NUTRITIONAL STATUS RESPONSE of Daphnia Laevis and Moina
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FOOD QUALITY AND NUTRITIONAL STATUS OF TROPICAL CLADOCERANS 555 NUTRITIONAL STATUS RESPONSE OF Daphnia laevis AND Moina micrura FROM A TROPICAL RESERVOIR TO DIFFERENT ALGAL DIETS: Scenedesmus quadricauda AND Ankistrodesmus gracilis MACEDO, C. F. and PINTO-COELHO, R. M. Programa de Pós-graduação em Ecologia, Conservação e Manejo da Vida Silvestre (ECMVS), Universidade Federal de Minas Gerais (UFMG), C.P. 486, CEP 30161-970, Belo Horizonte, MG, Brazil Correspondence to: Carla Fernandes Macedo, Rua Visconde do Rio Branco, 72/201, Centro, CEP 36770-000, Cataguases, MG, Brazil, e-mail: [email protected] Received June 21, 2000 – Accepted August 28, 2000 – Distributed November 30, 2001 (With 4 figures) ABSTRACT The accumulation of lipid reserves was investigated in two common cladoceran species typical of neotropical freshwaters. Experiments were performed in order to measure differential accumulation of lipid reserves in response to two algal diets, S. quadricauda and A. gracilis. The larger D. laevis fed with the Ankistrodesmus diet accumulated a higher amount of total lipids. The average lipid content for both diets was 11.1% and 22.1% dry weight for Scenedesmus and Ankistrodemsus, respectively. This difference was found to be highly significant. The superior nutritional quality of A. gracilis was confirmed by the experiments using the smaller Moina micrura. This cladoceran accumulated 11.4% and 19.9% of the average dry weight of lipids from Scenedesmus and Ankistrodesmus, respectively. The study also found that most lipid reserves are concentrated as triacylglycerols in both cladocer- ans. The relative contribution of this class of lipids also varied according to the diet. For Daphnia, for instance, the average triacylglycerol levels varied from 6.2 to 11.0 with the Scenedesmus and An- kistrodesmus diets, respectively. Key words: variation nutritional, cladocerans, chlorophyceae, experiments. RESUMO Respostas ao estado nutricional de Daphnia laevis e Moina micrura em um reservatório tropical para diferentes dietas algais: Scenedesmus quadricauda e Ankistrodesmus gracilis O presente estudo investigou o acúmulo de reservas de lipídeo em duas espécies comuns de cladóceros neotropicais, típicas de água doce. Foram desenvolvidos experimentos para medir o acúmulo dife- renciado de reservas lipídicas em resposta a duas dietas algais, S. quadricauda e A. gracilis. O cladócero maior, D. laevis, alimentado com a dieta Ankistrodesmus acumulou uma quantidade maior de lipídeo total. As médias de lipídeo total para ambas as dietas foram 11,1% e 22,1% de peso seco para Scenedesmus e Ankistrodesmus, respectivamente. Essa diferença encontrada foi altamente significativa. A qualidade nutricional mais elevada de A. gracilis foi confirmada pelos experimentos em que foi utilizada uma espécie menor de cladócero, Moina micrura. Esse cladócero acumulou valores médios de lipídeo total de 11,4% e 19,9% de peso seco para Scenedesmus e Ankistrodesmus, respecitvamente. Este estudo também revelou que a maior parte dos lipídeos estocados em ambos os cladóceros refere- se a triglicérides. A contribuição relativa dessa classe de lipídeo também variou de acordo com a dieta. Para Daphnia, por exemplo, os teores médios de triglicérides variaram de 6,2% a 11,0% de peso seco total com as dietas de Scenedesmus e Ankistrodesmus, respectivamente. Palavras-chave: variação nutricional, cladóceros, clorofíceas, experimentos. Braz. J. Biol., 61(4): 555-562 556 MACEDO, C. F. and PINTO-COELHO, R. M. INTRODUCTION According to Goulden & Henry (1988), the energy reserves of cladocerans seem to be used Lipids are the most important energy reser- in two different ways: a) the adults use them for ves for marine as well as freshwater zooplankton. their own metabolic needs; or b) the reserves are Contrasting with the large literature about lipids on transferred to the offspring through eggs and are marine zooplankton, there is a scarcity of published used by the young animals in their initial deve- works about lipids in freshwater zooplankton (Far- lopment. The plankton microcrustaceans represent kas, 1970; Goulden & Horning, 1980; Ahlgren et one of the most important links between the pro- al., 1990; Vanderploeg et al., 1992; Ahlgren et al., ducers and the fishes in the aquatic food chain. 1997). In planktonic crustaceans, the lipids may have Thus, the bioenergetics of zooplankton affects the important functional and structural properties. Since biochemical composition and energetic dynamics these compounds are of low density, high percentiles of fishes as well (Farkas, 1970). According to the of lipids in the biomass may be helpful for animal same author, several methodological problems have flotation. Lipids are also colorless, an important contributed to the scarcity of published information characteristic that protects against visual predators concerning the dynamics of fats in planktonic in a highly transparent environment. The lipids also crustaceans. These include sample treatment and have the capacity to accumulate nearly twice as much appropriate methods since most of the existing energy as proteins and carbohydrates in their procedures are not suitable for measuring lipids chemical bonds. Lipid levels can vary with several in such organisms (Gardner et al., 1985). environmental factors. They also can be used as The availability of new methodologies of indicators of the degree of disturbance of an eco- sample treatment (lyophilization, deep-freezing) system (Arts, 1999). The crustacean zooplankton as well as the adaptation or improvement of older frequently contains a large amount of lipids (Sterner analytical techniques have opened the possibility et al., 1992). of performing simple, precise, and reliable quan- Most copepods and cladocerans accumulate titative measurements of different lipid classes such large amounts of lipids as energy reserves. In ma- as triacylglycerols, phospholipids and fatty acids, rine copepods living in the deep sea or in the Arctic in addition to total lipids (De Mott & Müller-Na- Ocean, up to 60% of their biomass can consist of varra, 1997; Lürling et al., 1997; Weers et al., lipids reserves, mainly as fatty acids. Copepods 1997; Sekino et al., 1997; Weers & Gulati, 1997). living in superficial water of temperate oceans and In the neotropics, zooplankton is typically freshwater copepods and cladocerans have tria- dominated by small organisms, especially copepods cylglycerol as the main form of energy reserve. of the genus Thermocyclops spp., cladocerans such These compounds can reach up to 40% of body as Bosmina and Ceriodaphnia, and several species biomass. These reserves can be used by the adults of rotatoria (Lopes et al., 1997; Attayde & Bozelli, to satisfy their metabolic needs during periods of 1998). However, under certain circumstances, i.e., food limitation (Goulden & Henry, 1988). in the presence of eutrophication, the proportion Abyssal marine copepods accumulate tria- of larger cladocerans in the zooplankton increases cylglycerols and fatty acids. The triacylglycerols and they may often become dominant or sub-do- can be promptly used to sustain basal metabolism minant. Some of these species are: Daphnia laevis, during brief periods of hunger. The fatty acids, Diaphanosoma birgei, and Moina micrura (Pinto- in turn, are used when the animals are exposed to Coelho, 1998). several months of low food levels. Species of The objective of the present investigation was marine copepods living at the surface accumulate to answer the following questions: a) how much only triacylglycerol that can be quickly metabolized lipid can a large tropical cladoceran accumulate to sustain vital activities when food is temporarily under optimal food conditions? b) Is triacylglicerol scarce, or when they are looking for a better feeding the major lipid class accumulated by such orga- place. The energy reserves of freshwater species, nisms? c) Is there a significant difference in lipid such as Daphnia, are also mainly triacylglycerols accumulation between two often dominant cla- (Goulden & Horning, 1980). docerans in neotropical freshwaters? d) To what Braz. J. Biol., 61(4): 555-562 FOOD QUALITY AND NUTRITIONAL STATUS OF TROPICAL CLADOCERANS 557 extent can variations in food quality affect lipid with hot concentrated sulfuric acid. After the reserves? addition of a solution of vanillin and phosphoric acid, a red complex is formed which is measured MATERIAL AND METHODS with a Shimadzu UV 1201 spectrophotometer (546 nm). The lipid values are expressed as dry weight Two zooplankton organisms, Daphnia laevis percent. and Moina micrura, were collected in a nearby The triacylglycerol analyses were carried out eutrophic reservoir, the Pampulha Reservoir, lo- colorimetrically after lipid extraction using a mo- cated in Belo Horizonte, Brazil, and cultivated in dified Folch method (Folch et al., 1957). The lipid the laboratory. The organisms received a diet extraction procedure began with a conventional consisting of 3.0 to 4.0 × 104 cells.ml–1 of a mixed extraction using a chrophorm: methanol solution culture of Scenedesmus quadricauda and Ankis- (2:1), followed by different washings and centri- trodesmus gracilis algae. The water of these cul- fugation with methanol and chlorophorm with the tures was first changed every three days using addition of NaCl and Foch solution (Chlorophor- reservoir water. It was then replaced with recons- mic: Methanol: water, 8:4:3). The samples were tituted water as proposed by Cetesb (1994). dried under pure gaseous N2 and