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In pursuit of Irving and Scholander: a review of oxygen store management in seals and penguins

Ponganis, Paul J.; Meir, Jessica U.; Williams, Cassondra L.

Journal of Experimental Biology, 2011

Peer-reviewednarrative reviewmixed

A review of how diving birds and mammals store and ration oxygen, drawing on blood and muscle oxygen measurements, heart-rate records and back-mounted sensors from freely diving animals rather than from forced laboratory dives.

Studied in
Aptenodytes forsteri, Pygoscelis adeliae, Phocidae
Sample size
review of diving physiology across seals and penguins

Diving capacity rests on large oxygen stores in blood and muscle rather than in the lungs, on high myoglobin concentrations, and on active management: heart rate falls sharply and blood flow is redistributed so that oxygen goes where it is needed. Emperor penguins routinely dive beyond the point at which their oxygen stores would be exhausted at surface metabolic rates, tolerating lactate accumulation on the longest dives.

What it means

The authors' reading, and ours. Where they differ, that difference is the point.

How the authors put it

Diving performance is a matter of oxygen store size combined with regulated distribution, and freely diving animals behave differently from restrained ones, which had distorted the earlier literature.

How NatureHQ reads it

The picture people carry is a big breath held for a long time, and almost every part of that is wrong. An emperor penguin exhales rather than inflating, because air is buoyant and compressible and both are problems at depth. The oxygen is carried in blood and muscle, and the animal then rations it — dropping heart rate, shutting down non-essential circulation. What looks like breath-holding is closer to controlled shutdown. The review also makes the honest point that much of the older literature came from animals forced to dive, which measures something different from what a bird does when it chooses.

  • A review across two very different groups; the shared principles hide substantial differences.
  • Direct blood oxygen measurement in free-diving birds is possible only in a few species and a few individuals.
  • Instrumented animals may dive differently from uninstrumented ones.

What this study is used for on NatureHQ

One study can inform several subjects. Here is everywhere this one is cited.

Published by: The Company of BiologistsPublished by The Company of Biologists.

doi.org/10.1242/jeb.031252

This reference was resolved automatically against Crossref on 2026-08-11.

NatureHQ summarises research in its own words and does not reproduce published text. Reviewed 2026-08-11.