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Ancel, A, Starke LN, Ponganis PJ, Van Dam R, Kooyman GL.  2000.  Energetics of surface swimming in Brandt's cormorants (Phalacrocorax penicillatus Brandt). Journal of Experimental Biology. 203:3727-3731. AbstractWebsite

The energy requirements of Brandt's cormorants (Phalacrocorax penicillatus) during surface swimming were measured in birds swimming under a metabolic chamber in a water flume. From the oxygen consumption recordings, we extrapolated the metabolic rate and cost of transport at water speeds ranging from 0 to 1.3 ms(-1). In still water, the birds' mean mass-specific rate of oxygen consumption ((V)over dot(O2),) while floating at the surface was 20.2ml O-2 min(-1) kg(-1), 2.1 times the predicted resting metabolic rate. During steady-state voluntary swimming against a how, their Po, increased with water speed, reaching 74 mi O-2 min(-1) kg(-1) at 1.3 ms(-1), which corresponded to an increase in metabolic rate from 11 to 25 W kg(-1). The cost of transport decreased,vith swimming velocity, approaching a minimum of 19 J kg(-1) m(-1) for a swimming speed of 1.3 m s(-1) Surface swimming in the cormorant costs approximately 18% less than sub-surface swimming. This confirms similar findings in tufted ducks (Aythya fuligula) and supports the hypothesis that increased energy requirements are necessary in these bird diving to overcome buoyancy and heat submergence.

Ancel, A, Kooyman GL, Ponganis PJ, Gendner JP, Lignon J, Mestre X, Huin N, Thorson PH, Robisson P, Lemaho Y.  1992.  Foraging behaviour of emperor penguins as a resource detector in winter and summer. Nature. 360:336-339.   10.1038/360336a0   AbstractWebsite

The emperor penguin (Aptenodytes forsteri), which feeds only at sea, is restricted to the higher latitudes of the antarctic sea-ice habitat1-3. It breeds on the winter fast ice when temperatures are -30-degrees-C and high winds are frequent3. Assuming entirely the task of incubating the single egg, the male fasts for about 120 days in the most severe conditions. When it is relieved by the female around hatching time, the distance between the colony and the open sea may be 100 km or more4,5, but where emperors go to forage at that time or during the summer is unknown. The polynias are areas of open water in sea-ice and during winter, with the under-ice habitats at any time of the year, they are among the most difficult of all Antarctic areas to sample. Here we monitor by satellite the routes taken by emperor penguins for foraging and compare them with satellite images of sea-ice. Winter birds walking over fast ice travelled up to 296 km to feed in polynias, whereas those swimming in light pack-ice travelled as far as 895 km from the breeding colony. One record of diving showed that although most dives are to mid-water depths, some are near the bottom. Obtaining such detailed information on foraging in emperor penguins means that this bird now offers a unique opportunity to investigate the Antarctic sea-ice habitat.

Barber-Meyer, SM, Kooyman GL, Ponganis PJ.  2008.  Trends in western Ross Sea emperor penguin chick abundances and their relationships to climate. Antarctic Science. 20:3-11.   10.1017/s0954102007000673   AbstractWebsite

The emperor penguin (Aptenodytes forsteri) is extremely dependent on the extent and stability of sea ice, which may make the species particularly susceptible to environmental change. In order to appraise the stability of the emperor penguin populations at six colonies in the western Ross Sea, we used linear regression analysis to evaluate chick abundance trends (1983-2005) and Pearson's r correlation to assess their relation to two local and two large-scale climate variables. We detected only one significant abundance trend; the Cape Roget colony increased from 1983 to 1996 (n = 6). Higher coefficients of variation in chick abundances at smaller colonies (Cape Crozier, Beaufort Island, Franklin Island) suggest that such colonies occupy marginal habitat, and are more susceptible to environmental change. We determined chick abundance to be most often correlated with local Ross Sea climate variables (sea ice extent and sea surface temperature), but not in consistent patterns across the colonies. We propose that chick abundance is most impacted by fine scale sea ice extent and local weather events, which are best evaluated by on-site assessments. We did not find sufficient evidence to reject the hypothesis that the overall emperor penguin population in the Ross Sea was stable during this period.

Barber-Meyer, SM, Kooyman GL, Ponganis PJ.  2007.  Estimating the relative abundance of emperor penguins at inaccessible colonies using satellite imagery. Polar Biology. 30:1565-1570.   10.1007/s00300-007-0317-8   AbstractWebsite

Emperor penguin (Aptenodytes forsteri) populations are useful environmental indicators due to the bird's extreme reliance on sea ice. We used remote sensing technology to estimate relative adult bird abundance at two inaccessible emperor penguin colonies in the Ross Sea, Antarctica. We performed supervised classification of 12 panchromatic satellite images of the seven known Ross Sea colonies. We used regression to predict adult bird counts at the inaccessible colonies by relating the number of pixels classified as "penguin" in the satellite images of the accessible colonies to corresponding known adult bird counts from aerial photographs or ground counts. While our analysis was hampered by excessive guano and shadows, we used satellite imagery to differentiate between relatively small (< 3,000 adult birds) and larger colonies (> 5,000 adult birds). Remote sensing technology is logistically less intense and less costly than aerial or ground censuses when the objective is to document penguin presence and/or large emperor penguin population changes (e.g., catastrophic changes). Improvements expected soon in the resolution of the satellite images should allow for more accurate abundance estimates.

Blight, LK, Ainley DG, Ackley SF, Ballard G, Ballerini T, Brownell RL, Cheng CHC, Chiantore M, Costa D, Coulter MC, Dayton P, Devries AL, Dunbar R, Earle S, Eastman JT, Emslie SD, Evans CW, Garrott RA, Kim S, Kooyman G, Lescroel A, Lizotte M, Massaro M, Olmastroni S, Ponganis PJ, Russell J, Siniff DB, Smith WO, Stewart BS, Stirling I, Willis J, Wilson P, Woehler EJ.  2010.  Fishing for data in the Ross Sea. Science. 330:1316-1316.   10.1126/science.330.6009.1316   AbstractWebsite
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.

Cristofari, R, Bertorelle G, Ancel A, Benazzo A, Lemaho Y, Ponganis PJ, Stenseth NC, Trathan PN, Whittington JD, Zanetti E, Zitterbart DP, Le Bohec C, Trucchi E.  2016.  Full circumpolar migration ensures evolutionary unity in the Emperor penguin. Nature Communications. 7   10.1038/ncomms11842   AbstractWebsite

Defining reliable demographic models is essential to understand the threats of ongoing environmental change. Yet, in the most remote and threatened areas, models are often based on the survey of a single population, assuming stationarity and independence in population responses. This is the case for the Emperor penguin Aptenodytes forsteri, a flagship Antarctic species that may be at high risk continent-wide before 2100. Here, using genome-wide data from the whole Antarctic continent, we reveal that this top-predator is organized as one single global population with a shared demography since the late Quaternary. We refute the view of the local population as a relevant demographic unit, and highlight that (i) robust extinction risk estimations are only possible by including dispersal rates and (ii) colony-scaled population size is rather indicative of local stochastic events, whereas the species' response to global environmental change is likely to follow a shared evolutionary trajectory.

Crognale, MA, Levenson DH, Ponganis PJ, Deegan JF, Jacobs GH.  1998.  Cone spectral sensitivity in the harbor seal (Phoca vitulina) and implications for color vision. Canadian Journal of Zoology-Revue Canadienne De Zoologie. 76:2114-2118.   10.1139/cjz-76-11-2114   AbstractWebsite

The retinas of harbor seals (Phoca vitulina) contain two morphologically distinct photoreceptor types: rods and cones. The spectral properties of the cones have not been previously studied. The spectral sensitivities of the cones of harbor seals were measured using a retinal gross potential technique, flicker photometric electroretinography. We found a cone spectral sensitivity curve with a peak at about 510 nm. The shape of the spectral sensitivity curve remained invariant despite large changes in chromatic adaptation, implying that harbor seals have only a single cone photopigment. This means that harbor seals must lack color vision at photopic light levels. Any color discrimination in this species would have to be based on combined input from rods and cones and thus restricted to mesopic light levels. The spectral sensitivity of the cone pigment in the harbor seal is shifted to shorter wavelengths than those of terrestrial carnivores, consistent with adaptation to the aquatic photic environment.

Dolar, MLL, Suarez P, Ponganis PJ, Kooyman GL.  1999.  Myoglobin in pelagic small cetaceans. Journal of Experimental Biology. 202:227-236. AbstractWebsite

Although myoglobin (Mb) is considered to contribute significantly to the oxygen and diving capacity of marine mammals, few data are available for cetaceans, Cetacean by-catch in the tuna driftnet fisheries in the Sulu Sea, Philippines, afforded the opportunity to examine Mb content and distribution, and to determine muscle mass composition, in Fraser's (Lagenodelphis hosei) and spinner (Stenella longirostris) dolphins and a pygmy killer whale (Feresa attenuata). Age was estimated by body length determination. Stomach contents were analyzed for the presence or absence of milk and solid foods. It was hypothesized (a) that Mb concentration ([Mb]) would be higher in Fraser's and spinner dolphins than in other small cetaceans because of the known mesopelagic distribution of their prey, (b) that [Mb] would vary among different muscles according to function during diving, and (c) that [Mb] would increase with age during development. The results were as follows. ii) Myoglobin concentrations of the longissimus muscle in adult Fraser's (6.8-7.2 g 100 g(-1) muscle) and spinner (5-6 g 100 g(-1) muscle) dolphins and in an immature pygmy killer whale (5.7 g 100 g(-1) muscle) were higher than those reported previously for small cetaceans, (2) [Mb] varied significantly among the different muscle types in adult dolphins but not in calves; in adults, swimming muscles had significantly higher [Mb] than did non-swimming muscles, contained 82-86 % of total Mb, and constituted 75-80 % of total muscle mass. (3) Myoglobin concentrations in Fraser's and spinner dolphins increased with size and age and were 3-4 times greater in adults than in calves, The high Mb concentrations measured in the primary locomotory muscles of these pelagic dolphins are consistent with the known mesopelagic foraging behaviour of Fraser's and spinner dolphins and suggest that the pygmy killer whale is also a deep-diving species. The high Mb concentrations in epaxial, hypaxial and abdominal muscle groups also support the primary locomotory functions suggested for these muscles in other anatomical studies. As in other species. the increase in [Mb] during development probably parallels the development of diving capacity.

Eckert, SA, Eckert KL, Ponganis P, Kooyman GL.  1989.  Diving and foraging behavior of leatherback sea turtles (Dermochelys coriacea). Canadian Journal of Zoology-Revue Canadienne De Zoologie. 67:2834-2840.   10.1139/z89-399   AbstractWebsite

Remote time–depth recorders (TDR) were deployed on six gravid leatherbacks nesting on Sandy Point, St. Croix. Dive behavior was monitored continuously for each turtle during internesting intervals ranging from 9 to 11 days. Dive duration averaged 9.9 min/dive (SD = 5.3, n = 5096); mean depth was 61.6 m (SD = 59.1, n = 5096). One turtle dived twice beyond the range of her TDR to depths we estimate >1000 m. Postdive surfacing intervals averaged 4.9 min/dive (SD = 13.1, n = 5090). Differences in mean dive depth, dive duration, and surface intervals among turtles were not attributable to differences in body size (length or mass). Distinct diel periodicity was observed in dive behavior; submergence intervals were longest at dawn, declined throughout the day, and were shortest at dusk. Night dives (19:00–04:59) were shorter, shallower, and more frequent than day dives (05:00–18:59). Dive depth was less variable at night than during the day. The dive pattern suggests nocturnal foraging within the deep scattering layer, a hypothesis that is corroborated by seasonal weight loss data.

Goforth, H, Ponganis PJ, Eggerton E.  1987.  Glycogenolytic responses and force production characteristics of a bottlenose dolphin (Tursiops truncatus). Seventh biennial Conference on the Biology of Marine Mammals, abstracts, December 5-9, 1987, Miami, Florida. :1., Miami, FL: s.n. Abstract
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.

Jobsis, PD, Ponganis PJ, Kooyman GL.  2001.  Effects of training on forced submersion responses in harbor seals. Journal of Experimental Biology. 204:3877-3885. AbstractWebsite

In several pinniped species, the heart rates observed during unrestrained dives are frequently higher than the severe bradycardias recorded during forced submersions. To examine other physiological components of the classic 'dive response' during such moderate bradycardias, a training protocol was developed to habituate harbor seals (Phoca vitulina) to short forced submersions. Significant changes were observed between physiological measurements made during naive and trained submersions (3-3.5min). Differences were found in measurements of heart rate during submersion (naive 18 +/-4.3 beats min(-1) versus trained 35 +/-3.4 beats min(-1)), muscle blood flow measured using laser-Doppler flowmetry (naive 1.8 +/-0.8 ml min(-1) 100 g(-1) versus trained 5.8 +/-3.9 ml min(-1) 100 g(-1)), change in venous P-O 2 (naive -0.44 +/-1.25 kPa versus trained -1.48 +/-0.76 kPa) and muscle deoxygenation rate (naive -0.67 +/-0.27 mvd s(-1) versus trained -0.51 +/-0.18 mvd s(-1), a relative measure of muscle oxygenation provided by the Vander Niroscope, where mvd are milli-vander units). In contrast to the naive situation, the post-submersion increase in plasma lactate levels was only rarely significant in trained seals. Resting eupneic (while breathing) heart rate and total oxygen consumption rates (measured in two seals) were not significantly different between the naive and trained states. This training protocol revealed that the higher heart rate and greater muscle blood flow in the trained seals were associated with a lower muscle deoxygenation rate, presumably secondary to greater extraction of blood O-2 during trained submersions. Supplementation of muscle oxygenation by blood O-2 delivery during diving would increase the rate of blood O-2 depletion but could prolong the duration of aerobic muscle metabolism during diving. This alteration of the dive response may increase the metabolic efficiency of diving.

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.

Kooyman, GL, Ponganis PJ.  2014.  Chick production at the largest emperor penguin colony decreases by 50% from 2008-10. Antarctic Science. 26:33-37.   10.1017/s0954102013000515   AbstractWebsite

The emperor penguin colony at Coulman Island is reputedly the largest known. This reputation is based on intermittent ground and aerial surveys performed since 1958. From an aerial survey obtained on 28 October 2010 we discovered that the total number of chicks was 56% of the lowest previous estimate of 2006 and only 41% of the most recent estimate in 2008. All of the counts tallied since 1983 were determined either by ground counts or from aerial film or digital photographs, or estimates from adult counts. We also determined the sea ice conditions in autumn, which is close to the time the adults arrive to breed. We present three hypotheses of what might have happened from 2008-10 to cause the step change in chick production, the small recovery of chick numbers in 2011, and the complete recovery of number of adults from 2010-11. We conclude that local circumstances may have strongly influenced the breeding behaviour of the emperor penguins in 2010 and to a lesser degree in 2011 when many adults elected not to breed.

Kooyman, GL, Ponganis PJ.  2017.  Rise and fall of Ross Sea emperor penguin colony populations: 2000 to 2012. Antarctic Science. 29:201-208.   10.1017/s0954102016000559   AbstractWebsite

There are seven emperor penguin (Aptenodytes forsteri) colonies distributed throughout the traditional boundaries of the Ross Sea from Cape Roget to Cape Colbeck. This coastline is c. 10% of the entire coast of Antarctica. From 2000 to 2012, there has been a nearly continuous record of population size of most, and sometimes all, of these colonies. Data were obtained by analysing aerial photographs. We found large annual variations in populations of individual colonies, and conclude that a trend from a single emperor penguin colony may not be a good environmental sentinel. There are at least four possibilities for census count fluctuations: i) this species is not bound to a nesting site like other penguins, and birds move within the colony and possibly to other colonies, ii) harsh environmental conditions cause a die-off of chicks in the colony or of adults elsewhere, iii) the adults skip a year of breeding if pre-breeding foraging is inadequate and iv) if sea ice conditions are unsatisfactory at autumn arrival of the adults, they skip breeding or go elsewhere. Such variability indicates that birds at all Ross Sea colonies should be counted annually if there is to be any possibility of understanding the causes of population changes.

Kooyman, GL, Ponganis PJ, Howard RS.  1999.  Diving Animals. The lung at depth. ( Lundgren CEG, Miller JN, Eds.).:587-620., New York: Marcel Dekker Abstract
Kooyman, GL, Ponganis PJ.  1994.  Emperor penguin oxygen consumption, heart rate and plasma lactate levels during graded swimming exercise. Journal of Experimental Biology. 195:199-209. AbstractWebsite

Oxygen consumption (V-O2), heart rate and blood chemistry were measured in four emperor penguins, Aptenodytes forsteri (Gray), during graded swimming exercise. The maximum V-O2, obtained, 52ml O-2 kg(-1) min(-1), was 7.8 times the measured resting V-O2 of 6.7 ml O-2 kg(-1) min(-1) and 9.1 times the predicted resting V-O2. As the swimming effort rose, a linear increase in surface and submerged heart rates (fH) occurred. The highest average maximum surface and submersion heart rates of any bird were 213 and 210 beats min(-1), respectively. No increase in plasma lactate concentrations occurred until V-O2 was greater than 25 ml O-2 kg(-1) min(-1). At the highest V-O2 values measured, plasma lactate concentration reached 9.4 mmol l(-1). In comparison with other animals of approximately the same mass, the aerobic capacity of the emperor penguin is less than those of the emu and dog but about the same as those of the seal, sea lion and domestic goat. For aquatic animals, a low aerobic capacity seems to be consistent with the needs of parsimonious oxygen utilization while breath-holding.

Kooyman, GL, Ainley DG, Ballard G, Ponganis PJ.  2007.  Effects of giant icebergs on two emperor penguin colonies in the Ross Sea, Antarctica. Antarctic Science. 19:31-38.   10.1017/s0954102007000065   AbstractWebsite

The arrival in January 2001 in the south-west Ross Sea of two giant icebergs, C16 and Bl5A, subsequently had dramatic affects on two emperor penguin colonies. B15A collided with the north-west tongue of the Ross Ice Shelf at Cape Crozier, Ross Island, in the following months and destroyed the penguins' nesting habitat. The colony totally failed in 2001, and years after, with the icebergs still in place, exhibited reduced production that ranged from 0 to 40% of the 1201 chicks produced in 2000. At Beaufort Island, 70 km NW of Crozier, chick production declined to 6% of the 2000 count by 2004. Collisions with the Ross Ice Shelf at Cape Crozier caused incubating adults to be crushed, trapped in ravines, or to abandon the colony and, since 2001, to occupy poorer habitat. The icebergs separated Beaufort Island from the Ross Sea Polynya, formerly an easy route to feeding and wintering areas. This episode has provided a glimpse of events which have probably occurred infrequently since the West Antarctic Ice Sheet began to retreat 12 000 years ago. The results allow assessment of recovery rates for one colony decimated by both adult and chick mortality, and the other colony by adult abandonment and chick mortality.

Kooyman, G, Ponganis PJ.  1990.  Behavior and physiology of diving in emperor and king penguins. Penguin biology. ( Davis L, Darby JT, Eds.).:14., San Diego: Academic Press Abstract
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.

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.

Kooyman, GL, Ponganis PJ.  2007.  The initial journey of juvenile emperor penguins. Aquatic Conservation-Marine and Freshwater Ecosystems. 17:S37-S43.   10.1002/aqc.930   AbstractWebsite

1. The first major journey of emperor penguins, among several in their lifetime, is the juveniles' dispersal from their natal colony on a trip that takes them beyond Antarctic waters. The route taken by fledglings from Cape Washington (74.5 degrees S; 165.4 degrees E) was Studied by applying satellite transmitters to ten individuals during December 1994-1996. In January 2001 transmitters with longer transmission capacity were also applied to six hand-fed fledglings, which had been held captive for one month while attaining a body mass exceeding that of wild birds. These post-captive birds were released at the ice edge of McMurdo Sound (77.5 degrees S; 165.0 degrees E), which is in the vicinity of other emperor penguin colonies, and 320km south of their natal colony of Cape Washington. 2. Independent of their parents, the wild birds travelled north-east for the next two months, reaching locations as low as 57 degrees S. The post-captive birds travelled north also, but their trek reached only to about 63 degrees S before they turned south, or remained near their most northerly position from March through May. 3. It was concluded that among colonies in the southern Ross Sea: (a) most healthy fledglings Survive at least the first two months at sea, feeding themselves as they go; (b) the Cape Washington fledglings travelled as far north as 57 degrees S, and much of this journey was in ice free waters; (c) by April, the post-captive birds reached at least as far as the large-scale pack ice edge and possibly beyond the edge Lit 63 degrees S; (d) by early March the trend north ends, and by about late March the birds travel to, or remain near the northern ice edge. 4. The reason the birds travel so far north remains a mystery. Copyright (c) 2008 John Wiley & Sons, Ltd.

Kooyman, GL, Ponganis PJ.  2004.  The icing of external recorders during the polar winter. Memoirs of National Institute of Polar Research Special Issue. 58:183-187. AbstractWebsite
Kooyman, GL, Ponganis PJ.  1997.  The challenges of diving to depth. American Scientist. 85:530-539. AbstractWebsite