Publications

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1992
Castellini, MA, Kooyman GL, Ponganis PJ.  1992.  Metabolic rates of freely diving Weddell seals: correlations with oxygen stores, swim velocity and diving duration. Journal of Experimental Biology. 165:181-194. AbstractWebsite

The metabolic rates of freely diving Weddell seals were measured using modern methods of on-line computer analysis coupled to oxygen consumption instrumentation. Oxygen consumption values were collected during sleep, resting periods while awake and during diving periods with the seals breathing at the surface of the water in an experimental sea-ice hole in Antarctica. Oxygen consumption during diving was not elevated over resting values but was statistically about 1.5 times greater than sleeping values. The metabolic rate of diving declined with increasing dive duration, but there was no significant difference between resting rates and rates in dives lasting up to 82 min. Swimming speed, measured with a microprocessor velocity recorder, was constant in each animal. Calculations of the aerobic dive limit of these seals were made from the oxygen consumption values and demonstrated that most dives were within this theoretical limit. The results indicate that the cost of diving is remarkably low in Weddell seals relative to other diving mammals and birds.

2008
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.

2009
Meir, JU, Champagne CD, Costa DP, Williams CL, Ponganis PJ.  2009.  Extreme hypoxemic tolerance and blood oxygen depletion in diving elephant seals. American Journal of Physiology-Regulatory Integrative and Comparative Physiology. 297:R927-R939.   10.1152/ajpregu.00247.2009   AbstractWebsite

Meir JU, Champagne CD, Costa DP, Williams CL, Ponganis PJ. Extreme hypoxemic tolerance and blood oxygen depletion in diving elephant seals. Am J Physiol Regul Integr Comp Physiol 297: R927-R939, 2009. First published July 29, 2009; doi: 10.1152/ajpregu.00247.2009.-Species that maintain aerobic metabolism when the oxygen (O(2)) supply is limited represent ideal models to examine the mechanisms underlying tolerance to hypoxia. The repetitive, long dives of northern elephant seals (Mirounga angustirostris) have remained a physiological enigma as O(2) stores appear inadequate to maintain aerobic metabolism. We evaluated hypoxemic tolerance and blood O(2) depletion by 1) measuring arterial and venous O(2) partial pressure (PO(2)) during dives with a PO(2)/temperature recorder on elephant seals, 2) characterizing the O(2) hemoglobin (O(2)-Hb) dissociation curve of this species, 3) applying the dissociation curve to PO(2) profiles to obtain %Hb saturation (SO(2)), and 4) calculating blood O(2) store depletion during diving. Optimization of O(2) stores was achieved by high venous O(2) loading and almost complete depletion of blood O(2) stores during dives, with net O(2) content depletion values up to 91% (arterial) and 100% (venous). In routine dives (>10 min) Pv(O2) and Pa(O2) values reached 2-10 and 12-23 mmHg, respectively. This corresponds to SO(2) of 1-26% and O(2) contents of 0.3 (venous) and 2.7 ml O(2)/dl blood (arterial), demonstrating remarkable hypoxemic tolerance as PaO(2) is nearly equivalent to the arterial hypoxemic threshold of seals. The contribution of the blood O(2) store alone to metabolic rate was nearly equivalent to resting metabolic rate, and mean temperature remained near 37 degrees C. These data suggest that elephant seals routinely tolerate extreme hypoxemia during dives to completely utilize the blood O(2) store and maximize aerobic dive duration.

2010
Shiomi, K, Narazaki T, Sato K, Shimatani K, Arai N, Ponganis PJ, Miyazaki N.  2010.  Data-processing artefacts in three-dimensional dive path reconstruction from geomagnetic and acceleration data. Aquatic Biology. 8:299-304.   10.3354/ab00239   AbstractWebsite

Tri-axis magnetism and acceleration data loggers have recently been used to obtain time-series headings and, consequently, the 3-dimensional dive paths of aquatic animals. However, problems may arise in the resulting calculation process with multiple parameters. In this study, the dive paths of loggerhead turtles and emperor penguins were reconstructed. For both species, apparently unrealistic movements were found. Time-series heading data of turtles showed small regular fluctuations synchronous with stroking. In the dive paths of penguins, infrequent abrupt changes in heading were observed during stroke cycles. These were unlikely to represent true behaviours according to observations of underwater behaviour and tri-axis magnetism and acceleration data. Based on the relationship between sampling frequency and frequency of body posture change, we suggest that (1) the changes in the animals' posture concurrent with strokes and (2) the mismatched treatment (i.e. filtering and non-filtering) of the acceleration and magnetism data caused the artefacts. These inferences are supported by the results of simulations. For data sets obtained at a given sampling frequency, the error pattern in calculated dive paths is likely to differ depending on the frequency and amplitude of body posture changes and in swim speed. In order to avoid misinterpretation, it is necessary to understand the assumptions and inherent problems of the calculation methods as well as the behavioural characteristics of the study animals.

Houser, DS, Dankiewicz-Talmadge LA, Stockard TK, Ponganis PJ.  2010.  Investigation of the potential for vascular bubble formation in a repetitively diving dolphin. Journal of Experimental Biology. 213:52-62.   10.1242/jeb.028365   AbstractWebsite

The production of venous gas emboli (VGE) resulting from altered dive behavior is postulated as contributing to the stranding of beaked whales exposed to mid-frequency active sonar. To test whether nitrogen gas uptake during repetitive breath-hold diving is sufficient for asymptomatic VGE formation in odontocetes, a bottlenose dolphin (Tursiops truncatus Montagu) was trained to perform 10-12 serial dives with 60s surface intervals to depths of 30, 50, 70 or 100m. The dolphin remained at the bottom depth for 90s on each dive. Doppler and/or two-dimensional imaging ultrasound did not detect VGE in the portal and brachiocephalic veins following a dive series. Van Slyke analyses of serial, post-dive blood samples drawn from the fluke yielded blood nitrogen partial pressure (P(N2)) values that were negligibly different from control samples. Mean heart rate (HR; +/-1. s.d.) recorded during diving was 50+/-3. beats min(-1) and was not significantly different between the 50, 70 and 100 m dive sessions. The absence of VGE and elevated blood P(N2) during post-dive periods do not support the hypothesis that N(2) supersaturation during repetitive dives contributes to VGE formation in the dolphin. The diving HR pattern and the presumed rapid N(2) washout during the surface-interval tachycardia probably minimized N(2) accumulation in the blood during dive sessions.

2011
Ponganis, PJ, Meir JU, Williams CL.  2011.  In pursuit of Irving and Scholander: a review of oxygen store management in seals and penguins. Journal of Experimental Biology. 214:3325-3339.   10.1242/jeb.031252   AbstractWebsite

Since the introduction of the aerobic dive limit (ADL) 30 years ago, the concept that most dives of marine mammals and sea birds are aerobic in nature has dominated the interpretation of their diving behavior and foraging ecology. Although there have been many measurements of body oxygen stores, there have been few investigations of the actual depletion of those stores during dives. Yet, it is the pattern, rate and magnitude of depletion of O(2) stores that underlie the ADL. Therefore, in order to assess strategies of O(2) store management, we review (a) the magnitude of O(2) stores, (b) past studies of O(2) store depletion and (c) our recent investigations of O(2) store utilization during sleep apnea and dives of elephant seals (Mirounga angustirostris) and during dives of emperor penguins (Aptenodytes forsteri). We conclude with the implications of these findings for (a) the physiological responses underlying O(2) store utilization, (b) the physiological basis of the ADL and (c) the value of extreme hypoxemic tolerance and the significance of the avoidance of re-perfusion injury in these animals.

2017
Tift, MS, Huckstadt LA, McDonald BI, Thorson PH, Ponganis PJ.  2017.  Flipper stroke rate and venous oxygen levels in free-ranging California sea lions. Journal of Experimental Biology. 220:1533-1540.   10.1242/jeb.152314   AbstractWebsite

The depletion rate of the blood oxygen store, development of hypoxemia and dive capacity are dependent on the distribution and rate of blood oxygen delivery to tissues while diving. Although blood oxygen extraction by working muscle would increase the blood oxygen depletion rate in a swimming animal, there is little information on the relationship between muscle workload and blood oxygen depletion during dives. Therefore, we examined flipper stroke rate, a proxy of muscle workload, and posterior vena cava oxygen profiles in four adult female California sea lions (Zalophus californianus) during foraging trips at sea. Flipper stroke rate analysis revealed that sea lions minimized muscle metabolism with a stroke-glide strategy when diving, and exhibited prolonged glides during the descent of deeper dives (>100 m). During the descent phase of these deep dives, 55 +/- 21% of descent was spent gliding, with the longest glides lasting over 160 s and covering a vertical distance of 340 m. Animals also consistently glided to the surface from 15 to 25 m depth during these deeper dives. Venous hemoglobin saturation (SO2) profiles were highly variable throughout dives, with values occasionally increasing during shallow dives. The relationship between SO2 and flipper stroke rate was weak during deeper dives, while this relationship was stronger during shallow dives. We conclude that (1) the depletion of oxygen in the posterior vena cava in deep-diving sea lions is not dependent on stroke effort, and (2) stroke-glide patterns during dives contribute to a reduction of muscle metabolic rate.