SUMMARY To test the hypothesis that the preference for ketone bodies rather than lipids as oxidative fuel in elasmobranchs evolved in response to the appearance of urea-based osmoregulation, we measured total non-esterified fatty acids (NEFA) in plasma as well as maximal activities of enzymes of intermediary metabolism in tissues from marine and freshwater elasmobranchs,including: the river stingray Potamotrygon motoro (<1 mmol l –1 plasma urea); the marine stingray Taeniura lymma , and the marine shark Chiloscyllium punctatum (>300 mmol l –1 plasma urea); and the euryhaline freshwater stingray Himantura signifer , which possesses intermediate levels of urea. H. signifer also were acclimated to half-strength seawater(15‰) for 2 weeks to ascertain the metabolic effects of the higher urea level that results from salinity acclimation. Our results do not support the urea hypothesis. Enzyme activities and plasma NEFA in salinity-challenged H. signifer were largely unchanged from the freshwater controls, and the freshwater elasmobranchs did not show an enhanced capacity for extrahepatic lipid oxidation relative to the marine species. Importantly, and contrary to previous studies, extrahepatic lipid oxidation does occur in elasmobranchs, based on high carnitine palmitoyl transferase (CPT) activities in kidney and rectal gland. Heart CPT in the stingrays was detectable but low,indicating some capacity for lipid oxidation. CPT was undetectable in red muscle, and almost undetectable in heart, from C. punctatum as well as in white muscle from T. lymma . We propose a revised model of tissue-specific lipid oxidation in elasmobranchs, with high levels in liver,kidney and rectal gland, low or undetectable levels in heart, and none in red or white muscle. Plasma NEFA levels were low in all species, as previously noted in elasmobranchs. D-β-hydroxybutyrate dehydrogenase( d -β-HBDH) was high in most tissues confirming the importance of ketone bodies in elasmobranchs. However, very low d -β-HBDH in kidney from T. lymma indicates that interspecific variability in ketone body utilization occurs. A negative relationship was observed across species between liver glutamate dehydrogenase activity and tissue or plasma urea levels, suggesting that glutamate is preferentially deaminated in freshwater elasmobranchs because it does not need to be shunted to urea production as in marine elasmobranchs.

SUMMARY The effects of long-term fasting and subsequent refeeding on digestive physiology and energy metabolism were investigated in a subterranean aquatic crustacean, Stenasellus virei , and in a morphologically similar surface-dwelling species, Asellus aquaticus . Metabolic response to food deprivation was monophasic in A. aquaticus , with an immediate,large decrease in all energy reserves. In contrast, S. virei displayed three successive periods of phosphageno-glucidic, lipidic and,finally, proteo-lipidic-dominant catabolism over the course of the nutritional stress. To represent the responses of subterranean crustaceans to food stress and renutrition, a sequential energy strategy was hypothesized, suggesting that four successive phases (called stress, transition, adaptation and recovery) can be distinguished. Based on these results, a general adaptive strategy for groundwater organisms was proposed. Their remarkable resistance to long-term fasting may be partly explained by (1) a depressed metabolism,during which they mainly subsist on lipid stores, (2) a prolonged state of glycogen- and protein-sparing, (3) low energetic requirements and (4) large body stores. In addition, these groundwater species displayed high recovery abilities during refeeding, showing an optimal utilization of available food and a rapid restoration of their body reserves. These adaptive responses might be considered for numerous subterranean organisms as an efficient energy-saving strategy in a harsh and unpredictable environment where fasting(and/or hypoxic) periods of variable duration alternate with sporadic feeding events (and/or normoxic periods). Therefore, food-limited and/or hypoxia-tolerant groundwater species appear to be good examples of animals representing a low-energy system.