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Book Chapter
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
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.

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
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
Conference Proceedings
Ponganis, PJ, Gentry RL, Ponganis EP, Ponganis K.  1989.  Analysis of swimming velocity in deep and shallow dives of two northern fur seals, Callorhinus ursinus. Proceedings of the Eighth Biennial Conference on the Biology of Marine Mammals. , Pacific Grove, Calif. Abstract
Ponganis, PJ, Kooyman GL.  1990.  Diving physiology of penguins. Acta XX Congressus Internationalis Ornithologici, Christchurch, New Zealand, 2-9 December 1990. ( Butler PJ, Jones DR, Eds.).:6., Wellington, N.Z.: New Zealand Ornithological Congress Trust Board Abstract
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
Journal Article
Watanabe, S, Sato K, Ponganis PJ.  2012.  Activity time budget during foraging trips of emperor penguins. Plos One. 7   10.1371/journal.pone.0050357   AbstractWebsite

We developed an automated method using depth and one axis of body acceleration data recorded by animal-borne data loggers to identify activities of penguins over long-term deployments. Using this technique, we evaluated the activity time budget of emperor penguins (n = 10) both in water and on sea ice during foraging trips in chick-rearing season. During the foraging trips, emperor penguins alternated dive bouts (4.8 +/- 4.5 h) and rest periods on sea ice (2.5 +/- 2.3 h). After recorder deployment and release near the colony, the birds spent 17.9 +/- 8.4% of their time traveling until they reached the ice edge. Once at the ice edge, they stayed there more than 4 hours before the first dive. After the first dive, the mean proportions of time spent on the ice and in water were 30.8 +/- 7.4% and 69.2 +/- 7.4%, respectively. When in the water, they spent 67.9 +/- 3.1% of time making dives deeper than 5 m. Dive activity had no typical diurnal pattern for individual birds. While in the water between dives, the birds had short resting periods (1.2 +/- 1.7 min) and periods of swimming at depths shallower than 5 m (0.25 +/- 0.38 min). When the birds were on the ice, they primarily used time for resting (90.3 +/- 4.1% of time) and spent only 9.7 +/- 4.1% of time traveling. Thus, it appears that, during foraging trips at sea, emperor penguins traveled during dives >5 m depth, and that sea ice was primarily used for resting. Sea ice probably provides refuge from natural predators such as leopard seals. We also suggest that 24 hours of sunlight and the cycling of dive bouts with short rest periods on sea ice allow emperor penguins to dive continuously throughout the day during foraging trips to sea.

Williams, CL, Sato K, Ponganis PJ.  2019.  Activity, not submergence, explains diving heart rates of captive loggerhead sea turtles. Journal of Experimental Biology. 222   10.1242/jeb.200824   AbstractWebsite

Marine turtles spend their life at sea and can rest on the seafloor for hours. As air-breathers, the breath-hold capacity of marine turtles is a function of oxygen (O-2) stores, O-2 consumption during dives and hypoxia tolerance. However, some physiological adaptations to diving observed in mammals are absent in marine turtles. This study examined cardiovascular responses in loggerhead sea turtles, which have even fewer adaptations to diving than other marine turtles, but can dive for extended durations. Heart rates (f(H)) of eight undisturbed loggerhead turtles in shallow tanks were measured using self-contained ECG data loggers under five conditions: spontaneous dives, resting motionless on the tank bottom, resting in shallow water with their head out of water, feeding on squid, and swimming at the surface between dives. There was no significant difference between resting f(H) while resting on the bottom of the tank, diving or resting in shallow water with their head out of water. f(H) rose as soon as turtles began to move and was highest between dives when turtles were swimming at the surface. These results suggest cardiovascular responses in captive loggerhead turtles are driven by activity and apneic f(H) is not reduced by submergence under these conditions.

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.

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.

Ponganis, PJ, Gentry RL, Ponganis EP, Ponganis KV.  1992.  Analysis of swim velocities during deep and shallow dives of two northern fur seals, Callorhinus ursinus. Marine Mammal Science. 8:69-75.   10.1111/j.1748-7692.1992.tb00126.x   AbstractWebsite

Swim velocities at 15-sec intervals and maximum depth per dive were recorded by microprocessor units on two "mixed diver" adult female northern fur seals during summer foraging trips. These records allowed comparison of swim velocities of deep (> 75 m) and shallow (< 75 m) dives. Deep dives averaged 120 m depth and 3 min duration; shallow dives averaged 30 m and 1.2 min. Mean swim velocities on deep dives were 1.8 and 1.5 m/ sec for the two animals; mean swim velocities on shallow dives were 1.5 and 1.2 m/sec. The number of minutes per hour spent diving during the deep and shallow dive patterns were 11 and 27 min, respectively. Swim velocity, and hence, relative metabolic rate, did not account for the differences in dive durations between deep and shallow dives. The long surface durations associated with deep dives, and estimates of metabolic rates for the observed swim velocities, suggest that deep dives involve significant anaerobic metabolism.

Ponganis, PJ, Stockard TK.  2007.  The Antarctic toothfish: how common a prey for Weddell seals? Antarctic Science. 19:441-442.   10.1017/s0954102007000715   AbstractWebsite

The Antarctic toothfish (Dissostichus mawsoni Norman) has been considered an occasional large prey item of the Weddell seal (Leptonychotes weddellii Lesson) (Kooyman 1967, Calhaem & Christoffel 1969, Testa et al. 1985, Castellini et al. 1992, Davis et al. 1999, Fuiman et al. 2002). The seal's most common prey is the Antarctic silverfish (Pleuragramma antarcticum Boulenger) as well as benthic and sub-ice fish, cephalopods, and crustaceans (Dearborn 1965, Green & Burton 1987, Plotz 1987, Plotz et al. 1991, Castellini et al. 1992, Burns et al. 1998).

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.

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

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

Ponganis, PJ, Kreutzer U, Stockard TK, Lin PC, Sailasuta N, Tran TK, Hurd R, Jue T.  2008.  Blood flow and metabolic regulation in seal muscle during apnea. Journal of Experimental Biology. 211:3323-3332.   10.1242/jeb.018887   AbstractWebsite

In order to examine myoglobin (Mb) function and metabolic responses of seal muscle during progressive ischemia and hypoxemia, Mb saturation and high-energy phosphate levels were monitored with NMR spectroscopy during sleep apnea in elephant seals (Mirounga angustirostris). Muscle blood flow (MBF) was measured with laser-Doppler flowmetry (LDF). During six, spontaneous, 8-12 min apneas of an unrestrained juvenile seal, apneic MBF decreased to 46 +/- 10% of the mean eupneic MBF. By the end of apnea, MBF reached 31 +/- 8% of the eupneic value. The t(1/2) for 90% decline in apneic MBF was 1.9 +/- 1.2 min. The initial post-apneic peak in MBF occurred within 0.20 +/- 0.04 min after the start of eupnea. NMR measurements revealed that Mb desaturated rapidly from its eupenic resting level to a lower steady state value within 4 min after the onset of apnea at rates between 1.7 +/- 1.0 and 3.8 +/- 1.5% min(-1), which corresponded to a muscle O(2) depletion rate of 1-2.3 ml O(2)kg(-1) min(-1). High-energy phosphate levels did not change with apnea. During the transition from apnea to eupnea, Mb resaturated to 95% of its resting level within the first minute. Despite the high Mb concentration in seal muscle, experiments detected Mb diffusing with a translational diffusion coefficient of 4.5 x 10(-7) cm(2) s(-1), consistent with the value observed in rat myocardium. Equipoise P(O2) analysis revealed that Mb is the predominant intracellular O(2) transporter in elephant seals during eupnea and apnea.

Stockard, TK, Levenson DH, Berg L, Fransioli JR, Baranov EA, Ponganis PJ.  2007.  Blood oxygen depletion during rest-associated apneas of northern elephant seals (Mirounga angustirostris). Journal of Experimental Biology. 210:2607-2617.   10.1242/jeb.008078   AbstractWebsite

Blood gases (P-O2, P-CO2, pH), oxygen content, hematocrit and hemoglobin concentration were measured during rest-associated apneas of nine juvenile northern elephant seals. In conjunction with blood volume determinations, these data were used to determine total blood oxygen stores, the rate and magnitude of blood O-2 depletion, the contribution of the blood O-2 store to apneic metabolic rate, and the egree of hypoxemia that occurs during these breath-holds. Mean body mass was 66 +/- 9.7 kg (+/- s.d.); blood volume was 196 +/- 20 ml kg(-1); and hemoglobin concentration was 23.5 +/- 1.5 g dl(-1). Rest apneas ranged in duration from 3.1 to 10.9 min. Arterial P-O2 declined exponentially during apnea, ranging between a maximum of 108 mmHg and a minimum of 18 mmHg after a 9.1 min breath-hold. Venous P-O2 values were indistinguishable from arterial values after the first minute of apnea; the lowest venous P-O2 recorded was 15 mmHg after a 7.8 min apnea. O-2 contents were also similar between the arterial and venous systems, declining linearly at rates of 2.3 and 2.0 ml O-2 dl(-1) min (-1), respectively, from mean initial values of 27.2 and 26.0 ml O-2 dl(-1). These blood O-2 depletion rates are approximately twice the reported values during forced submersion and are consistent with maintenance of previously measured high cardiac outputs during rest-associated breath-holds. During a typical 7-min apnea, seals consumed, on average, 56% of the initial blood O-2 store of 52 ml O-2 kg(-1); this contributed 4.2 ml O-2 kg(-1) min(-1) to total body metabolic rate during the breath-hold. Extreme hypoxemic tolerance in these seals was demonstrated by arterial P-O2 values during late apnea that were less than human thresholds for shallow-water blackout. Despite such low P-O2s, there was no evidence of significant anaerobic metabolism, as changes in blood pH were minimal and attributable to increased P-CO2. These findings and the previously reported lack of lactate accumulation during these breath- holds are consistent with the maintenance of aerobic metabolism even at low oxygen tensions during rest- associated apneas. Such hypoxemic tolerance is necessary in order to allow dissociation of O-2 from hemoglobin and provide effective utilization of the blood O-2 store.

Meir, JU, Robinson PW, Vilchis LI, Kooyman GL, Costa DP, Ponganis PJ.  2013.  Blood oxygen depletion is independent of dive function in a deep diving vertebrate, the northern elephant seal. Plos One. 8   10.1371/journal.pone.0083248   AbstractWebsite

Although energetics is fundamental to animal ecology, traditional methods of determining metabolic rate are neither direct nor instantaneous. Recently, continuous blood oxygen (O-2) measurements were used to assess energy expenditure in diving elephant seals (Mirounga angustirostris), demonstrating that an exceptional hypoxemic tolerance and exquisite management of blood O-2 stores underlie the extraordinary diving capability of this consummate diver. As the detailed relationship of energy expenditure and dive behavior remains unknown, we integrated behavior, ecology, and physiology to characterize the costs of different types of dives of elephant seals. Elephant seal dive profiles were analyzed and O-2 utilization was classified according to dive type (overall function of dive: transit, foraging, food processing/rest). This is the first account linking behavior at this level with in vivo blood O-2 measurements in an animal freely diving at sea, allowing us to assess patterns of O-2 utilization and energy expenditure between various behaviors and activities in an animal in the wild. In routine dives of elephant seals, the blood O-2 store was significantly depleted to a similar range irrespective of dive function, suggesting that all dive types have equal costs in terms of blood O-2 depletion. Here, we present the first physiological evidence that all dive types have similarly high blood O-2 demands, supporting an energy balance strategy achieved by devoting one major task to a given dive, thereby separating dive functions into distinct dive types. This strategy may optimize O-2 store utilization and recovery, consequently maximizing time underwater and allowing these animals to take full advantage of their underwater resources. This approach may be important to optimizing energy expenditure throughout a dive bout or at-sea foraging trip and is well suited to the lifestyle of an elephant seal, which spends >90% of its time at sea submerged making diving its most "natural" state.

Meir, JU, Ponganis PJ.  2010.  Blood temperature profiles of diving elephant seals. Physiological and Biochemical Zoology. 83:531-540.   10.1086/651070   AbstractWebsite

Hypothermia-induced reductions in metabolic rate have been proposed to suppress metabolism and prolong the duration of aerobic metabolism during dives of marine mammals and birds. To determine whether core hypothermia might contribute to the repetitive long-duration dives of the northern elephant seal Mirounga angustirostris, blood temperature profiles were obtained in translocated juvenile elephant seals equipped with a thermistor and backpack recorder. Representative temperature (the y-intercept of the mean temperature vs. dive duration relationship) was 37.2 degrees +/- 0.6 degrees C (n=3 seals) in the extradural vein, 38.1 degrees +/- 0.7 degrees C (n=4 seals) in the hepatic sinus, and 38.8 degrees +/- 16 degrees C (n=6 seals) in the aorta. Mean temperature was significantly though weakly negatively related to dive duration in all but one seal. Mean venous temperatures of all dives of individual seals ranged between 36 degrees and 38 degrees C, while mean arterial temperatures ranged between 35 degrees and 39 degrees C. Transient decreases in venous and arterial temperatures to as low as 30 degrees-33 degrees C occurred in some dives >30 min (0.1% of dives in the study). The lack of significant core hypothermia during routine dives (10-30 min) and only a weak negative correlation of mean temperature with dive duration do not support the hypothesis that a cold-induced Q(10) effect contributes to metabolic suppression of central tissues during dives. The wide range of arterial temperatures while diving and the transient declines in temperature during long dives suggest that alterations in blood flow patterns and peripheral heat loss contribute to thermoregulation during diving.

Ponganis, PJ, Stockard TK, Levenson DH, Berg L, Baranov EA.  2006.  Cardiac output and muscle blood flow during rest-associated apneas of elephant seals. Comparative Biochemistry and Physiology a-Molecular & Integrative Physiology. 144:105-111.   10.1016/j.cbpa.2006.02.009   AbstractWebsite

In order to evaluate hemodynamics and blood flow during rest-associated apnea in young elephant seals (Mirounga angustirostris), cardiac outputs (CO, thermodilution), heart rates (HR), and muscle blood flow (MBF, laser Doppler flowmetry) were measured.. Mean apneic COs and HRs of three seals were 46% and 39% less than eupneic values, respectively (2.1 +/- 0.3 vs. 4.0 +/- 0.1 mL kg(-1) s(-1), and 54 6 vs. 89 14 beats min(-1)). The mean apneic stroke volume (SV) was not significantly different from the eupneic value (2.3 +/- 0.2 vs. 2.7 +/- 0.5 mL kg(-1)). Mean apneic MBF of three seals was 51% of the eupneic value. The decline in MBF during apnea was gradual, and variable in both rate and magnitude. In contrast to values previously documented in seals during forced submersions (FS), CO and SV during rest-associated apneas were maintained at levels characteristic of previously published values in similarly-sized terrestrial mammals at rest. Apneic COs of such magnitude and incomplete muscle ischemia during the apnea suggest that (1) most organs are not ischemic during rest-associated apneas, (2) the blood O-2 depletion rate is greater during rest-associated apneas than during FS, and (3) the blood O-2 store is not completely isolated from muscle during rest-associated apneas. (c) 2006 Elsevier Inc. All rights reserved.

Ponganis, PJ, Kooyman GL, Zornow MH, Castellini MA, Croll DA.  1990.  Cardiac output and stroke volume in swimming harbor seals. Journal of Comparative Physiology B-Biochemical Systemic and Environmental Physiology. 160:473-482.   10.1007/BF00258974   AbstractWebsite

Cardiac output was measured by the thermodilution method in three young harbor seals, at rest and while swimming up to the maximum effort for which they could be trained. Stroke volume was determined by counting heart rate simultaneously with determination of cardiac output. Cardiac outputs varied widely between surface breathing (7.8 and breath-holding while swimming under water (1.8 Stroke volume while at the surface was almost twice the volume while submerged. Surface cardiac output was always near maximal despite work effort, whereas submerged cardiac output gradually increased at higher work efforts. The cardiovascular performance of seals at the maximum MO2 we could induce from them is equivalent to that of the domestic goat.

Ponganis, PJ, Kooyman GL, Zornow MH.  1991.  Cardiac output in swimming California sea lions, Zalophus californianus. Physiological Zoology. 64:1296-1306. AbstractWebsite

Cardiac output was determined by the thermodilution technique in three California sea lions while resting and while swimming. Metabolic rates increased seven-to ninefold above resting rates during maximal exercise. While the sea lions were at rest, stroke volume was also determined by simultaneously counting heart rate during cardiac output determinations. At rest, cardiac output (2.5-3.0 mL kg-1s-1) and stroke volume (2 mL kg-1) were similar to those of harbor seals and terrestrial mammals of similar mass. During exercise, mean cardiac output increased linearly with work load and surface/submerged intervals were short and frequent. The exercise capacity of swimming sea lions appears similar to that of harbor seals, but the exercise response resembles that of terrestrial mammals more than that of harbor seals.

Kooyman, GL, Ponganis PJ.  1997.  The challenges of diving to depth. American Scientist. 85:530-539. AbstractWebsite


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.