Energetics: The Costs of Living and Reproducing for an Individual Cephalopod
Cephalopods, like all other animals, have to decide how to allocate resources; maintenance processes, growth of somatic and reproductive tissues, and locomotor activity all have costs. We should like to be able to identify these costs and discover how efficiently cephalopods make use of the prey that they capture and digest. Cephalopods generally grow fast and mature rapidly; a first task is to determine how accurately laboratory studies reflect growth in the wild, because much of the information we need (such as food conversion efficiencies, excretion rates or the costs of locomotion) can be collected only from animals kept in the laboratory. Comparison of laboratory feeding and growth rates for octopods, sepioids and teuthoids with fisheries data suggests that data collected from cephalopods fed ad libitum in the laboratory may be used validly to construct energy budgets representative of individuals in the wild. The immediate cost of feeding (the specific dynamic action) has been thoroughly documented in Octopus, as has the longer-term elevation or depression of metabolic rate by feeding or starvation; it is assumed that similar costs will be found in squid. The cost of locomotion has been studied in both octopods and squid, but we have only limited data on how much time the animals spend moving, and how rapidly, in the wild. Excretory and faecal losses are assessed from laboratory studies, and maintenance costs estimated from feeding rates that just maintain body mass in the laboratory. Comparison of gross and net food conversion efficiencies suggest that squid convert food into tissues less efficiently than octopods, owing primarily to their greater time spent in locomotion. We present a representative series of energy budgets for octopods (based on Octopus) and squids (based on Illex and Loligo), for starving, feeding, migrating and maturing individuals. A major contrast is provided by Nautilus, which lives for ten or twenty years and grows only slowly. Finally we speculate on the possible biochemical and historical factors that may have limited the adaptive radiation of cephalopods, resulting in a group lacking herbivores, detritivores or filter-feeders but extremely successful as carnivores.