Oxygen extraction and jet propulsion in cephalopods

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Abstract
The ventilatory system of cephalopods has evolved so that the animals minimise the energetic cost of either oxygen extraction or jet propulsion. Optimal design for jet propulsion requires a large stroke volume moving water through the system with minimal acceleration, so the oxygen-carrying capacity of the ventilatory stream is always greatly in excess of requirements. Oxygen uptake in a jet-propelled animal must be independent of the volume of the jet to avoid locking locomotion to oxygen uptake. Any such link is incompatible with the repayment of an oxygen debt after exercise, with added oxygen demand during digestion, and with regulation of uptake under hypoxic conditions. Cephalopods evolved for efficient jet propulsion must be able to alter the rate of oxygen extraction from the ventilatory stream. Squids and Nautilus do this; oxygen extraction is normally low, in the range 5-10% during jet-propelled cruising at subcritical speeds, but can rise to as much as 25% in acute hypoxia or at rest after exhausting exercise. Other cephalopods, such as Octopus, minimise the cost of oxygen extraction by propelling a minimal volume of water through the gills, extracting 35-50% (exceptionally 75%) of the available oxygen. This leaves little scope for a further increase in extraction, and any added demand is met by increasing the ventilation stroke volume. A consequence is that jet propulsion becomes extravagant; octopuses showed much greater jet pressures than squid when they choose to swim. The two alternative specialisations are linked to life-style and cut across systematic boundaries.

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