked in each tank. The reproductive characteristics of broodstock, such as mating and spawning rate, egg production, reproductive index and fecundity, and the egg diameter, weight, biochemical composition and fatty acid composition of embryos were analysed. The results showed that female crabs can mate but did not spawn at salinity 3 ppt, and the lowest salinity for spawning was 6 ppt. When the salinity was 18 ppt, the egg production, reproductive index and fecundity of the broodstock reached the highest. The embryo diameter was the largest at salinity 6 ppt, which was significantly higher than that in other groups (p < 0.05). When the salinity was 12 ppt, the wet weight and dry weight of a single embryo were the highest. With the increase in salinity, the contents of crude protein, total lipid and moisture in the embryo of E. sinensis increased first and then decreased. When the salinity was 15 ppt, the contents of crude protein, total lipid and ash in embryos reached the highest. At salinity 21 ppt, the total contents of saturated fatty acids (SFA) and polyunsaturated fatty acids (PUFA) were the highest, and there were significant differences with other groups (p < 0.05). The total amount of MUFA was the highest at salinity 12 ppt. The results provided a theoretical basis and reference for artificial culture and breeding of E. sinensis.
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The present study was designed to investigate the efficiency of lipid‐enriched brood stock diets on the reproductive performance, such as fecundity, egg volume, fatty acid profile of newly extruded (stage 1) embryos and the starvation threshold of the newly hatched larvae of the marine ornamental ‘hinge‐beak’ shrimp Rhynchocinetes durbanensis Gordon. Three dietary treatments (50, 100 and 150 g/kg lipids) were formulated to understand their influence on the reproduction of R. durbanensis under captive condition and compared with the wild‐caught shrimps carrying embryos in the abdomen. The reproductive parameters varied significantly between the treatments (captive and wild). Further, the fatty acid profile revealed that essential fatty acid levels of newly extruded embryos in the wild collected shrimps were almost like that of embryos produced in 100 and 150 g/kg of lipid‐enriched diets. The results of linear discriminant analysis (LDA) suggest that the fatty acid profile of embryos has confirmed the separation of four centroids indicating comprehensive differences among the embryos of captive‐reared and wild‐caught shrimps. Hence, it is recommended that the commercial diets formulated with essential nutrients would play a major role in enhancing the reproductive performance of marine ornamental shrimps.
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The maternal environment may influence the quantity and quality of resources invested in offspring (per-offspring provisioning) and this trait, in turn, affects larval fitness and may carry-over into early juvenile life. Here, per-offspring investment was measured across three consecutive breeding seasons for the caridean shrimp, Palaemon varians. Egg and hatchling larval dry weight as well as hatchling larval biochemical composition were measured. Results indicate that egg volume is positively correlated with dry weight, but egg volume as a proxy for dry weight should be used with caution. Correlations were identified between per-offspring investment and average breeding season temperature, but the period over which temperature was averaged was important to whether these correlations were positive or negative, highlighting the complexity of identifying environmental influence on phenotypic traits. The most important factor determining offspring size and per-offspring investment was maternal size, which may be influenced by the environment as carapace length varied significantly between breeding seasons. These data demonstrate variation in egg and larval composition, related to maternal size, which will affect early life traits and survivorship for P. varians.
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Brown shrimp (Crangon crangon, L.) are subjected to a huge annual temperature range, and certain thermal conditions during winter have been identified to affect the brown shrimp population. Despite that, little is known about its thermal biology with regard to critically low temperatures. In the present study, we determined the critical thermal minima (CTmin) and the critical lethal minima (CLmin) of male and female brown shrimp of different body sizes in laboratory-based experiments. For the CTmin trials, shrimp were acclimated to 4.0, 9.0, and 14.0 °C and exposed to a cooling rate of −0.2 °C min−1. In the CLmin trials, brown shrimp were exposed to a cooling rate of −1.0 °C day−1 without prior thermal acclimation. Acclimation temperature significantly affected the temperature tolerance of brown shrimp (p < 0.001). CTmin among the experimental groups just varied slightly, and no clear effect of gender or body size was observed. In the CLmin trials, brown shrimp even tolerated the coldest temperature of −1.7 °C that could be established in the experimental setup. However, we observed a negative relationship between temperature and reactivity within the range of 7.0 and 1.0 °C that was determined by means of the flicking response. This relationship suddenly broke between 1.0 and 0.0 °C where an abrupt drop in the reactivity of the shrimp became apparent. The results of this study revealed that brown shrimp hold a wider thermal range as originally reported and that it can cope with subzero temperatures. Implications of low-temperature tolerance are discussed in the context of the brown shrimp’s ecology as well as stock assessment.
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Seasonal variation on fatty acid composition of 51 algal species from Gulf of Mannar Marine Biosphere Reserve has been studied . Analysis of fatty acids by gas chromatography revealed the presence of higher amount of saturated fatty acids than unsaturated fatty acids. Among the saturated fatty acids, the amount of palmitic acid was higher (96.88%) in all the seasons and species studied. In the present investigation, the highest relative percentage of lauric acid (31.58%), tridecanic acid (4.46%), nonadecanoic acid (2.06%), heneicosanoic acid (23.9%), oleic acid (21.38%) and linolenic acid (10.42%) were recorded during spring season from Ulva lactuca, Acanthophora spicifera, Valoniopsis pachynema, Turbinaria decurrens, Stoechospermum marginatum and Cheilosporum spectabile respectively. Highest percentage of heptadecanoic acid (12.36%) and behenic acid (8.92%) were recorded during monsoon season from Caulerpa taxifolia and Terbinaria ornate. Myristic acid (24.71%), pentadecanoic acid (3.29%), palmitic acid (96.88%), stearic acid (6.99%) and linoleic acid (28.61%) were high in Halimeda tuna, Valoniopsis pachynema, Ulva fasciata, Bryopsis plumose and Ulva reticulata respectively during post monsoon season. Padina tetrastromatica showed the highest relative percentage of arachidic acid (12.6%) during summer.
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The hippolytid shrimp Nauticaris magellanica, with a known geographical distribution covering approximately 35 degrees of latitude, was selected to conduct a latitudinal comparison regarding volume, biomass, and fatty acid changes during embryogenesis. Ovigerous females were collected from populations in northern (Guanaqueros) and central-southern Chile (Metri and Putemun). Recently produced eggs from the 3 populations sampled wen: similar in size (ranging from 0.031-0.038 min(3)). Embryos close to hatching, however, were considerably larger in central-southern Chile (Metri: 0.072 mm(3); Putemun: 0.091 mm(3)) compared with those from Guanaqueros (0.054 mml). Egg volume increase during the incubation period varied between 74% (Guanaqueros) and 160% (Putemun). Wet mass and water content of embryos increased, while dry and ash mass decreased during embryogenesis. Analyses of fatty acids revealed similar results for eggs from the 3 study sites and different developmental stages. The overall utilization of fatty acids, however, was elevated in embryos from the most southern location (Putemun) compared with that found in embryos from the other sampling sites. Main fatty acids of eggs and newly hatched larvae were the polyunsaturates 20:5 (n-3) and 22:6 (n-3), and the saturate 16:0, comprising 21, 16, and 15%, respectively, of the total. The pattern of fatty acid utilization during embryogenesis is characterized by a sharp decline of the 16:1 (n-7) fatty acid. Our results confirm a latitudinal dine in egg volume in N. magellanica. The differences observed among populations may be attributed, however, to differences in the ambient conditions (e.g., temperature, salinity, feeding) of the habitats rather than simply to its northern and southern location. In addition, the lipid biochemistry of developing eggs seems to be unaffected by latitude.
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Changes in biomass and lipid biochemistry during egg development were studied in the tropical shrimps, Alpheussaxidomus and Palaemonetesschmitti, from Pacific Costa Rica. Freshly-laid eggs of P. schmitti were substantially smaller than those of A. saxidomus; dry mass decreased during embryogenesis in the former species but remained almost constant in the latter one. Water content of eggs close to hatching were similar among both species (roughly 75%). Newly-produced eggs of the two species contained ≈20% fatty acids per egg dry mass; a comparison with data concerning decapods inhabiting tropical and temperate waters revealed that eggs produced by shrimps inhabiting tropical waters tend to have a higher lipid egg content per dry mass than those from temperate regions. Major lipid classes in the eggs of both species were phospholipids and triacylglycerols which increased and decreased during the incubation period, respectively. The predominant fatty acids of P. schmitti eggs were 16:0, 20:5(n-3) and 22:6(n-3) whereas eggs of A. saxidomus showed high amounts of 16:0, 20:5(n-3) and 16:1(n-7), and remarkably low values of 22:6(n-3) fatty acid. Lipid utilization was more pronounced in P. schmitti; in A. saxidomus, eggs close to hatching still contained 70% of the initially deposited fatty acid content which may indicate an enhanced independence of the newly-hatched larvae on external energy resources. The observed differences may partially be related to different habitat preferences, however, the role of adaptation and phylogeny as determinants of egg lipid biochemistry in caridean shrimps remains to be clarified.
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This review aims to update and extend the synopsis by Tiews (1970) on the biology and fisheries of Crangon Crangon (L.). Its wide distributional range along the European coast from the White Sea to Morocco within the Atlantic and throughout the Mediterranean and Black Seas reflects the capability of C. Crangon to cope with a wide range of temperature and salinity conditions and is further explained by its migratory capacity. Present knowledge suggests that the limiting factor at the northern cold water edge of its distribution is formed by egg and larval development and at the southern warm water edge by maintenance costs. No information is available about the genetic population structure, but patterns in isoenzymes and in morphometric characters indicate the existence of various subpopulations. Over its distributional range, especially along the north Atlantic coast, clear trends in life-history parameters are observed, most likely reflecting temperature conditions. Due to its generally high abundance, the common shrimp forms a key component in the functioning of coastal shallow ecosystems; however, it is unclear whether the population dynamics of the species is subject to top-down or bottom-up controL. On the one hand, C. Crangon is an opportunistic feeder with a wide prey spectrum though it remains to be solved whether growth conditions are optimal and only determined by prevailing water temperatures, or whether food limitation is a regulating mechanism. On the other hand, top-down control by predation cannot be excluded since C. Crangon is also an important food item for a variety of predators, especially fish species. There are strong indications that predation by C. Crangon might regulate some of their prey species. Topics for further research include (1) the analysis of the genetic population structure by means of molecular tools; (2) the study of growth and reproduction in relation to latitude; (3) the application of dynamic energy budgets for the analysis in terms of energy of the various trade-offs, including growth versus reproduction; and (4) the analysis of the mechanisms determining recruitment, especially whether top-down or bottom-up control is occurring. © R.N. Gibson, R.J.A. Atkinson, and J.D.M. Gordon, Editors Taylor & Francis.
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The present study investigates amino and fatty acid dynamics of embryos of different-sized simultaneous hermaphrodite shrimp (SH) (Lysmata seticaudata) during early (ERS) and late reproductive seasons (LRS). A significant relative decrease in total amino acids and essential amino acids (EAA) was recorded (PPn-9 and 18:1n-7 and the polyunsaturates 20:4n-6 (arachidonic acid, ARA), 20:5n-3 (eicosapentaenoic acid, EPA) and 22:6n-3 (docosahexaenoic acid, DHA). Monounsaturates were used at a higher rate, and embryos produced by SH shrimp displayed similar consumption rates of saturated and polyunsaturated fatty acids. Considering individual fatty acids, no clear utilization pattern between different-sized SH shrimp in ERS and LRS was recorded. The inexistence of consistent differences between amino and fatty acid utilization during embryogenesis among different-sized SH shrimp in ERS and LRS emphasizes the variability affecting offspring in decapod crustaceans.
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The present work is a comprehensive study of reproduction and embryonic development of Armases cinereum. Ovigerous A. cinereum (Bosc, 1802) females from Sebastian Inlet, Florida (9.88–19.4mm CW) lay 2,000–12,000 eggs per brood, depending on their CW (mm): fecundity=24.662CW1.9432. A. cinereum displayed significant brood loss through development (ca. 500 eggs per brood) independently from their CW (no senescence). However, since smaller females lay fewer eggs than larger ones, the percentage of eggs lost during embryonic development is greater in smaller females. The number of eggs carried on a later stage of development (potential fertility=5.5593CW2.4417) is a more accurate estimate of the reproductive output and subsequent recruitment. Egg volume increased during development (64%, 0.025–0.041mm3 or 0.36–0.43mm of diameter, N=270) and was strongly correlated with egg water content increase (19.21%, r=0.89). Lipids, particularly fatty acids, seem to be the major energy source for embryonic development, decreasing 56.31 and 37.08% (respectively) during embryonic development; both are negatively correlated with egg volume (r=−0.90). The utilization of fatty acids through the different developmental stages of A. cinereum is presented. The most consumed fatty acids are the monounsatured (43.33μgmg−1dw), followed by the saturated (29.91μgmg−1dw) and polyunsaturated (24.03μgmg−1). Palmitic (16:0) and linoleic (18:2n-6) acids are preferentially consumed (19.5 and 17.9μgmg−1dw, respectively). The high proportion of essential polyunsaturated fatty acids of C18 and C20 reflects the consumption of primary producers such as mangrove leaves. EPA/DHA ratio (2.85–3.84) and low DHA content indicated that this species appears in a medium-low level of the trophic chain. The low ratio of 18:1n-7/18:1n-9 and high percentage of 18:1n-9 (marker of carnivory) may be a sign of the consumption of juvenile invertebrates. The high percentage of odd-numbered FA indicated the occurrence of detritivores/scavenger behaviours. The fatty acid composition of the eggs reflects adult feeding ecology (omnivorous) and habitat.
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Larval and early post-larval growth has been investigated in H. araneus L. (Majidae) reared in the laboratory. Growth was measured as dry weight (DW), ash-free dry weight (AFDW), carbon (C), nitrogen (N), hydrogen (H), gross biochemical constituents (protein, lipid, carbohydrate, chitin, ash) and energy (calculated separately from carbon and biochemical composition). During larval development, i.e. from freshly hatched zoea-I to late megalopa, all these criteria of biomass increase by factors ranging between 5 and 14; carbohydrate shows the lowest, chitin the highest increment. There are indications of loss in organic body weight during the latest period preceding metamorphosis to the crab stage. When no food is offered during this time, megalopae lose significantly more biomass than control larvae. This suggests that food is still required, but feeding activity is reduced to a level below maintenance ingestion rate. Following metamorphosis, the juvenile crab accumulates biomass at a far higher absolute rate (expressed as µg/d) than all larval stages.
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Variations in total lipid content (g of lipid, and as % of wet and dry body weights) of the prawn Pandalus borealis, ranging from eggs to 52 mo old adults, indicate that lipid growth exhibits marked seasonal oscillations while growth in carapace length shows only comparatively weak seasonality. Positive (anabolic) lipid growth occurs from about April to September inclusive (with up to 40 % of dry body weight as lipid), the major production season for both phytoplankton and herbivorous zoo-plankton. Negative (catabolic) lipid growth (with lipid being reduced to about 10 % of dry weight) is prominent during the winter. Correspondence analysis involving the relative body-content of total lipid, total protein, ash and water indicates that both 'age' and 'seasonally' related trends are visible; lipid tends to play an increasingly important role with age but lipid levels oscillate between peaks in late summer (August/September) and troughs in late winter (February/March). Qualitative investigations of gut contents and analyses of lipid-class composition and fatty acid food-chain markers of selected sizes and ages of prawn indicate that lipid-rich zooplankton (mainly copepods and krill) form the bulk of the food of prawns from 1 to 4 yr old. Although benthic polychaetes and detrital material tend to increase in relative importance with increasing size and age of prawn, they are of secondary importance compared to pelagic organisms. The older prawns frequently contained substantial numbers of capelin scales, probably from feeding on discarded fish from prawn trawling. This study thus reveals that P. borealis in north Norwegian fjords is linked trophically to the highly seasonal pelagic production cycle. The lipid percentage of P. borealis is, however, less than that of herbivorous zooplankton but substantially greater than that of the lipid-deficient benthos.
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Zoea-1 larvae ofHyas araneus were kept under different nutritional conditions. Their midgut glands were investigated with a transmission electron microscope. The glandular epithelium consists of the cell types known from adult decapods. It is mainly the R-cell type that undergoes ultrastructural alterations which reflect nutritional conditions. R-cells of fed larvae are characterized by large lipid inclusions; after a certain period of food deprivation (point-of-no-return) the original ultrastructure cannot be reestablished. Refeeding results in large glycogen deposits in these cells.
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Juvenile shrimp Crangon crangon (L.) were sampled frequently on a tidal flat in the westernmost part of the Wadden Sea during spring and summer from 1983 to 1991. High densities (several tens per m2) of small shrimp (length range 5 to 25 mm, mean generally <15 mm) were observed at low tide during late spring, summer and early autumn. The low numbers (ca 1 m-2) of overwintered shrimps observed on the tidal flats in March were positively correlated with the mean temperature of the foregoing months. Settlement of a new generation of postlarvae (ca 5 mm) started in April after mild winters and in May after cold winters. Maximal densities were observed in June after mild winters and in July after cold winters. Settlement continued with fluctuating intensity throughout summer and early autumn. Growth was rapid: as early as ca 1 mo after settlement the shrimps reached a length of 20 to 25 mm, at which point they leave (the higher parts of) the tidal flats. Thus, juvenile shrimp use tidal flats as a transit nursery, where several successive cohorts grow up during any one summer season. Earlier published estimates of time of recruitment, mean size and growth rate of shrimp living in the same area are refuted. In spring (up to mid-June), shrimp biomass on the tidal flat was much lower after cold than after mild winters. This difference could also help explain the generally highly successful recruitment ot the bivalve Macoma balthica after cold winters.
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Larvae of the shrimpsCrangon crangon L. andC. allmanni Kinahan were reared in the laboratory from hatching through metamorphosis. Effects of rearing methods (larval density, application of streptomycin, food) and of salinity on larval development were tested only inC. crangon, influence of temperature was studied in both species. Best results were obtained when larvae were reared individually, with a mixture ofArtemia sp. and the rotiferBrachionus plicatilis as food. Streptomycin had partly negative effects and was thus not adopted for standard rearing techniques. All factors tested in this study influenced not only the rates of larval survival and moulting, but also morphogenesis. In both species, in particular inC. crangon, a high degree of variability in larval morphology and in developmental pathways was observed. Unsuitable conditions, e.g. crowding in mass culture, application of antibiotics, unsuitable food (rotifers, phytoplankton), extreme temperatures and salinities, tend to increase the number of larval instars and of morphological forms. The frequency of moulting is controlled mainly by temperature. Regression equations describing the relations between the durations of larval instars and temperature are given for bothCrangon species. The number of moults is a linear function of larval age and a power function of temperature. There is high variation in growth (measured as carapace length), moulting frequency, morphogenesis, and survival among hatches originating from different females. The interrelations between these different measures of larval development in shrimps and prawns are discussed.
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Groups of shrimp (Pandalus borealis) larvae were reared under different food concentrations and types to assess changes in condition and survival during development. As expected, rate of growth (wet weight) was higher for larvae fed on 300 Artemia nauplii∙L−1 (San Francisco strain: 0.0528 mg∙d−1) Canada strain: 0.0355 mg∙d−1) compared with those reared at 150 nauplii∙L−1 (San Francisco strain: 0.0211 mg∙d−1), larvae fed Isochrysis cells only, and no food. Shrimp larvae reared without food and larvae fed on Isochrysis cells showed a decreasing trend in wet weight during development and did not survive beyond stage II. The triacylglycerol (TAG) content of larvae accumulated rapidly during the initial phase of intermoult followed by a decline to a minimum coincident with ecdysis. High mortality (average 2.2%∙d−1) occurred during the first phases of development (first 25 d) for all rations. Differences among experiments in the magnitude of the mortality (from 0.86 to 4.66%∙d−1) at the first moult were related to the proportion of larvae in poor TAG condition. The results are consistent with the concept that the TAG condition index can be used to forecast, on a relative basis, differential survival among larval groups.
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Laboratory experiments on newly hatched larvae of the sand shrimp Crangon septemspinosa were conducted to study larval vulnerability to food deprivation. Larvae subjected to 6 different feeding regimes were well adapted to temporary lack of food, dependent on time and duration of starvation periods. Critical periods for larval development were the first 24 h after hatching, stage III and, to a lesser extent, metamorphosis. Food deprivation during the first 7 d resulted in a prolongation of stage I and more so stage II. After this period, larvae in the starvation treatments tended to reduce the duration of subsequent larval stages. Number of molts prior to metamorphosis ranged from 4 to 7 and were fewest in larvae fed every day. A majority (57.9%) of the larvae reached the juvenile stage after 5 molts. Neither the duration of larval development nor the size of the juveniles obtained from the various treatments showed significant differences. The possible importance of starvation as a recruitment regulatory process for C. septemspinosa is discussed in terms of food availability during the major hatching period in Chesapeake Bay and adjacent waters.
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Larvae of the spider crabHyas araneus were reared in the laboratory at constant conditions (12°C; 32‰S), and their feeding rate (F), oxygen consumption (R), nitrogen excretion (U), and growth were measured in regular intervals of time during development from hatching to metamorphosis. Growth was measured as dry weight (W), carbon (C), nitrogen (N), hydrogen (H) protein, and lipid. All these physiological and biochemical traits revealed significant changes both from instar to instar and during individual larval moult cycles. AverageF was low in the zoea I, reached a maximum in the zoea II, and decreased again in the megalopa. In the zoeal instars, it showed a bell-shaped pattern, with a maximum in the middle (zoea I) or during the first half of the moult cycle (zoea II). MaximumF in the megalopa was observed still earlier, during postmoult. Respiration (R) increased in the zoeal instars as a linear function of time, whereas it showed a sinusoidal pattern in the megalopa. These findings on variation inF andR during larval development confirm results obtained in previous studies onH. araneus and other decapod species. Excretion (U) was measured for the first time with a high temporal resolution in crab larvae. It showed in all three larval instars a bell-shaped variation pattern, with a maximum near the middle of the moult cycle, and significantly increasing average values from instar to instar. The atomicO/N ratio followed an inverse pattern, suggesting a maximum utilization of protein as a metabolic substrate during intermoult. Growth data from the present study and from a number of previous studies were compiled, showing consistency of growth patterns, but a considerable degree of variability between larvae from different hatches reared under identical conditions. The data show the following consistent tendencies: during the first part of each larval moult cycle (in postmoult, partly in intermoult), lipids are accumulated at a higher rate than protein, whereas an inverse growth patterns is typical of the later (premoult) stages. These two different growth phases are interpreted as periods dominated by reserve accumulation in the hepatopancreas, and epidermal growth and reconstruction (morphogenesis), respectively. Differences between individual larval instars in average biochemical composition and growth patterns may be related to different strategies: the zoeal instars and the early megalopa are pelagic feeding stages, accumulating energy reserves (principally lipids) necessary for the completion of larval development, whereas the later (premoult) megalopa is a semibenthic settling stage that converts a significant part of this energy to epidermal protein. The megalopa shifts in behaviour and energy partitioning from intense feeding activity and body growth to habitat selection and morphogenesis, preparing itself for metamorphosis, i.e. it shows an increasing degree of lecithotrophy. Data from numerous parallel elemental and biochemical analyses are compiled to show quantitative relationships betweenW, C, N, H, lipid, and protein. These regressions may be used as empirical conversion equations for estimates of single chemical components in larvalHyas araneus, and, possibly, other decapods.
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Effects of feeding and starvation on the moult cycle and on the ultrastructure of hepatopancreas cells were studied in Stage I lobster larvae (Homarus americanus Milne-Edwards). The relative significance of yolk and first food was quite different in larvae originating from two females. This difference was evident also in the amounts of stored lipid in the R-cells of the larval hepatopancreas. Most larvae from one hatch were, in principle, able to develop exclusively with yolk reserves (without food) to the second instar. The larvae from the second hatch showed lecithotrophic development only to the transition between late intermoult and early premoult (Stages C/D0 of Drachs's moult cycle) of the first larval instar. When initial starvation in this group lasted for 3 days or more, the point of no return (PNR) was exceeded. After the PNR, consumption of food was still possible, but development ceased in the transition C/D0 or in late premoult (D3–4). It is suggested that these stages of the moult cycle are critical points were cessation of development and increased mortality are particularly likely in early larval lobsters under nutritional stress. Examination of hepatopancreas R-cells suggested that the PNR is caused by an irreversible loss of the ability to restore lipid reserves depleted during initial starvation. Initial periods of starvation ending before the PNR prolonged mainly Stage D0 of the same instar (I). During this delay, structural changes in the R-cells caused by the preceding period of starvation were reversed: reduced lipid inclusions, swollen mitochondria, an increased number of residual bodies indicating autolysis, and a reduction of the microvillous processes. Continually starved larvae which showed lecithotrophic development throughout the first instar and were then re-fed after moulting successfully, had later a prolonged intermoult (Stage C) period in the second instar. This shows that, despite occasional lecithotrophy, food is an important factor in early larval development of the lobster.
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Plankton samples were collected from January 1985 to January 1986 three times per week at Helgoland to study seasonal occurrence and abundance of caridean shrimp larvae. A total of eleven species were obtained. Ninety-one % of all larvae collected during the sample period belonged toCrangon crangon L. andCrangon allmanni Kinahan, 6% toPhilocheras trispinosus Hailstone and 3% to the remaining eight species. Collections were generally dominated byC. crangon larvae. However,C. allmanni larvae were most abundant in June coinciding with hatching activities of the population near Helgoland.C. allmanni was observed to have the highest density of all species with approximately 8 larvae per m3. Larvae ofEualus occultus (Lebour),Eualus pusiolus (Kroyer),Hippolyte varians Leach andAthanas nitescens Leach were most likely released by populations inhabiting the rocky intertidal zone around Helgoland. The presence ofProcessa modica Williamson & Rochanaburanon andProcessa nouveli holthuisi Al-Adhub & Williamson in the German Bight was verified by observations of a series of different developmental stages. Larvae of the rare speciesCaridion steveni Lebour were also recorded. The observed shrimp species were placed into three different groups with respect to their seasonal occurrence. Possible advantages of the timing of larval dispersal relative to predation and food availability are given. The results on seasonal occurrence and relative abundance are discussed in relation to environmental factors (temperature, salinity) as well as to the geographical distribution of the species.
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C and N lost or gained during development and the substratum utilized were determined from measurements of the C and N content of laboratory-reared larvae of the Antarctic krill Euphausia superba Dana from hatching through mid-Calyptopis 1 (Cl, the first feeding stage) at 0, 1 and 2 °C. The relative and absolute use of C and N during early development supported the hypothesis that lipid stores were the primary catabolic substratum during development to Cl and mid-way through Cl. Both measured and calculated estimates of the C required to develop from hatching to Cl indicated that larvae developing at higher temperatures used less energy to reach the Cl stage. After metamorphosis to the Cl stage, internal C reserves continued to satisfy ≈90% of calculated C requirements of fed larvae but the availability of food during this time may be important for development of the digestive enzyme system or of efficient feeding behavior. Development, growth, survival and elemental composition of larvae that experienced delays in food availability after metamorphosis to Cl were observed at 0 and 2 °C. Results of these experiments were used to determine the point-of-no-return (PNR) and quantify the importance of energy reserves to the first feeding stage. If initial delays in food availability exceed a certain point (PNR), starved larvae cannot recover when refed and eventually die. Short delays in food availability affected the timing of development, not the ability to continue to develop, but after iong delays the larvae lost the ability to metamorphose. The PNR depends not only on the initial energy reserves of the embryo but also on the amount of energy needed to reach Cl which in turn depends on the effect of temperature on O2 consumption rates and developmental times. The PNR of 10–14 days from the food availability experiments was similar to the calculated PNR from metabolic requirements and known lipid reserves. The calculated PNR increased with increasing temperature, from 9 to 15 days within the environmental range. Despite the narrow temperature range in the Southern Ocean, both temperature and food availability will affect the survival and recruitment of the early larvae of E. superba.
Article
The lipid biochemistry of environmentally stressed larval fish, bivalves and crustaceans is presented to illustrate the utility of a larval condition index based on lipid composition. Larvae under environmental stress are often unable to obtain sufficient energy from exogenous sources and, as a result, endogenous energy reserves are catabolised to maintain basal metabolism. The storage lipid triacylglycerol (TAG) is of particular importance in this respect, TAG content consequently correlating with the physiological condition of a larva. However, absolute TAG content cannot be directly correlated with larval condition because of its dependency on larval size. It is proposed that TAG content can be correlated with larval condition when expressed in the form of a TAG-sterol ratio that accounts for the size dependency of TAG content. This proposition is supported by data that illustrate a highly positive correlation between sterol content and dry weight for larval herring and larval American lobster. Examples of TAG-sterol ratios are calculated from a survey of studies relating to the lipid class composition of nutritionally and pollutant stressed larvae of some marine fish and crustaceans.
Article
Analysis of the carotenoid pigments of decapod crustaceans has indicated that the visible differences in the appearance of species from different depth horizons are largely due to the disposition of pigments and not to differences in total amount of pigment relative to body size. Species from the upper mesopelagic zone have relatively few large chromatophores, and their chitin is either unpigmented or feebly pigmented. Species from deeper zones have very many small chromatophores and chitin heavily pigmented by unesterified astaxanthin. The major carotenoid pigments of almost all species are astaxanthin and its esters. Lipid analyses have shown that several species of Caridea have very high levels of lipid in the body, and that in many deep-water species wax esters comprise a large proportion of the total lipid. Specific patterns of distribution of the various lipid classes are found in different tissues, with wax esters particularly dominating the hepatopancreas lipids of deep-living species. The patterns of pigment and lipid distribution are discussed in relation to the depth distribution, light environment, and metabolic requirements of the different species.
Article
Since November 1954, the natant epifauna in depths greater than 20 fm. off the Northumberland coast has been sampled at a number of stations with the object of obtaining information on its constitution and fluctuations. The survey has been limited to those species caught with a net of f in. mesh. Seasonal and yearly fluctuations in this fauna must have a considerable effect on the inshore fisheries, for these animals form an important part of the food of marketable fish, yet, this fauna has never beensubject to a detailed analysis over a number of years. Samples have been taken as regularly as weather and other laboratory commitments have allowed. In practice this has been of the order of one sample each month, butat times sampling has been more frequent than this (Table 8). Although sampling is being continued, the material collected so far gives much information on the epifaunaas a whole and the biology of the constituent natant Crustacea and Mollusca.
Article
Large decapod eggs have a relatively high lipid content and low density when compared with small eggs. This relationship is probably a consequence of abbreviated development in those species with large eggs.
Article
1. Changes in chemical composition and caloric content as well as the cumulative efficiencies of yolk utilization have been studied in the developing eggs and freshly hatched protozoea of the shrimpCrangon crangon L. 2. Per unit dry weight of the fresh egg the following relative increases were observed during the development: 16.8% water, 5.9% ash, 10.6% protein, and 0.5% non-protein nitrogen. During the same period (fresh egg to freshly hatched protozoea) fat content decreased from 32.6% to 15.6% and energy content from 6443 to 5287 cal/g dry organic substance. 3. The cumulative efficiencies of yolk utilization for the different constituents varied; they were 70.3% for dry weight, 54.0% for total energy, 83.0% for protein, and 33.6% for fat. 4. Of the 0.0453 cal expended on the metabolic processes of the embryo, only 20.8% was drawn from the oxidation of protein, while fat oxidation contributed as much as 75.0%. 5. Considerable quantities of inorganic salts (0.29µg/egg) were absorbed from the surrounding sea-water by the egg during its development.
Article
Changes in the lipid class and fatty acid composition of developing eggs and unfed larvae of cod (Gadus morhua L.) were studied with the objective of determining probable requirements of cod larvae for dietary lipid. The eggs were collected on 24 March 1985 from holding tanks containing cod which had been caught off the northwest coast of Scotland. Phosphatidylcholine (PC) was the only lipid class to decline in absolute terms during embryogenesis. Catabolism of neutral lipid was initiated during the first week after hatching, and the rate of neutral lipid utilisation increased after the larvae had completely absorbed their yolk sacs. The quantity of triacylglycerol (TAG) remained constant during embryogenesis, but the percentage of 22:6(n-3) int TAG increased substantially during this period. It was calculated that ca. 33% of the 22:6(m-3) released during the process of PC catabolism was incorporated into TAG and sterol ester. The results suggest that PC, replete in appropriate essential fatty acids, should represent a major proportion of the lipid in artificial diets for fish and crustacean larvae.
Article
The influence of starvation on respiration (R), dry weight (W), carbon (C), nitrogen (N), hydrogen (H), and energy content (E; calculated fromC) of spider crab (Hyas araneus L.) larvae was studied in the laboratory. In all larval stages (zoea I and II, megalopa)W increased during postmoult, independent of food, and decreased subsequently. The final reduction inW after continued starvation increased from stage to stage (9, 13, and 20% respectively), but it was always much lower than the decrease inC (44 to 52%),N (42 to 46%),H (50 to 58%), andE (53 to 62%). Individual (R) and weight-specific respiration rates (QO 2) were reduced by 83 to 88%. The time-dependence of these reductions in metabolism and biomass as well as the rates of change in all parameters studied were described with non-linear regression models and differential equations, respectively. Rates and total amounts of energy lost during starvation were independently calculated fromC andR values, and similar results were mostly obtained. Only in the megalopa stage was there a conspicuous difference between the two estimates: higher losses were calculated fromR. This shows that further (biochemical) data are required for a more complete understanding of the energetics of this stage. Estimates of total protein (fromN) and lipid (fromC) suggest that both constituents serve as metabolic substrates during starvation, but most of the energy originates from the breakdown of protein.
Article
Fatty acid biosynthesis in the larval stages of Penaeus japonicus Bate was examined by feeding microencapsulated diets containing (1-14C) palmitic acid, and fat-free diets supplemented with defined fatty acids. Highest larval growth rates were achieved on diets containing Tapes philippinarum lipid and, when defined fatty acids were substituted, on diets containing 20:63 fatty acid. The radioactive tracer experiments indicate that 16:17, 18:0 and 18:19 fatty acids may be synthesised from palmitic acid and that P. japonicus larvae may possess the ability to elongate 18:33 to 20:53 and 22:63, and 18:26 to 20:46. However, the rates of these reactions appear to be too slow to meet the larval requirements for essential fatty acids and the 3 series of polyunsaturated fatty acids must be provided in the diet.
Article
In prior studies, critical points have been found in the interaction between exogenous (food) and endogenous (hormonal) control of the larval development of decapod crustaceans. One of them was termed “point of reserve saturation” (PRS). If it has been passed under conditions of sufficient availability of food, decapod larvae may complete their moulting cycle and develop, independent of food, to the next instar. In the present investigation the PRS and the moulting cycle were studied simultaneously in first instar larvae of nine decapod species belonging to eight families and four higher taxa (Caridea, Macrura, Anomura, and Brachyura). The experimental results suggest that the PRS was in general near the transition between stages C and D0 of Drach's classification system, independent of the species, instar duration, and temperature. When temperature was kept constant throughout the experiment, the critical point was reached after one third to one half of total time of first instar development. If postmoult and intermoult (stages A–C) were prolonged by lower initial temperature, the PRS was delayed but found again near the transition. If starvation began before the PRS, the reserves accumulated from food did not allow successful development to the second instar. Depending on the time of initial feeding, the moulting cycle was arrested at one of the two following critical points
Article
A rapid and simple gas-liquid chromatographic analysis of fatty acids and alcohols of lipid extracts is described. Calanoid copepods were chosen because they contain large amounts of was exters. Lipid extracts were analysed directly after trans-esterification and subsequent hexane extraction. In a single chromatogram fatty acid methyl esters and fatty alcohols were analysed simultaneously on capillary columns coated with SILAR 10 c and CP SIL 5 CB. The major and most of the minor compounds can be determined. Time-consuming separation and purification steps, e.g. by thin-layer chromatography, which can cause loss of polyunsaturated fatty acids and alcohols, are thus avoided.
Article
Larvae of the mud crab Eurypanopeus depressus (Smith) showed higher survival and faster development rate from hatching to the megalopa stage when fed a diet of brine shrimp nauplii than when fed rotifers. When the rotifer diet was supplemented with the lipid-free fraction of brine shrimp in encapsulated form, larval growth was not enhanced. However, when only the lipid fraction of brine shrimp was added in encapsulated form to the rotifer diet, larvae showed enhanced survival and development rate to the megalopa as compared to rotifer-fed controls. When the rotifer diet was supplemented with microcapsules containing specific polyunsaturated fatty acid, enhanced larval survival and development rate resulted. The greatest effect was achieved on docosahexaenoic acid (22 : 6ω3). The results indicate that long-chain polyunsaturated fatty acids, especially 20 : 5ω3 and 22 : 6ω3, are significant in promoting successful development to the megalopa.
Article
A method is described whereby a complete analysis of individual neutral lipid and phospholipid classes in marine animal total lipid can be achieved using an latroscan TLC-FID analyser. The method involves separate analyses of two samples of total lipid in solvents designed to separate neutral and polar lipid classes, together with calibration by a composite standard similar in composition to the sample under analysis. The method does not depend on the degree of unsaturation of the fatty acids present, is rapid and compares well in accuracy with conventional combined gravimetric, colouritnetric, and densitometric procedures.
Article
Lipids are a prominent biochemical class in marine invertebrates, characteristic of wella-fed organisms, and are utilized when the animals are starved. There has been little demonstration of such usage in the field, however, and little consideration of the relationship between the amount of lipid stores and the characteristic life styles of benthic macroinvertebrates. The amount of lipids stored by zooplankton and pelagic invertebrates has been interpreted primarily in terms of availability and variation of food supply and in involvement in buoyancy. Lipid stores in marine invertebrates may serve a variety of roles in the same organism; a primary goal of future work should be to identify and discriminate among them.
Article
This chapter describes the flame ionization detection (FID) applied to thin-layer chromatography (TLC) on coated quartz rods. Iatroscan combines TLC with the FID, and the TLC/FID combination extends TLC sensitivity by two or more orders of magnitude into the range achieved only with gas chromatography. A Chromarod, after development and freeing of solvent, is passed through the hydrogen flame and the carbon ions produced are collected and amplified. In the actual apparatus the housing supports a moving frame, which in the TH-10 model that accepts any number of Chromarods up to 10. The short development time of the Chromarod-S is an encouragement to the sequential use of different solvent systems. By judicious choice of solvent systems, complex separations can often be effected that are not possible with one solvent system, and others can be much improved. A complex mixture of hydrocarbon in a heavy fuel oil could be resolved on a Chromarod-S into four main fractions, which includes asphaltines and polar compounds, resins, polynuclear aromatics and alkyl aromatics, and aliphatics.
The life cycle of Crangon allmanni Kinahan in the Southern North Sea
  • Creutzberg
Creutzberg F. and van Leeuwen F. (1980) The life cycle of Crangon allmanni Kinahan in the Southern North Sea. ICES C.M. 1980/L:71, 11 pp. Criales M. M. (1985) Untersuchungen zur Larval6kologie yon Crangon crangon L. und Crangon allmanni Kinahan (Decapoda, Natantia, Caridea).
Feld- und Laboruntersuchungen über natante Dekapodenlarven in der Deutschen Bucht.
  • Wehrtmann
Wehrtmann I. S. (1986) Feld-und Laboruntersuchungen fiber natante Dekapodenlarven in der Deutschen Bucht. Dipl. Thesis, pp.
Lipids. In The Biochemical Chemistry of Marine Copepods
  • J R Sargent
  • R J Henderson
Sargent J. R. and Henderson R. J. (1986) Lipids. In The Biochemical Chemistry of Marine Copepods (Edited by Corner E. D. S. and O'Hara S. C. M.), pp. 59-108.
Experimental studies on the larval development of the shrimps Crangon crangon and C. allmanni. Helg. Meet
  • Dissertation
Dissertation, pp. 233. Univ. Kiel. Criales M. M. and Anger K. (1986) Experimental studies on the larval development of the shrimps Crangon crangon and C. allmanni. Helg. Meet. 40, 241-265.
Lipid reserves and energy metabolism in the larvae of benthic marine invertebrates
  • Holland
Lipid condition and survival in shrimp (Pandalus borealis)
  • Quellet
Untersuchungen zur Larvalökologie von Crangon crangon L. und Crangon allmanni Kinahan (Decapoda, Natantia, Caridea)
  • Criales
Management of brown shrimp (Crangon crangon) stock in Dutch coastal waters
  • Boddeke