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Kooyman, GL, Ponganis PJ, Castellini MA, Ponganis EP, Ponganis KV, Thorson PH, Eckert SA, Lemaho Y.  1992.  Heart rates and swim speeds of Emperor penguins diving under sea ice. Journal of Experimental Biology. 165:161-180. AbstractWebsite

Heart rate during overnight rest and while diving were recorded from five emperor penguins with a microprocessor-controlled submersible recorder. Heart rate, cardiac output and stroke volume were also measured in two resting emperor penguins using standard electrocardiography and thermodilution measurements. Swim velocities from eight birds were obtained with the submersible recorder. The resting average of the mean heart rates was 72 beats min-1. Diving heart rates were about 15% lower than resting rates. Cardiac outputs of 1.9-2.9 ml kg-1 s-1 and stroke volumes of 1.6-2.7 ml kg-1 were similar to values recorded from mammals of the same body mass. Swim velocities averaged 3 m s-1. The swim speeds and heart rates suggest that muscle O2 depletion must occur frequently: therefore, many dives require a significant energy contribution from anaerobic glycolysis.

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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.

Ponganis, PJ.  2007.  Bio-logging of physiological parameters in higher marine vertebrates. Deep-Sea Research Part Ii-Topical Studies in Oceanography. 54:183-192.   10.1016/j.dsr2.2006.11.009   AbstractWebsite

Bio-logging of physiological parameters in higher marine vertebrates had its origins in the field of bio-telemetry in the 1960s and 1970s. The development of microprocessor technology allowed its first application to bio-logging investigations of Weddell seal diving physiology in the early 1980s. Since that time, with the use of increased memory capacity, new sensor technology, and novel data processing techniques, investigators have examined heart rate, temperature, swim speed, stroke frequency, stomach function (gastric pH and motility), heat flux, muscle oxygenation, respiratory rate, diving air volume, and oxygen partial pressure (PO(2)) during diving. Swim speed, heart rate, and body temperature have been the most commonly studied parameters. Bio-logging investigation of pressure effects has only been conducted with the use of blood samplers and nitrogen analyses on animals diving at isolated dive holes. The advantages/disadvantages and limitations of recording techniques, probe placement, calibration techniques, and study conditions are reviewed. (c) 2007 Elsevier Ltd. All rights reserved.

Ponganis, PJ, Kooyman GL.  2000.  Diving physiology of birds: a history of studies on polar species. Comparative Biochemistry and Physiology a-Molecular and Integrative Physiology. 126:143-151.   10.1016/s1095-6433(00)00208-7   AbstractWebsite

Our knowledge of avian diving physiology has been based primarily on research with polar species. Since Scholander's 1940 monograph, research has expanded from examination of the 'diving reflex' to studies of free-diving birds, and has included laboratory investigations of oxygen stores, muscle adaptations, pressure effects, and cardiovascular/metabolic responses to swimming exercise. Behavioral and energetic studies at sea have shown that common diving durations of many avian species exceed the calculated aerobic diving limits (ADL). Current physiological research is focused on factors, such as heart rate and temperature, which potentially affect the diving metabolic rate and duration of aerobic diving. (C) 2000 Elsevier Science Inc. All rights reserved.

Ponganis, PJ, Van Dam RP, Levenson DH, Knower T, Ponganis KV, Marshall G.  2003.  Regional heterothermy and conservation of core temperature in emperor penguins diving under sea ice. Comparative Biochemistry and Physiology a-Molecular & Integrative Physiology. 135:477-487.   10.1016/s1095-6433(03)00133-8   AbstractWebsite

Temperatures were recorded at several body sites in emperor penguins (Aptenodytes forsteri) diving at an isolated dive hole in order to document temperature profiles during diving and to evaluate the role of hypothermia in this well-studied model of penguin diving physiology. Grand mean temperatures (+/-S.E.) in central body sites during dives were: stomach: 37.1 +/- 0.2 degreesC (n = 101 dives in five birds), pectoral muscle: 37.8 +/- 0.1 degreesC (n = 71 dives in three birds) and axillary/brachial veins: 37.9 +/- 0.1 degreesC (n = 97 dives in three birds). Mean diving temperature and duration correlated negatively at only one site in one bird (femoral vein, r = -0.59, P < 0.05; range < 1 degreesC). In contrast, grand mean temperatures in the wing vein, foot vein and lumbar subcutaneous tissue during dives were 7.6 +/- 0.7 degreesC (n = 157 dives in three birds), 20.2 +/- 1.2 degreesC (n = 69 in three birds) and 35.2 +/- 0.2 degreesC (n = 261 in six birds), respectively. Mean limb temperature during dives negatively correlated with diving duration in all six birds (r = -0.29 to -0.60, P < 0.05). In two of six birds, mean diving subcutaneous temperature negatively correlated with diving duration (r = -0.49 and -0.78, P < 0.05). Sub-feather temperatures decreased from 31 to 35 T during rest periods to a grand mean of 15.0 +/- 0.7 degreesC during 68 dives of three birds; mean diving temperature and duration correlated negatively in one bird (r = -0.42, P < 0.05). In general, pectoral, deep venous and even stomach temperatures during diving reflected previously measured vena caval temperatures of 37-39 degreesC more closely than the anterior abdominal temperatures (19-30 degreesC) recently recorded in diving emperors. Although prey ingestion can result in cooling in the stomach, these findings and the lack of negative correlations between internal temperatures and diving duration do not support a role for hypothermia-induced metabolic suppression of the abdominal organs as a mechanism of extension of aerobic dive time in emperor penguins diving at the isolated dive hole. Such high temperatures within the body and the observed decreases in limb, anterior abdomen, subcutaneous and sub-feather temperatures are consistent with preservation of core temperature and cooling of an outer body shell secondary to peripheral vasoconstriction, decreased insulation of the feather layer, and conductive/convective heat loss to the water environment during the diving of these emperor penguins. (C) 2003 Elsevier Science Inc. All fights reserved.

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Shiomi, K, Sato K, Mitamura H, Arai N, Naito Y, Ponganis PJ.  2008.  Effect of ocean current on the dead-reckoning estimation of 3-D dive paths of emperor penguins. Aquatic Biology. 3:265-270.   10.3354/ab00087   AbstractWebsite

The dead-reckoning technique is a useful method for obtaining 3-D movement data of aquatic animals. However, such positional data include an accumulative error. Understanding the source of the error is important for proper data interpretation. In order to determine whether ocean currents affect dive paths calculated by dead-reckoning, as has previously been hypothesized, we examined the directions of the estimated positions relative to the known real points (error direction) and the relationship between the error direction and the current direction. 3-D dive paths of emperor penguins Aptenodytes forsteri diving at isolated dive holes in eastern McMurdo Sound were reconstructed by dead-reckoning, and the net error and error direction were calculated. The net error correlated positively with the dive duration. The error directions were not distributed uniformly, and the mean error direction tended to be north of the starting point of dives. Because there was a southward-flowing current in eastern McMurdo Sound, the ocean current was likely to affect the calculated dive paths. Therefore, the method of error correction generally used, in which the net error divided by the dive duration is applied to each estimated position, is realistically appropriate, provided that the current does not change significantly during a dive.

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Wright, AK, Ponganis KV, McDonald BI, Ponganis PJ.  2014.  Heart rates of emperor penguins diving at sea: implications for oxygen store management. Marine Ecology Progress Series. 496:85-98.   10.3354/meps10592   AbstractWebsite

Heart rate (f(H)) contributes to control of blood oxygen (O-2) depletion through regulation of the magnitude of pulmonary gas exchange and of peripheral blood flow in diving vertebrates such as penguins. Therefore, we measured H during foraging trip dives of emperor penguins Aptenodytes forsteri equipped with digital electrocardiogram (ECG) recorders and time depth recorders (TDRs). Median dive f(H) (total heartbeats/duration, 64 beats min(-1)) was higher than resting H (56 beats min(-1)) and was negatively related to dive duration. Median dive f(H) in dives greater than the 5.6 min aerobic dive limit (ADL; dive duration associated with the onset of a net accumulation of lactic acid above resting levels) was significantly less than the median dive f(H) of dives less than the ADL (58 vs. 66 beats min(-1)). f(H) profile patterns differed between shallow (<50 m) and deep dives (>250 m), with values usually declining to levels near resting f(H) in shallow, short-duration dives, and to levels as low as 10 beats min(-1) during the deepest segments of deep dives. The total number of heartbeats in a dive was variable in shallow dives and consistently high in deep dives. A true bradycardia (f(H) below resting levels) during segments of 31% of shallow and deep dives of emperor penguins is consistent with reliance on myoglobin-bound O-2 stores for aerobic muscle metabolism that is especially accentuated during the severe bradycardias of deep dives. Although f(H) is low during the deepest segments of deep dives, the total number and distribution of heartbeats in deep, long dives suggest that pulmonary gas exchange and peripheral blood flow primarily occur at shallow depths.