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Spragg, RG, Ponganis PJ, Marsh JJ, Rau GA, Bernhard W.  2004.  Surfactant from diving aquatic mammals. Journal of Applied Physiology. 96:1626-1632.   10.1152/japplphysiol.00898.2003   AbstractWebsite

Diving mammals that descend to depths of 50 - 70 m or greater fully collapse the gas exchanging portions of their lungs and then reexpand these areas with ascent. To investigate whether these animals may have evolved a uniquely developed surfactant system to facilitate repetitive alveolar collapse and expansion, we have analyzed surfactant in bronchoalveolar lavage fluid (BAL) obtained from nine pinnipeds and from pigs and humans. In contrast to BAL from terrestrial mammals, BAL from pinnipeds has a higher concentration of phospholipid and relatively more fluidic phosphatidylcholine molecular species, perhaps to facilitate rapid spreading during alveolar reexpansion. Normalized concentrations of hydrophobic surfactant proteins B and C were not significantly different among pinnipeds and terrestrial mammals by immunologic assay, but separation of proteins by gel electrophoresis indicated a greater content of surfactant protein B in elephant seal surfactant than in human surfactant. Remarkably, surfactant from the deepest diving pinnipeds produced moderately elevated in vitro minimum surface tension measurements, a finding not explained by the presence of protein or neutral lipid inhibitors. Further study of the composition and function of pinniped surfactants may contribute to the design of optimized therapeutic surfactants.

Ponganis, PJ, van Dam RP, Knower T, Levenson DH, Ponganis KV.  2004.  Deep dives and aortic temperatures of emperor penguins: new directions for bio-logging at the isolated dive hole. Memoirs of National Institute of Polar Research Special Issue. 58:155-161. AbstractWebsite
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
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.

Ponganis, PJ, Kooyman GL, h. Ridgway S.  2003.  Comparative Diving Physiology. Bennett and Elliott's physiology and medicine of diving. ( Brubakk AO, Neuman TS, Bennett PB, Elliott DH, Eds.).:16., Edinburgh; New York: Saunders Abstract
Ponganis, PJ, Kreutzer U, Sailasuta N, Knower T, Hurd R, Jue T.  2002.  Detection of myoglobin desaturation in Mirounga angustirostris during apnea. American Journal of Physiology-Regulatory Integrative and Comparative Physiology. 282:R267-R272. AbstractWebsite

H-1 NMR solution-state study of elephant seal (Mirounga angustirostris) myoglobin (Mb) and hemoglobin (Hb) establishes the temperature-dependent chemical shifts of the proximal histidyl NdeltaH signal, which reflects the respective intracellular and vascular PO2 in vivo. Both proteins exist predominantly in one major isoform and do not exhibit any conformational heterogeneity. The Mb and Hb signals are detectable in M. angustirostris tissue in vivo. During eupnea M. angustirostris muscle maintains a well-saturated MbO(2). However, during apnea, the deoxymyoglobin proximal histidyl NdeltaH signal becomes visible, reflecting a declining tissue PO2. The study establishes a firm methodological basis for using NMR to investigate the metabolic responses during sleep apnea of the elephant seal and to secure insights into oxygen regulation in diving mammals.

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.

Ponganis, PJ.  2002.  Circulatory System. Encyclopedia of marine mammals. ( Perrin WF, Würsig BG, Thewissen J, Eds.).:229-232., San Diego: Academic Press Abstract

This encyclopedia is a comprehensive, scientifically accurate work devoted to all aspects of marine mammals, including their anatomy, physiology, evolution, behavior, reproduction, ecology, and disease, as well as issues of exploitation, conservation, and management.

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.

Nagy, KA, Kooyman GL, Ponganis PJ.  2001.  Energetic cost of foraging in free-diving emperor penguins. Physiological and Biochemical Zoology. 74:541-547.   10.1086/322165   AbstractWebsite

Hypothesizing that emperor penguins (Aptenodytes forsteri) would have higher daily energy expenditures when foraging for their food than when being hand-fed and that the increased expenditure could represent their foraging cost, we measured field metabolic rates (FMR; using doubly labeled water) over 4-d periods when 10 penguins either foraged under sea ice or were not allowed to dive but were fed fish by hand. Surprisingly, penguins did not have higher rates of energy expenditure when they dove and captured their own food than when they did not forage but were given food. Analysis of time-activity and energy budgets indicated that FMR was about 1.7 x BMR (basal metabolic rate) during the 12 h d(-1) that penguins were lying on sea ice. During the remaining 12 h d(-1), which we termed their "foraging period" of the day, the birds were alert and active (standing, preening, walking, and either free diving or being hand-fed), and their FMR was about 4.1 x BMR. This is the lowest cost of foraging estimated to date among the eight penguin species studied. The calculated aerobic diving limit (ADL(C)), determined with the foraging period metabolic rate of 4.1 x BMR and known O-2 stores, was only 2.6 min, which is far less than the 6-min ADL previously measured with postdive lactate analyses in emperors diving under similar conditions. This indicates that calculating ADL(C) from an at-sea or foraging-period metabolic rate in penguins is not appropriate. The relatively low foraging cost for emperor penguins contributes to their relatively low total daily FMR (2.9 x BMR). The allometric relationship for FMR in eight penguin species, including the smallest and largest living representatives, is kJ d(-1) = 1,185 kg(0.705).

Ponganis, PJ, Van Dam RP, Knower T, Levenson DH.  2001.  Temperature regulation in emperor penguins foraging under sea ice. Comparative Biochemistry and Physiology a-Molecular and Integrative Physiology. 129:811-820.   10.1016/s1095-6433(01)00349-x   AbstractWebsite

Inferior vena caval (IVC) and anterior abdominal (AA) temperatures were recorded in seven emperor penguins (Aptenodytes foresteri) foraging under sea ice in order to evaluate the hypothesis that hypothermia-induced metabolic suppression might extend aerobic diving time. Diving durations ranged from 1 to 12.5 min, with 39% of dives greater than the measured aerobic dive limit of 5.6 min. Anterior abdominal temperature decreased progressively throughout dives, and partially returned to pre-dive values during surface intervals. The lowest AA temperature was 19 degreesC. However, mean AA temperatures during dives did not correlate with diving durations. In six of seven penguins, only minor fluctuations in IVC temperatures occurred during diving. These changes were often elevations in temperature. In the one exception, although IVC temperatures decreased, the reductions were less than those in the anterior abdomen and did not correlate with diving durations. Because of these findings, we consider it unlikely that regional hypothermia in emperor penguins leads to a significant reduction in oxygen consumption of the major organs within the abdominal core. Rather, temperature profiles during dives are consistent with a model of regional heterothermy with conservation of core temperature, peripheral vasoconstriction, and cooling of an outer body shell. (C) 2001 Elsevier Science Inc. All rights reserved.

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.

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.

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.

Ponganis, PJ, Kooyman GL.  1999.  Heart rate and electrocardiogram characteristics of a young California gray whale (Eschrichtius robustus). Marine Mammal Science. 15:1198-1207.   10.1111/j.1748-7692.1999.tb00885.x   AbstractWebsite

Electrocardiogram (ECG) analyses of Holter monitor recordings from a young California gray whale were performed to determine ECG waveform characteristics, evaluate the heart rate pattern for sinus arrhythmia, obtain resting heart rates at known body masses as the whale increased in size, and compare those heart rates with predicted heart rates from allometric equations. The PR and QRS intervals (475 +/- 35 msec, 208 +/- 24 msec, respectively, n = 20) support the concept (Meijler et al. 1992) that atrioventricular transmission and ventricular excitation times do not increase linearly in very large mammals. A sinus arrhythmia pattern at rest (apneic heart rates of 15-25 beats per min [bpm] and eupneic heart rates of 34-40 bpm) is consistent with a relative eupneic tachycardia and apneic bradycardia during diving activity of whales. The heart rate-body mass measurements (35-24 bpm at body masses of 3,531-8,200 kg) in this study (1) extend the range of allometric heart rate and body mass data in mammals a full order of magnitude, to almost 10,000 kg, (2) support the use of allometric equations (based primarily on mammals <1,000 kg in body mass) in estimating resting heart rates in whales, and (3) demonstrate that previously reported heart rates in large whales are not representative of resting heart rate, probably secondary to circumstances during measurement.

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.

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.

Ponganis, PJ, Starke LN, Horning M, Kooyman GL.  1999.  Development of diving capacity in emperor penguins. Journal of Experimental Biology. 202:781-786. AbstractWebsite

To compare the diving capacities of juvenile and adult emperor penguins Aptenodytes forsteri, and to determine the physiological variables underlying the diving ability of juveniles, we monitored diving activity in juvenile penguins fitted with satellite-linked time/depth recorders and examined developmental changes in body mass (M-b), hemoglobin concentration, myoglobin (Mb) content and muscle citrate synthase and lactate dehydrogenase activities, Diving depth, diving duration and time-at-depth histograms were obtained from two fledged juveniles during the first 2.5 months after their departure from the Cape Washingon colony in the Ross Sea, Antarctica. During this period, values of all three diving variables increased progressively. After 8-10 weeks at sea, 24-41% of transmitted maximum diving depths were between 80 and 200 m, Although most dives lasted less than 2 min during the 2 month period, 8-25% of transmitted dives in the last 2 weeks lasted 2-4 min. These values are lower than those previously recorded in adults during foraging trips. Of the physiological variables examined during chick and juvenile development, only M-b and Mb content did not approach adult values, In both near-hedge chicks and juveniles, Mb was 50-60% of adult values and Mb content was 24-31% of adult values. This suggests that the increase in diving capacity of juveniles at sea will be most dependent on changes in these factors.

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
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.

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.  1997.  The challenges of diving to depth. American Scientist. 85:530-539. AbstractWebsite
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.

Ponganis, PJ, Costello ML, Starke LN, MathieuCostello O, Kooyman GL.  1997.  Structural and biochemical characteristics of locomotory muscles of emperor penguins, Aptenodytes forsteri. Respiration Physiology. 109:73-80.   10.1016/s0034-5687(97)84031-5   AbstractWebsite

Structural and biochemical characteristics of the primary muscles used for swimming (pectoralis, PEC and supracoracoideus, SC) were compared to those of leg muscles in emperor penguins (Aptenodytes forsteri). The mass of PEG-SC was four times that of the leg musculature, and mitochondrial volume density in PEC and SC (4%) was two-thirds that in sartorius (S) and gastrocnemius. The differences in muscle mass and mitochondrial density yielded a 2.2-fold greater total mitochondrial content in PEG-SC than leg muscles, which appears to account for the 1.8-fold greater whole-body highest oxygen consumption previously recorded in emperor penguins during swimming compared to walking. Calculation of maximal mitochondrial O-2 consumption in PEG-SC and leg muscle yielded values of 5.8-6.9 mi O-2 ml(-1) min(-1), which are similar to those in locomotory muscles of most mammals and birds. A distinct feature of emperor penguin muscle was its myoglobin content, with concentrations in PEG-SC (6.4 g 100 g(-1)) among the highest measured in any species. This resulted in a PEG-SC O-2 store greater than that of the entire blood. In addition, ratios of myoglobin content to mitochondrial volume density and to citrate synthase activity were 4.4 and 2.5 times greater in PEG than in S, indicative of the significant role of myoglobin in the adaptation of muscle to cardiovascular adjustments during diving. (C) 1997 Elsevier Science B.V.

Ponganis, PJ, Kooyman GL, Winter LM, Starke LN.  1997.  Heart rate and plasma lactate responses during submerged swimming and trained diving in California sea lions, Zalophus californianus. Journal of Comparative Physiology B-Biochemical Systemic and Environmental Physiology. 167:9-16.   10.1007/s003600050042   AbstractWebsite

California sea lions, Zalophus californianus, were trained to elicit maximum voluntary breath holds during stationary underwater targeting, submerged swimming, and trained diving. Lowest heart rate during rest periods was 57 bpm. The heart rate profiles in all three protocols were dominated by a bradycardia of 20-50 bpm, and demonstrated that otariid diving heart rates were at or below resting heart rate. Venous blood samples were collected after submerged swimming periods of 1-3 min. Plasma lactate began to increase only after 2.3-min submersions. This rise in lactate and our inability to train sea lions to dive or swim submerged for periods longer than 3 min lead us to conclude that an aerobic limit had been reached. Due to the similarity of heart rate responses and swimming velocities recorded during submerged swimming and trained diving, this 2.3-min limit should approximate the aerobic dive limit in these 40-kg sea lions. Total body O-2 stores, based on measurements of blood and muscle O-2 stores in these animals, and prior lung O-2 Store analyses, were 37-43 ml O-2 kg(-1). The aerobic dive limit, calculated with these O-2 stores and prior measurements of at-sea metabolic rates of sea lions, is 1.8-2 min, similar to that measured by the change in post-submersion lactate concentration.