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Biochemical Composition of Ovary, Embryo, and Hepatopancreas in the Grapsoid Crabs Armases Cinereum and Sesarma Nr

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Biochemical Composition of Ovary, Embryo, and Hepatopancreas in the Grapsoid Crabs Armases Cinereum and Sesarma Nr CBP ELSEVIER Comparative Biochemistry and Physiology, Part B 140 (2005) 455 -463 www.elsevier.comllocate/cbpb Biochemical composition of ovary, embryo, and hepatopancreas In the grapsoid crabs Armases cinereum and Sesarma nr. reticulatum (Crustacea, Decapoda) RE. Hasek], D.L. Felder" Department ofBiology. University ofLouisiana at Lafayette, Po. Box 42451. .' -ayette, LA 70504. USA Received 8 July 2004; received in revised fonn 12 November 2004; accepted 15 November 2004 Abstract Biochemical composition of ovary, embryo, and hepatopancreas tissues in wild populations of Armases cinereum and Sesarma nr. reticulatum were monitored during the reproductive season. Total lipid, carbon, nitrogen, CN ratio, and water concentration of the ovary, hepatopancreas and embryos were quantified over the course of ovarian maturation. Ovary nitrogen concentration decreased as ovaries matured. Ovary lipid and carbon concentration differed significantly over the course ofovarian maturation for both species, but there was no relationship between the concentration or total content of hepatopancreas lipid and the stage of ovarian development in females. Neither species showed a relationship between measures of hepatopancreas lipid and the gonadosomatic index. There was also no simultaneously measurable net decrease in mass of the females' hepatopancreas. Lipid demands of ovarian maturation thus appear to be met in large part by increased dietary intake rather than by substantial draw down of pre-existing lipid stores from the hepatopancreas. While these temperate grapsoid crabs live with putatively fluctuating quality and quantity offood resources, no evidence could be found to demonstrate depletion of lipid concentrations in the hepatopancreas concomitant with ovarian maturation. © 2004 Elsevier Inc. All rights reserved. Keywords: Carbon; Gonadosomatic index; Gulf of Mexico; Hepatopancreas index; Lipid; Ovarian maturation; Nitrogen I. Introduction often connect with burrows of other species (Zimmerman and Felder, 1991). S. nr. reticulatum occurs in densities as Populations of Armases cinereum Bose and Sesarma nr. high as 25 crabs m-2 and is thought to feed on live reticulatum Say are found commonly in intertidal areas near vegetation such as Spartina alterniflora as well as fiddler bayous and in well-drained salt marshes of Louisiana crabs, detritus, and algae (Zimmerman and Felder, 1991). In (Zimmerman and Felder, 1991). S. nr. reticulatum , a species populations of S. reticulatum on the southeastern US from the Gulf of Mexico near (nr.) S. reticulatum , is Atlantic coast a mature male and several females will morphologically, genetically, and physiologically distinct defend their interconnected burrows (Allen and Curran, from S. reticulatum of the Atlantic coast (Zimmerman and 1974; Seiple, 1979). Burrow guarding behaviors have not Felder, 1991; Staton and Felder, 1992; Felder and Staton, been studied in Gulf populations of S. nr. reticulatum, 1994; Mangum and McKenney, 1996). Burrows of S. nr. though we have observed that these crabs also live reticulatum are characterized by an above-ground hood and communally in burrow systems. In Louisiana, the repro­ ductive season of S. nr. reticulatum occurs from April to September (Zimmerman and Felder, 1991). A. cinereum prefers to live along intertidal margins of * Corresponding author. Tel.: +I 3374826743; fax: +I 3374825834. E-mail address: [email protected] (D.L. Felder). marshes under debris and dense vegetation or constructs I Current address. Pennington Biomedical Research Center, 6400 shallow burrows (Seiple, 1979). Commonly, A. cinereum Perkins Road, Baton Rouge, LA 70808, USA. will occupy the abandoned burrow of other species 1096-4959/$ - sec front matter e 2004 Elsevier Inc. All rights reserved. doi: I0.10 16/j.cbpc.2004.11.0 I0 456 B.E. Hasek. D.L. Felder / Comparative Biochem istry and Physiology; Part B 140 (2005) 455-463 (Seiple, 1979). This species does not defend permanent Cypremort Bayou and Vermilion Bay near Cypremort Point, burrows and spends less time in social interactions than LA, USA (29°43' 57.9"N; 91°50' 37.3"W). Specimens does S. reticulatum (Seiple and Salmon , 1982). A. were removed from burrows with shovels or caught as they cinereum appears to be a scavenger that feeds on ran for cover. Crabs were placed in individual vials and decaying plant matter and animals as well as shoots of transported to the laboratory for analysis. Spartina (Seiple and Salmon, 1982). It is primarily nocturnal and during the day hides under debris and 2.2. Morphometric analysis and ovary stage determination vegetation (Seiple and Salmon, 1982). The lipid and relative fatty acid concentration in the Carapace width (CW) was measured with dial calipers larvae of A. cinereum and S. reticula tum have been studied (±O.I mm). Body wet weight WW (WW, ±O.I g), as well in laboratory-hatched broods (Staton and Sulkin, 1991). as dissected ovary and hepatopancreas tissue WW (±O.OOOI Larvae ofA. cinereum had a lower lipid concentration (9%) g) were determined with an analytical balance. Embryos on than did those of S. reticulatum (12%) (Staton and Sulkin, pleopods of ovigerous females were removed and weighed . 1991). Staton and Sulkin (1991) suggested that S. retic­ All tissues were lightly blotted with tissue paper prior to ulatum had high nutritional flexibility, which they defined weighing to remove surface water. Ovary, embryo, and as an ability to develop without high levels of PUFA hepatopancreas tissues were then lyophilized and stored in (polyunsaturated fatty acid) in the diet. S. reticulatum glass vials with Teflonw tops at -80 °C for further analyses. showed resistance to starvation, while A. cinereum showed Gonadosomatic index (GSI) was equal to the ovary WW a complete lack of nutritional flexibility (Staton and Sulkin, divided by the total body WW multiplied by 100 (Giese, (991). It is not known if the lipid concentrations in Atlantic 1966). Digestive gland index (DGI) for females was equal to populations ofS. reticula/urn and the Gulfcoast populations the hepatopancreas WW divided by the total body WW of S. nr. reticula tum are similar. minus ovary WW multiplied by 100 (Clarke, 1977). DGI for Few studies document the dynamic between lipid males was equal to the hepatopancreas WW divided by the concentration and ovarian maturation for semi-terrestrial total body WW multiplied by 100. Ovaries were classified or intertidal erabs (Pillay and Nair, 1973; Mourcntc et al., into three developmental stages (I, II and III) on the basis of 1994; Wcn ct ai., 200 1), and the present study is the first color and size (Ajrnal Khan and Natarajan, 19RO; Mourcntc investigation of lipid concentration and content in er aI., 1994). grapsoid crabs from specifically the northwestern Gulf of Mexico. The purpose of this study was to measure the 2.3. Hepatopancreas color determination lipid, carbon (C), nitrogen (N), and water concentration of the ovary, hepatopancreas and embryos of wild caught During dissection, the hepatopancreas was classified into crabs. Measurement of lipid concentration throughout the one offive color classes. Each class approximated a color in stages of ovarian maturation provides insight to the role of the Pantonew Color Matching System (Pantone, Carlstadt, lipids in crustacean life histories and can indicate whether NJ 07072, USA; colors as numbered below) found in Adobe or not a given crustacean transfers lipid from the Photoshop® (Adobe Systems, San Jose, CA 95110 USA). hepatopancreas to the ovary during gonad maturation. The classes were brown (419CV), which ranged to a brown­ We hypothesized that lipid and C in the hepatopancreas black with black patches; yellow-ochre (l45CV), in which would decrease as ovarian maturation progressed. We the general mass of the hepatopancreas ranged to uniform expected the amounts of lipid and C in the ovary, dark brown-yellow or dark-yellow with brown spots hepatopancreas and embryos would differ between these throughout tubules; white (5875CV), in which hepatopan­ two species with putatively differing habitat and dietary creas tissues were uniform white or cream; lemon-yellow preferences. Finally, we hypothesized there would be (129CV) , in which the general mass of the hepatopancreas differences between males and females in the biochemical and tubules were a uniform yellow; and cadmium-yellow constituents of the hepatopancreas, most probably a result (102CV) , in which the general mass of the hepatopancreas of ovarian maturation influencing the biochemical contents and tubules were a uniform high intensity yellow with little of the hepatopancreas. white. 2.4. Lipid extraction and CHN analysis 2, Materials and methods Lipids were extracted from tissues of individual animals 2.1. Sampling using Parrish's modification of the Folch, Lees, and Sloane Stanley chlorofonnlmethanol extraction method (Parrish, Intermolt adult specimens of A. cinereum and S. nr. 1999). Lipid concentrations were determined gravimetri­ reticulatum (Crustacea, Decapoda, Sesarmidae) were cally by measuring mg of lipid (±O.I mg) g-I dry weight sampled during their reproductive season in March-July (DW) tissue with an analytical balance. Total lipid content 200 I. Crabs were collected from coastal wetlands between was determined by multiplying lipid concentration by organ BE Hasek, DL Felder / Comparative Biochemistry and Physiology, Part B 140 (2005) 455-463 457 Table I which factors were different. A significance level of 0.05 Biometric means±SD for females at different stages of ovarian maturation was applied for all tests in this study. One-factor ANOVA Armases cinereum Ovary stages was used to test for differences between ovary stage and I (II 6) II (n 4) III (n I") hepatopancreatic nutrient concentration as well as total CW 16.5±4 18.2±1.2 17.2±3.3 organ content; regression analyses was used to determine Total WW 3.3± I 3.7±1.1 3.2± 1.1 the relationship between OSI and hepatopancreatic nutrient b Ovary WW 0.04±0.0I' 0.1±0.04 0.16±0.06 concentration, as well as total organ content. GSI 1.2±0.5" 2.7±I' 5.312b Hepatopancreas WW 0.1810.07 0.310.1 0.16±0.04 Sesarma nr.
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