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2018
Tift, MS, Huckstadt LA, Ponganis PJ.  2018.  Anterior vena caval oxygen profiles in a deep-diving California sea lion: arteriovenous shunts, a central venous oxygen store and oxygenation during lung collapse. Journal of Experimental Biology. 221   10.1242/jeb.163428   AbstractWebsite

Deep-diving California sea lions (Zalophus californianus) can maintain arterial hemoglobin saturation (S-O2) above 90% despite lung collapse (lack of gas exchange) and extremely low posterior vena caval S-O2 in the middle of the dive. We investigated anterior vena caval P-O2 and S-O2 during dives of an adult female sea lion to investigate two hypotheses: (1) posterior vena caval S-O2 is not representative of the entire venous oxygen store and (2) a well-oxygenated (arterialized) central venous oxygen reservoir might account for maintenance of arterial S-O2 during lung collapse. During deep dives, initial anterior vena caval S-O2 was elevated at 83.6 +/- 8.4% (n = 102), presumably owing to arteriovenous shunting. It remained high until the bottom phase of the dive and then decreased during ascent, whereas previously determined posterior vena caval S-O2 declined during descent and then often increased during ascent. These divergent patterns confirmed that posterior vena caval S-O2 was not representative of the entire venous oxygen store. Prior to and early during descent of deep dives, the high S-O2 values of both the anterior and posterior venae cavae may enhance arterialization of a central venous oxygen store. However, anterior vena caval S-O2 values at depths beyond lung collapse reached levels as low as 40%, making it unlikely that even a completely arterialized central venous oxygen store could account for maintenance of high arterial S-O2. These findings suggest that maintenance of high arterial S-O2 during deep dives is due to persistence of some gas exchange at depths beyond presumed lung collapse.

2017
Ponganis, PJ, McDonald BI, Tift MS, Williams CL.  2017.  Heart rate regulation in diving sea lions: the vagus nerve rules. Journal of Experimental Biology. 220:1372-1381.   10.1242/jeb.146779   AbstractWebsite

Recent publications have emphasized the potential generation of morbid cardiac arrhythmias secondary to autonomic conflict in diving marine mammals. Such conflict, as typified by cardiovascular responses to cold water immersion in humans, has been proposed to result from exercise-related activation of cardiac sympathetic fibers to increase heart rate, combined with depth-related changes in parasympathetic tone to decrease heart rate. After reviewing the marine mammal literature and evaluating heart rate profiles of diving California sea lions (Zalophus californianus), we present an alternative interpretation of heart rate regulation that de-emphasizes the concept of autonomic conflict and the risk of morbid arrhythmias in marine mammals. We hypothesize that: (1) both the sympathetic cardiac accelerator fibers and the peripheral sympathetic vasomotor fibers are activated during dives even without exercise, and their activities are elevated at the lowest heart rates in a dive when vasoconstriction is maximal, (2) in diving animals, parasympathetic cardiac tone via the vagus nerve dominates over sympathetic cardiac tone during all phases of the dive, thus producing the bradycardia, (3) adjustment in vagal activity, which may be affected by many inputs, including exercise, is the primary regulator of heart rate and heart rate fluctuations during diving, and (4) heart beat fluctuations (benign arrhythmias) are common in marine mammals. Consistent with the literature and with these hypotheses, we believe that the generation of morbid arrhythmias because of exercise or stress during dives is unlikely in marine mammals.

2013
McDonald, BI, Ponganis PJ.  2013.  Insights from venous oxygen profiles: oxygen utilization and management in diving California sea lions. Journal of Experimental Biology. 216:3332-3341.   10.1242/jeb.085985   AbstractWebsite

The management and depletion of O-2 stores underlie the aerobic dive capacities of marine mammals. The California sea lion (Zalophus californianus) presumably optimizes O-2 store management during all dives, but approaches its physiological limits during deep dives to greater than 300. m depth. Blood O-2 comprises the largest component of total body O-2 stores in adult sea lions. Therefore, we investigated venous blood O-2 depletion during dives of California sea lions during maternal foraging trips to sea by: (1) recording venous partial pressure of O-2 (PO2) profiles during dives, (2) characterizing the O-2-hemoglobin (Hb) dissociation curve of sea lion Hb and (3) converting the PO2 profiles into percent Hb saturation (SO2) profiles using the dissociation curve. The O-2-Hb dissociation curve was typical of other pinnipeds (P-50=28 +/- 2mmHg at pH 7.4). In 43% of dives, initial venous SO2 values were greater than 78% (estimated resting venous SO2), indicative of arterialization of venous blood. Blood O-2 was far from depleted during routine shallow dives, with minimum venous SO2 values routinely greater than 50%. However, in deep dives greater than 4. min in duration, venous SO2 reached minimum values below 5% prior to the end of the dive, but then increased during the last 30-60s of ascent. These deep dive profiles were consistent with transient venous blood O-2 depletion followed by partial restoration of venous O-2 through pulmonary gas exchange and peripheral blood flow during ascent. These differences in venous O-2 profiles between shallow and deep dives of sea lions reflect distinct strategies of O-2 store management and suggest that underlying cardiovascular responses will also differ.

2007
Sato, K, Watanuki Y, Takahashi A, Miller PJO, Tanaka H, Kawabe R, Ponganis PJ, Handrich Y, Akamatsu T, Watanabe Y, Mitani Y, Costa DP, Bost CA, Aoki K, Amano M, Trathan P, Shapiro A, Naito Y.  2007.  Stroke frequency, but not swimming speed, is related to body size in free-ranging seabirds, pinnipeds and cetaceans. Proceedings of the Royal Society B-Biological Sciences. 274:471-477.   10.1098/rspb.2006.0005   AbstractWebsite

It is obvious, at least qualitatively, that small animals move their locomotory apparatus faster than large animals: small insects move their wings invisibly fast, while large birds flap their wings slowly. However, quantitative observations have been difficult to obtain from free-ranging swimming animals. We surveyed the swimming behaviour of animals ranging from 0.5 kg seabirds to 30 000 kg sperm whales using animal-borne accelerometers. Dominant stroke cycle frequencies of swimming specialist seabirds and marine mammals were proportional to mass(-0.29) (R-2=0.99, n=17 groups), while propulsive swimming speeds of 1-2 m s(-1) were independent of body size. This scaling relationship, obtained from breath-hold divers expected to swim optimally to conserve oxygen, does not agree with recent theoretical predictions for optimal swimming. Seabirds that use their wings for both swimming and flying stroked at a lower frequency than other swimming specialists of the same size, suggesting a morphological trade-off with wing size and stroke frequency representing a compromise. In contrast, foot-propelled diving birds such as shags had similar stroke frequencies as other swimming specialists. These results suggest that muscle characteristics may constrain swimming during cruising travel, with convergence among diving specialists in the proportions and contraction rates of propulsive muscles.

Ponganis, PJ, Stockard TK, Meir JU, Williams CL, Ponganis KV, Van Dam RP, Howard R.  2007.  Returning on empty: extreme blood O-2 depletion underlies dive capacity of emperor penguins. Journal of Experimental Biology. 210:4279-4285.   10.1242/jeb.011221   AbstractWebsite

Blood gas analyses from emperor penguins (Aptenodytes forsteri) at rest, and intravascular P-O2 profiles from free-diving birds were obtained in order to examine hypoxemic tolerance and utilization of the blood O-2 store during dives. Analysis of blood samples from penguins at rest revealed arterial P(O2)s and O-2 contents of 68 +/- 7 mmHg (1 mmHg= 133.3 Pa) and 22.5 +/- 1.3 ml O-2 dl(-1) (N= 3) and venous values of 41 +/- 10 mmHg and 17.4 +/- 2.9 ml O-2 dl(-1) (N= 9). Corresponding arterial and venous Hb saturations for a hemoglobin (Hb) concentration of 18 g dl(-1) were > 91% and 70%, respectively. Analysis of P-O2 profiles obtained from birds equipped with intravascular P-O2 electrodes and backpack recorders during dives revealed that (1) the decline of the final blood P-O2 of a dive in relation to dive duration was variable, (2) final venous P-O2 values spanned a 40-mmHg range at the previously measured aerobic dive limit (ADL; dive duration associated with onset of post-dive blood lactate accumulation), (3) final arterial, venous and previously measured air sac P-O2 values were indistinguishable in longer dives, and (4) final venous P-O2 values of longer dives were as low as 1-6 mmHg during dives. Although blood O-2 is not depleted at the ADL, nearly complete depletion of the blood O-2 store occurs in longer dives. This extreme hypoxemic tolerance, which would be catastrophic in many birds and mammals, necessitates biochemical and molecular adaptations, including a shift in the O-2-Hb dissociation curve of the emperor penguin in comparison to those of most birds. A relatively higher-affinity Hb is consistent with blood P-O2 values and O-2 contents of penguins at rest.

2005
Knower Stockard, T, Heil J, Meir JU, Sato K, Ponganis KV, Ponganis PJ.  2005.  Air sac P-O2 and oxygen depletion during dives of emperor penguins. Journal of Experimental Biology. 208:2973-2980.   10.1242/jeb.01687   AbstractWebsite

In order to determine the rate and magnitude of respiratory O-2 depletion during dives of emperor penguins (Aptenodytes forsteri), air sac O-2 partial pressure (PO2) was recorded in 73 dives of four birds at an isolated dive hole. These results were evaluated with respect to hypoxic tolerance, the aerobic dive limit (ADL; dive duration beyond which there is post-dive lactate accumulation) and previously measured field metabolic rates (FMRs). 55% of dives were greater in duration than the previously measured 5.6-min ADL. P-O2 and depth profiles revealed compression hyperoxia and gradual O-2 depletion during dives. 42% of final P(O2)s during the dives (recorded during the last 15 s of ascent) were < 20 mmHg (< 2.7 kPa). Assuming that the measured air sac P-O2 is representative of the entire respiratory system, this implies remarkable hypoxic tolerance in emperors. In dives of durations greater than the ADL, the calculated end-of-dive air sac O-2 fraction was < 4%. The respiratory O-2 store depletion rate of an entire dive, based on the change in O-2 fraction during a dive and previously measured diving respiratory volume, ranged from I to 5 ml O-2 kg(-1) min(-1) and decreased exponentially with diving duration. The mean value, 2.1 +/- 0.8 ml O-2 kg(-1) min(-1), was (1) 19-42% of previously measured respiratory O-2 depletion rates during forced submersions and simulated dives, (2) approximately one-third of the predicted total body resting metabolic rate and (3) approximately 10% of the measured FMR. These findings are consistent with a low total body metabolic rate during the dive.

2002
Van Dam, RP, Ponganis PJ, Ponganis KV, Levenson DH, Marshall G.  2002.  Stroke frequencies of emperor penguins diving under sea ice. Journal of Experimental Biology. 205:3769-3774. AbstractWebsite

During diving, intermittent swim stroke patterns, ranging from burst/coast locomotion to prolonged gliding, represent potential energy conservation mechanisms that could extend the duration of aerobic metabolism and, hence, increase the aerobic dive limit (ADL, dive duration associated with onset of lactate accumulation). A 5.6 min ADL for emperor penguins had been previously determined with lactate measurements after dives of <50 m depth. In order to assess locomotory patterns during such dives, longitudinal acceleration was measured with an attached accelerometer in 44 dives of seven adult birds diving from an isolated dive hole in the sea ice of McMurdo Sound, Antarctica. Detection of wing strokes in processed accelerometer data was verified in selected birds with analysis of simultaneous Crittercam underwater video footage. Mean dive duration of birds equipped with the accelerometer and a time-depth recorder (TDR) was 5.7+/-2.2 min; 48% of these dives were greater than the measured 5.6 min ADL (ADL(M)). Highest stroke frequencies (0.92+/-0.31Hz, N=981) occurred during the initial descent to 12 m depth. Swimming effort was reduced to a mean stroke frequency <0.70 Hz during other phases of the dive (while traveling below 12 m depth, during foraging ascents/descents to and from the sub-ice surface, and during final ascents to exit). The longest stroke interval (8.6 s) occurred during a feeding excursion to the undersurface of the ice. In dives >ADL(M), mean stroke frequency during travel segments was significantly less than that in dives 10 s) periods of prolonged gliding during these shallow (<60 m) foraging dives. However, a stroke/glide pattern was evident with more than 50% of strokes associated with a stroke interval >1.6 s, and with lower stroke frequency associated with increased dive duration.

2000
Ponganis, PJ, Van Dam RP, Marshall G, Knower T, Levenson DH.  2000.  Sub-ice foraging behavior of emperor penguins. Journal of Experimental Biology. 203:3275-3278. AbstractWebsite

Emperor penguins (Aptenodytes forsteri) were equipped with a remote underwater video camera, the Crittercam, to evaluate sub-ice foraging behavior while the birds dived from an isolated dive hole. Three birds dived and foraged successfully for Ih periods after being trained to wear and to dive with a harness for camera attachment. Video and depth profile recordings revealed that emperor penguins travel at shallow depths (<50 m), ascend to the undersurface of the ice to feed on fish, and descend back to depth to return to the exit hole. Although the mean durations of dives of individual birds with the Crittercam were 21-35 % shorter than the diving durations of these same birds without the camera, the dive profiles in both situations were similar, thus demonstrating a similar foraging strategy in birds diving without the camera. Despite shorter diving durations with the camera, the penguins were still successful at prey capture in 80 % of 91 dives greater than 1 min in duration. Prey included the sub-ice fish Pagothenia borchgrevinki. Hunting ascents (from depth to within 5 m of the surface) occurred in 85 % of dives, ranged from zero to three per dive, and were associated with successful prey capture in 77 % of 128 ascents, Occasionally, several fish were captured during a single ascent, These observations and this application of video technology create a model for further physiological and behavioral studies of foraging, and also emphasize the potential importance of shallow dives as sources of food intake for emperor penguins during foraging trips to sea.

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.

1997
Ponganis, PJ, Kooyman GL, Baranov EA, Thorson PH, Stewart BS.  1997.  The aerobic submersion limit of Baikal seals, Phoca sibirica. Canadian Journal of Zoology-Revue Canadienne De Zoologie. 75:1323-1327.   10.1139/z97-756   AbstractWebsite

An aerobic dive limit (ADL), the diving duration beyond which postdive lactate concentration increases above the resting level, has been estimated theoretically for many species. Such calculations have been based on an oxygen store/diving metabolic rate (MR) equation. Until now, an ADL has been determined empirically from measurements of blood lactate concentration only in the Weddell seal, Leptonychotes weddellii. We measured post-submergence plasma lactate concentrations during spontaneous voluntary submersions of three captive adult Baikal seals (Phoca sibirica). Two-phase regression analysis revealed a transition in the lactate concentration - submersion duration relationship after the animal had been diving for 15 min. Data collected in prior studies on oxygen stores and submersion metabolic rates of Baikal seals yielded a calculated aerobic limit of 16 min. As in Weddell seals, the empirically determined aerobic limit was very similar to the theoretical limit. Furthermore, most diving durations recorded during recent studies of free-ranging Baikal seals are under this limit. These data support the concept of an ADL and its estimation by means of an oxygen store/diving MR calculation.