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2015
Ponganis, PJ, St Leger J, Scadeng M.  2015.  Penguin lungs and air sacs: implications for baroprotection, oxygen stores and buoyancy. Journal of Experimental Biology. 218:720-730.   10.1242/jeb.113647   AbstractWebsite

The anatomy and volume of the penguin respiratory system contribute significantly to pulmonary baroprotection, the body O-2 store, buoyancy and hence the overall diving physiology of penguins. Therefore, three-dimensional reconstructions from computerized tomographic (CT) scans of live penguins were utilized to measure lung volumes, air sac volumes, tracheobronchial volumes and total body volumes at different inflation pressures in three species with different dive capacities [Adelie (Pygoscelis adeliae), king (Aptenodytes patagonicus) and emperor (A. forsteri) penguins]. Lung volumes scaled to body mass according to published avian allometrics. Air sac volumes at 30 cm H2O (2.94 kPa) inflation pressure, the assumed maximum volume possible prior to deep dives, were two to three times allometric air sac predictions and also two to three times previously determined end-of-dive total air volumes. Although it is unknown whether penguins inhale to such high volumes prior to dives, these values were supported by (a) body density/buoyancy calculations, (b) prior air volume measurements in free-diving ducks and (c) previous suggestions that penguins may exhale air prior to the final portions of deep dives. Based upon air capillary volumes, parabronchial volumes and tracheobronchial volumes estimated from the measured lung/airway volumes and the only available morphometry study of a penguin lung, the presumed maximum air sac volumes resulted in air sac volume to air capillary/parabronchial/tracheobronchial volume ratios that were not large enough to prevent barotrauma to the non-collapsing, rigid air capillaries during the deepest dives of all three species, and during many routine dives of king and emperor penguins. We conclude that volume reduction of airways and lung air spaces, via compression, constriction or blood engorgement, must occur to provide pulmonary baroprotection at depth. It is also possible that relative air capillary and parabronchial volumes are smaller in these deeper-diving species than in the spheniscid penguin of the morphometry study. If penguins do inhale to this maximum air sac volume prior to their deepest dives, the magnitude and distribution of the body O-2 store would change considerably. In emperor penguins, total body O-2 would increase by 75%, and the respiratory fraction would increase from 33% to 61%. We emphasize that the maximum pre-dive respiratory air volume is still unknown in penguins. However, even lesser increases in air sac volume prior to a dive would still significantly increase the O-2 store. More refined evaluations of the respiratory O-2 store and baroprotective mechanisms in penguins await further investigation of species-specific lung morphometry, start-of-dive air volumes and body buoyancy, and the possibility of air exhalation during dives.

2012
Shiomi, K, Sato K, Ponganis PJ.  2012.  Point of no return in diving emperor penguins: is the timing of the decision to return limited by the number of strokes? Journal of Experimental Biology. 215:135-140.   10.1242/jeb.064568   AbstractWebsite

At some point in a dive, breath-hold divers must decide to return to the surface to breathe. The issue of when to end a dive has been discussed intensively in terms of foraging ecology and behavioral physiology, using dive duration as a temporal parameter. Inevitably, however, a time lag exists between the decision of animals to start returning to the surface and the end of the dive, especially in deep dives. In the present study, we examined the decision time in emperor penguins under two different conditions: during foraging trips at sea and during dives at an artificial isolated dive hole. It was found that there was an upper limit for the decision-to-return time irrespective of dive depth in birds diving at sea. However, in a large proportion of dives at the isolated dive hole, the decision-to-return time exceeded the upper limit at sea. This difference between the decision times in dives at sea versus the isolated dive hole was accounted for by a difference in stroke rate. The stroke rates were much lower in dives at the isolated hole and were inversely correlated with the upper limit of decision times in individual birds. Unlike the decision time to start returning, the cumulative number of strokes at the decision time fell within a similar range in the two experiments. This finding suggests that the number of strokes, but not elapsed time, constrained the decision of emperor penguins to return to the surface. While the decision to return and to end a dive may be determined by a variety of ecological, behavioral and physiological factors, the upper limit to that decision time may be related to cumulative muscle workload.

1999
Ponganis, PJ, Kooyman GL, Van Dam R, Lemaho Y.  1999.  Physiological responses of king penguins during simulated diving to 136 m depth. Journal of Experimental Biology. 202:2819-2822. AbstractWebsite

To evaluate blood N-2 uptake and the role of the respiratory volume (air sacs/lungs) as a N-2 and O-2 reservoir in deep-diving penguins, diving respiratory volume (V-DR), heart rate (f(H)), venous P-N2, blood volume (V-b) and hemoglobin (Hb) concentration were measured in king penguins (Aptenodytes patagonicus) during forced submersions and compressions equivalent to depths up to 136 m, V-DR was 69+/-18 ml kg(-1) (mean +/- S.D.) in 62 submersions ranging from 4.4 atmospheres absolute (ATA; 1 ATA=101 kPa) (34 m) to 14.6 ATA (136 m), Submersion f(H) averaged 30+/-7 beats min(-1) (N=18), approximately 20% of pre- and post-submersion values. Venous P-N2 values during and after submersions as deep as 11.2 ATA (102 m) were all less than 2.8 atmospheres N-2 (283 kPa) above ambient pressure, a previously measured threshold for symptomatic bubble formation. Mean V-b was 83+/-8 ml kg(-1) (N=6); [Hb] was 17.6+/-0.7 g dl(-1) (N=7), On a mass-specific basis, mean V-DR, and therefore total available N-2, is 41% of that in shallow-diving penguin species. Total body O-2 stores, calculated from measured V-DR, V-b, [Hb], muscle mass and myoglobin concentration, are 45 ml kg(-1), with 23 % in the respiratory system. This small respiratory fraction in comparison with that in shallow-diving penguins suggests a lesser reliance on the respiratory oxygen store for extended breath-holding and also a reduced uptake of nitrogen at depth.

1998
Kooyman, GL, Ponganis PJ.  1998.  The physiological basis of diving to depth: Birds and mammals. Annual Review of Physiology. 60:19-32.   10.1146/annurev.physiol.60.1.19   AbstractWebsite

There is wide diversity in the animals that dive to depth and in the distribution of their body oxygen stores. A hallmark of animals diving to depth is a substantial elevation of muscle myoglobin concentration. In deep divers, more than 80% of the oxygen store is in the blood and muscles. How these oxygen stores are managed, particularly within muscle, is unclear. The aerobic endurance of four species has now been measured. These measurements provide a standard for other species in which the limits cannot be measured. Diving to depth requires several adaptations to the effects of pressure. In mammals, one adaptation is lung collapse at shallow depths, which limits absorption of nitrogen. Blood Nz levels remain below the threshold for decompression sickness. No such adaptive model is known for birds. There appear to be two diving strategies used by animals that dive to depth. Seals, for example, seldom rely on anaerobic metabolism. Birds, on the other hand, frequently rely on anaerobic metabolism to exploit prey-rich depths otherwise unavailable to them.

1997
Ponganis, PJ, Kooyman GL, Starke LN, Kooyman CA, Kooyman TG.  1997.  Post-dive blood lactate concentrations in emperor penguins, Aptenodytes forsteri. Journal of Experimental Biology. 200:1623-1626. AbstractWebsite

In order to determine an aerobic diving limit (ADL) in emperor penguins (Aptenodytes forsteri), post-dive blood lactate concentrations were measured in penguins foraging at an isolated sea ice hole. Resting lactate concentrations were 1.2-2.7 mmol l(-1). Serial samples revealed that lactate level usually peaked within 5 min after dives and that 7-12 min was required for lactate concentrations to decrease from 5-8 mmol l(-1) to less than 2.5 mmol l(-1). Post-dive lactate level was not elevated above 3 mmol l(-1) for dives shorter than 5 min. Two-phase regression analysis revealed a transition at 5.6 min in the post-dive lactate level versus diving duration relationship. All dives longer than 7 min were associated with lactate concentrations greater than 5 mmol l(-1). We conclude that the ADL in emperor penguins ranges between 5 and 7 min. These are the first determinations of post-dive lactate concentrations in any free-diving bird and are currently the only physiological assessment of an ADL in an avian species.

1992
Ponganis, PJ, Kooyman GL, Sartoris D, Jobsis P.  1992.  Pinniped splenic volumes. American Journal of Physiology. 262:R322-R325. AbstractWebsite

Splenic volume was measured by computerized axial tomography in three harbor seals (Phoca vitulina) and two California sea lions (Zalophus californianus). Volumes ranged from 228 to 679 ml, representing 0.8-3.0% of calculated percentage body mass. Despite possible variation in the state of splenic contraction during the examination, these values are in the upper range of reported mammalian splenic volumes (as % of body mass). This reinforces the pinniped splenic erythrocyte storage concept.

Newsome, LR, Ponganis PJ, Reichman R, Nakaji N, Jaski B, Haltley M.  1992.  Portable percutaneous cardiopulmonary bypass: use in supported coronary angioplasty, aortic valvuloplasty, and cardiac arrest. Journal of Cardiothoracic and Vascular Anesthesia. 6:328-331.   10.1016/1053-0770(92)90151-V   Abstract

Portable cardiopulmonary bypass (CPB) systems consisting of a battery source and charger, centrifugal pump, hollow-fiber oxygenator, pump tubing, and large-bore thin-walled femoral arterial and venous cannulae have been commercially available for the past few years. Modifications of the Seldinger technique to allow percutaneous placement facilitate the expeditious institution of CPB in virtually any hospital setting.‘” As a result of this new technology, “supported” percutaneous transluminal coronary angioplasty (PTCA) and aortic valvuloplasty (AVP), which use the prophylactic institution of percutaneous CPB prior to the beginning of these procedures, were reported in 1990. Additionally, these portable systems have been used at various medical centers to help resuscitate patients suffering from cardiac arrest from a variety of causes. Two case histories representative of the use of such a system in both scenarios and including some anesthetic considerations for the use of such systems, specifically in supported angioplasties, are reported. In addition, the authors’ total experience with portable CPB is described.