Mysid crustaceans as standard models for the screening and testing of endocrine-disrupting chemicals

Author Posting. © Springer, 2007. This is the author's version of the work. It is posted here by permission of Springer for personal use, not for redistribution. The definitive version was published in Ecotoxicology 16 (2007): 205-219, doi:10.1007/s10646-006-0122-0.Investigative efforts into the potential endocrine-disrupting effects of chemicals have mainly concentrated on vertebrates, with significantly less attention paid to understanding potential endocrine disruption in the invertebrates. Given that invertebrates account for at least 95% of all known animal species and are critical to ecosystem structure and function, it remains essential to close this gap in knowledge and research. The lack of progress regarding endocrine disruption in invertebrates is still largely due to: (1) our ignorance of mode-of-action, physiological control, and hormone structure and function in invertebrates; (2) lack of a standardized invertebrate assay; (3) the irrelevance to most invertebrates of the proposed activity-based biological indicators for endocrine disruptor exposure (androgen, estrogen and thyroid); (4) limited field studies. Past and ongoing research efforts using the standard invertebrate toxicity test model, the mysid shrimp, have aimed at addressing some of these issues. The present review serves as an update to a previous publication on the use of mysid shrimp for the evaluation of endocrine disruptors (Verslycke et al., 2004a). It summarizes recent investigative efforts that have significantly advanced our understanding of invertebrate-specific endocrine toxicity, population modeling, field studies, and transgeneration standard test development using the mysid model.Supported by a Fellowship of the Belgian American Educational Foundation

Seasonal reproduction, multiple spawning, and fecundity in northern krill, Meganyctiphanes norvegica, and Antarctic krill, Euphausia superba

2nd International Symposium on Krill, UNIV CALIF SANTA CRUZ, SANTA CRUZ, CALIFORNIA, AUG 23-27, 1999Both northern krill, Meganyctiphanes norvegica, and Antarctic krill, Euphausia superba, release eggs seasonally in multiple spawning events. Fecundity estimates were determined based on counts of mature oocytes in the ovary of females in preserved field samples. Principal components analysis and allometry were used to analyze the covariation of egg-batch size with female characteristics. In both krill species, egg-batch size scaled isometrically with ovarian weight but showed a positive allometry with body size. Predicted egg-batch size ranged from 200 to 4000 eggs for M. norvegica of 25.5-35 mm body length and from 1500 to 6000 eggs for E. superba of 36-55 mm. Allometric relationships can be used to predict annual fecundity from size structure data of krill populations. In this study, the effect of temperature on the duration of molting and spawning cycles was used to infer a number of reproductive cycles per year. taking into account climate and seasonal variation

Life cycle strategies of Northern krill (Meganyctiphanes norvegica) for regulating growth, moult, and reproductive activity in various environments: the case of fjordic populations

Adaptive strategies of two fjord populations of Northern krill (Meganyctiphanes norvegica Sars) were studied and compared with other populations from different climatic and oceanic conditions. The Gullmarsfjord (West Sweden) and the Clyde Sea (West Scotland) resident populations followed the same basic pattern of development as the Kattegat (between Denmark and Sweden) and the Ligurian Sea (Northwest Mediterranean) populations, but the fjord krill reached a comparatively larger body size during their second year of life. The positive relationship between body size and fecundity means that fjord populations are potentially more productive than those of the open sea. High rates of moulting and spawning activity were limited to the spring and summer despite the fact that trophic conditions still seemed favourable in early autumn. We show here that, in autumn, the adult moult cycle became longer and growth stopped, large 2-year-old krill disappeared from the population and ovarian development was arrested. Resources already accumulated ill the oocytes (glycoproteic and lipid yolk) were recovered by oosorption and ovaries regressed for a winter rest. Autumn trophic conditions were still favourable in both sites, especially in terms of copepod abundance. However, the phytoplankton community changed from a dominance of diatoms in early season to dinoflagellates in late summer. We suggest that these changes triggered the autumn arrest of krill production

Ovarian development and spawning in relation to the moult cycle in Northern krill, Meganyctiphanes norvegica (Crustacea : Euphausiacea), along a climatic gradient

Adaptive processes linked to reproduction were studied comparatively for three populations of Northern krill, Meganyctiphanes norvegica (M. Sars, 1857), sampled during winter and summer cruises in the Clyde Sea (W Scotland), the Kattegat (E Denmark), and the Ligurian Sea (NW Mediterranean). The aim was to investigate the functional relationship between egg production and moulting under contrasted climatic and environmental conditions. A staging system for female sexual development established for live krill was complemented by a histological study of the ovary at various developmental steps. During the reproductive season, all adult female krill were engaged in cyclical egg production. During experiments, female krill released one batch of mature oocytes in one or two spawning events. The ovary of postspawn female krill still contained developing oocytes for another egg batch. In the non-reproductive period, all female krill had a resting ovary. Ovarian structure and pattern of egg production were identical in the three populations, but seasonal timing of egg production was different. The model proposed for the Ligurian population of the annual cycle of ovarian development can be extended to the other two populations, taking into account the seasonal characteristics of each site. Random field samples were staged simultaneously for moult cycle and for sexual development. Moult stages and the seasonal variation of the intermoult period were studied for the Kattegat population using multi-year data and compared to data obtained during summer/winter cruises in the Clyde and the Ligurian Sea. At the three sites, intermoult period was shorter and temperature-dependent during the reproductive period, concurrent with the season of greatest food availability. During most of the year and the period of sexual rest, moulting activity was reduced. The relationship between spawning and the moult cycle was studied comparatively for the three populations. Eggs were released during the premoult phase of a ``spawning moult cycle'', in one or two spawnings associated with apolysis and Moult Stage D1, respectively. Yolk accumulation for the next egg batch was completed during an alternating ``vitellogenic moult cycle''. A model for the timing of cyclical egg production in relation to moulting, as proposed for the Kattegat, can be extended to the other populations, taking into account intermoult period variation with temperature. Temperature appeared to be the principal environmental factor controlling growth (through moulting) and egg production during the reproductive season, in connection with favourable trophic conditions

External parasite infestation depends on moult-frequency and age in Antarctic krill (Euphausia superba)

Levels of infestation by the suctorian Ephelota sp. were determined in Antarctic krill taken in the north-eastern region of the Scotia sea, close to the island of South Georgia. Individuals caught in summer between 2002 and 2004 were measured for body length, eye diameter, moult stage, sex, sexual maturity and number of suctorian parasites. All krill samples were found to contain at least two modal size-classes and the complete range of maturity- and moult stages. Sample analysis for the different characters identified moult stage as an important determinant of infestation by Ephelota sp., with 66% of those about to moult (pre-moult individuals) being infested compared with 0% of post-moult individuals. Multiple Correspondence Analysis also identified a strong link between age and degree of suctorian infestation with large-eyed (and hence old) males having very high numbers of suctorian bodies. Moulting is costly in terms of energy and vulnerability, but it is believed that krill moult at relatively high rates throughout their life to facilitate growth and "shrinkage". Here, we demonstrate that the control of external parasitism is also a major advantage of moulting at high frequency. The capacity of krill to maintain a high moult rate decreases with age, leading to older krill enduring higher parasitic loads

Variation of lipid classes among organs of the Northern krill Meganyctiphanes norvegica, with respect to reproduction

Lipid content and class in the Northern krill Meganyctiphanes norvegica (digestive gland, stomach, gonad, fat body, abdomen) was investigated and correlated with sex and reproductive stage. Ready to spawn females, have high lipid content in ovaries, while in males and spent females, the major site of lipid deposits was the digestive gland, followed by the fat body. These differences among spawning and spent females are indicative of strong interactions between the ovary and digestive gland and the ovary and fat body during vitellogenesis. Triacylglycerols (TAG) were the major neutral lipid class with high levels in the digestive gland. The major phospholipid was phosphatidylcholine (PC) particularly in the muscular tissue of the abdomen. Phosphatidyl-ethanolamine (PE) and -serine-inositol (PS-PI), were present at intermediate levels. Reproductive males were depleted in TAG and diacylglycerols (DAG) in the digestive gland, gonad and fat body, and had 4 times lower cholesterol in the gonad than ready to spawn females. Furthermore, ready to spawn females had in the ovary higher amounts of TAG, DAG and phospholipids (PC, PE, PS-PI) than spent females. Linear relationships between lipid content and main lipid class (TAG, PC, PE, PS-PT) in different fractions of males and ready to spawn females showed that: (1). TAG was stored for both sexes in all cephalothorax fractions with highest values in the digestive gland and ovary fluid; (2). PC was accumulated for both sexes in the fat body and the gonad with a higher slope for females, with the highest values in the ovary fluid and in the abdomen of males and that (3). PS-PI was stored only in the ovary and abdomen of mature females. These results are discussed in terms of the strategy developed by Meganyctiphanes norvegica to allocate lipids to the next generation for optimised embryogenesis. (C) 2001 Elsevier Science Inc. All rights reserved

Vertical migration behaviour in the northern krill Meganyctiphanes norvegica is influenced by moult and reproductive processes

International audienceA population of the northern krill Meganyctiphanes norvegica was sampled between 24 July and 3 August 1998 in the Alkor Deep, Kattegat, in order to investigate the influence of sexual and developmental factors on the vertical distribution of adult krill. Depth-discrete samples of krill were taken with a 1 m(2) MOCNESS net at the cardinal times of day (midnight, midday, sunrise and sunset). Specimens were immediately measured and categorised for moulting and spawning status. Further samples were preserved for detailed analysis in the home laboratory. Results showed that the population concentrated in the deep (80 to 100 m) during day-time but segregated vertically during nighttime. Moulting occurred in the deep during night-time, away from the main part of the non-moulting population, which was located between 80 and 50 m. Spawning females were most evident in the up permost depth interval (30 to 5 m). Moulting at night in the deepest layers may be viewed as a mechanism to avoid cannibalism whilst in a vulnerable condition. Spawning in the warm upper layers accelerates reproductive processes and may also reduce the depth to which the eggs sink before hatching into nauplii

Chromosome diversity in Mediterranean and Antarctic euphausiid species (Euphausiacea)

Chromosome number and morphology were studied in gonadal tissue from eight euphausiid species, using an air-drying technique and Giemsa staining. Among Mediterranean species, haploid chromosome numbers were: n = 19 in Euphausia hemigibba, Euphausia brevis, and Nematoscelis megalops, n = 18 in Euphausia krohni, and n = 11 in Nyctiphanes couchi. Among Antarctic species, haploid chromosome numbers were: n = 17 in Euphausia superba, n = 20 in Thysanoessa macrura, and n = 13 in Thysanoessa vicina. Chromosome morphology was further studied for three species after chromosome measurements: Euphausia hemigibba karyotype showed 17 metacentric and 2 submetacentric chromosome pairs, Euphausia krohni 17 metacentric and 1 submetacentric chromosome pairs, and Nematoscelis megalops 17 metacentric and 2 submetacentric chromosome pairs. Three species sharing the same chromosome number of n = 19, i.e., Euphausia hemigibba, Nematoscelis megalops, and Meganyctiphanes norvegica were distinguished by a different slope of the decrease in chromosome size and by the presence/absence of metacentric and submetacentric chromosome pairs differing in number and position. Chromosome morphology of other species was derived from the observation of meiotic metaphases II. A majority of metacentric chromosomes was also displayed. Cytotaxonomic differences in haploid chromosome number allow the discrimination of morphologically neighboring sympatric species (e.g., Meganyctiphanes norvegica and Nyctiphanes couchi; Thysanoessa vicina and T. macrura). Euphausiids share common chromosomal features which clearly separate them from other eucarids, i.e., a narrow range of haploid chromosome number, a large chromosome size, and a majority of metacentric chromosomes

The scheduling of spawning with the moult cycle in Northern krill (Crustacea : Euphausiacea): a strategy for allocating lipids to reproduction

8th International Congress on Invertebrate Reproduction, AMSTERDAM, NETHERLANDS, AUG 10-14, 1998Euphausiids moult and grow throughout their life, which implies sharing of resources between growth and reproduction for adult krill. In the Northern krill, Meganyctiphanes norvegica (M. Sars), female krill produce eggs cyclically. Spawning moult cycles alternate with vitellogenic moult cycles for lipid yolk accumulation. Histology shows that lipids are associated with the R cells of the digestive gland in both sexes, with the yolk platelets of mature oocytes and with the fat body cell membranes and blood lacunae in reproducing females. Mature female krill can have a total lipid content twice as high as males, mostly due to accumulation in the ovary, the fat body and the haemolymph. In contrast, in males, as well as in non-reproducing females, the highest percentage of lipids is found in the digestive gland and the haemolymph. In Meganyctiphanes norvegica, the most abundant lipid fractions are polar lipids and triglycerides, the latter being relatively low in reproducing female gonad and fat body. Triglycerides are believed to be a pure energy source and polar lipids are essential for membrane development in embryos. The fatty acid content and composition of the triglyceride and polar lipid fractions in females are different from males, related to both reproductive and dietary processes. Higher levels of polyunsaturated fatty acids (PUFA) in the polar lipid fraction were found in reproductive females. During the non-reproductive season, the converse was found, indicating the specific role PUFA and other fatty acids play in growth and egg production. Adaptive processes linked to reproduction were studied comparatively in three populations of the Northern krill - Clyde Sea (W, Scotland), Kattegat (E, Denmark), Ligurian Sea (Mediterranean) - all differing considerably in climatic and trophic conditions. Such adjustments in lipid synthesis and storage are viewed as reproductive strategies developed by the Northern krill in response to different environmental conditions

Chapter 6.9 Southern Ocean Euphausiids

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