Physiological and biochemical changes during the larval development of a brachyuran crab reared under constant conditions in the laboratory

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.