Description: Vampyroteuthis infernalis is a cosmopolitan cephalopod that lives in the oxygen minimum layer between 600 and 800 m depth. Morphometric and physiological studies have indicated that V. infernalis has little capacity for jet propulsion and has the lowest metabolic rate ever measured for a cephalopod. Because fin swimming is inherently more efficient than jet propulsion, some of the reduction in energy usage relative to other cephalopods may result from the use of fins as the primary means of propulsion. V. infernalis undergoes a rapid metamorphosis which consists of changes in the position, size and shape of the fins. This suggests that there are changes in the selective factors affecting locomotion through ontogeny. The present study describes these changes in relation to models for underwater 'flight'. Citrate synthase (CS) and octopine dehydrogenase (ODH) activities, indicative of aerobic and anaerobic metabolism, respectively, were measured in fin, mantle and arm tissue across a range of body size of four orders of magnitude. The low enzymatic activities in both posterior and anterior fin tissue and the relatively high activity in mantle muscle prior to metamorphosis indicate that jet propulsion using mantle contraction is the primary means of propulsion in juvenile V. infernalis. The increase in CS activity with size after metamorphosis suggests an increased use of the fins for lift-based propulsion. Fin swimming appears to be the primary means of propulsion at all adult sizes. The negative allometry of CS activity in mantle and arm muscle is consistent with the scaling of oxygen consumption previously measured for V. infernalis and with the scaling of aerobic metabolism observed in most animals. The unusual positive allometry of fin muscle CS activity suggests that the use of fins is either relatively more important or more costly in larger animals. Positive scaling of ODH activity in all tissues suggests that fin propulsion, jet propulsion and medusoid 'bell-swimming' are all important for burst escape responses. Enzyme activities in Cirrothauma murrayi are consistent with fin-swimming observed from submersibles, while those in Opisthoteuthis californiana suggest a strong reliance on medusoid swimming using the arms. The transition from jet propulsion to paired-fin 'flight' with increasing body size in Vampyroteuthis infernalis appears functionally to be an ontogenetic 'gait-transition'.

Description: The survival of oceanic organisms in oxygen minimum zones (OMZs) depends on their total oxygen demand and the capacities for oxygen extraction and transport, anaerobic ATP production and metabolic suppression. Anaerobic metabolism and metabolic suppression are required for daytime forays into the most extreme OMZs. Critical oxygen partial pressures are, within a range, evolved to match the minimum oxygen level to which a species is exposed. This fact demands that low oxygen habitats be defined by the biological response to low oxygen rather than by some arbitrary oxygen concentration. A broad comparative analysis of oxygen tolerance facilitates the identification of two oxygen thresholds that may prove useful for policy makers as OMZs expand due to climate change. Between these thresholds, specific physiological adaptations to low oxygen are required of virtually all species. The lower threshold represents a limit to evolved oxygen extraction capacity. Climate change that pushes oxygen concentrations below the lower threshold (∼0.8 kPa) will certainly result in a transition from an ecosystem dominated by a diverse midwater fauna to one dominated by diel migrant biota that must return to surface waters at night. Animal physiology and, in particular, the response of animals to expanding hypoxia, is a critical, but understudied, component of biogeochemical cycles and oceanic ecology. Here, I discuss the definition of hypoxia and critical oxygen levels, review adaptations of animals to OMZs and discuss the capacity for, and prevalence of, metabolic suppression as a response to temporary residence in OMZs and the possible consequences of climate change on OMZ ecology.