SENSORY BIOPHYSICS OF MARINE MAMMALS

The underwater existence of marine mammals has encouraged a variety of special biophysical adaptations to their environment. Their sensory and communication systems reflect the transmission properties of sea water. For example, vision is keen in spectra that penetrate water best, vocalization is broadband and used at the frequencies that appear to fit their activities best—the differences in sensory use match the intriguing variety of behavior observed for each species. To date most of the observations of animal interactions with their marine environment have dealt with sound. There has been some work on vision and studies are underway to determine animal sensitivities to hydrodynamic pressure, chemical traces and magnetic fields. The species that have been recorded to date are listed and vocalizations are generally compared. Methods for observation of sensory mechanisms are noted along with a discussion of other aspects of marine mammal biophysics including vibrissal sensation and the biophysics of movement in a fluid environment.

[1]  R S Wolfe,et al.  Magnetite in Freshwater Magnetotactic Bacteria , 1979, Science.

[2]  E. Wever,et al.  Sound Reception in the Porpoise as it Relates to Echolocation , 1980 .

[3]  William A. Watkins,et al.  Underwater Sounds of Monodon (Narwhal) , 1971 .

[4]  J. L. Fobes,et al.  Sensory capacities of marine mammals. , 1981, Psychological bulletin.

[5]  J. Terhune,et al.  Masked hearing thresholds of ringed seals. , 1975, The Journal of the Acoustical Society of America.

[6]  J. Terhune,et al.  The harp seal, Pagophilus groenlandicus (Erxleben, 1777). X. The air audiogram. , 1971, Canadian journal of zoology.

[7]  Max O. Kramer,et al.  Hydrodynamics of the Dolphin , 1965 .

[8]  W. Au,et al.  Critical ratio and bandwidth of the Atlantic bottlenose dolphin (Tursiops truncatus) , 1983 .

[9]  T. Bullock,et al.  Soviet literature on the nervous system and psychobiology of Cetacea. , 1979, International review of neurobiology.

[10]  Douglas Wartzok,et al.  Color discrimination by a bering sea spotted seal, Phoca largha , 1978, Vision Research.

[11]  R. Schusterman,et al.  Vocal displays under water by the gray seal, the harbor seal,. and the stellar sea lion , 1970 .

[12]  W. F. Flanigan,et al.  Marine mammal chemoreception , 1980 .

[13]  L. Herman,et al.  Bottle-nosed dolphin: double-slit pupil yields equivalent aerial and underwater diurnal acuity. , 1975, Science.

[14]  M. Fuller,et al.  Magnetic material in the head of the common Pacific dolphin. , 1981, Science.

[15]  F. T. Awbrey,et al.  A classification and comparison of vocalizations of captive killer whales (Orcinus orca) , 1982 .

[16]  William A. Watkins,et al.  Click Sounds from Animals at Sea , 1980 .

[17]  R J Schusterman,et al.  Underwater audiogram of the California sea lion by the conditioned vocalization technique. , 1972, Journal of the experimental analysis of behavior.

[18]  T. Raslear The use of the cochlear microphonic response as an indicant of auditory sensitivity: review and evaluation. , 1974, Psychological bulletin.

[19]  R. W. Floyd,et al.  Measurement of echolocation signals of the Atlantic bottlenose dolphin, Tursiops truncatus Montagu, in open waters. , 1974, The Journal of the Acoustical Society of America.

[20]  D. W. Watkins,et al.  Grating resolution and refraction in the cat estimated from evoked cerebral potentials. , 1973, Vision research.

[21]  J. Lythgoe Visual pigments and visual range underwater. , 1968, Vision research.

[22]  J. L. Gould,et al.  Bees Have Magnetic Remanence , 1978, Science.

[23]  M. Matthews,et al.  Critical flicker frequency in a harp seal: Evidence for duplex retinal organization , 1975, Vision Research.

[24]  Richard A. Altes,et al.  Sonar for generalized target description and its similarity to animal echolocation systems. , 1975, The Journal of the Acoustical Society of America.

[25]  K. Pryor,et al.  Observations on a Pygmy Killer Whale (Feresa Attenuata Gray) from Hawaii , 1965 .

[26]  L. V. Worthington,et al.  Underwater Sounds heard from Sperm Whales , 1957, Nature.

[27]  C W Clark,et al.  Sound Playback Experiments with Southern Right Whales (Eubalaena australis) , 1980, Science.

[28]  William A. Watkins,et al.  Sperm whale codas , 1977 .

[29]  R. Pepper,et al.  In-air visual acuity of the bottlenose dolphin. , 1973, Experimental neurology.

[30]  D E Mitchell,et al.  Visual Resolution and Experience: Acuity Deficits in Cats Following Early Selective Visual Deprivation , 1973, Science.

[31]  D. K. Caldwell,et al.  Individualized Whistle Contours in Bottle-nosed Dolphins (Tursiops truncatus) , 1965, Nature.

[32]  A. Popper,et al.  Sound localization by the bottlenose porpoise Tursiops truncatus. , 1975, The Journal of experimental biology.

[33]  C. K. Tayler,et al.  Diurnal Activity Cycles in Captive and Free-Ranging Indian Ocean Bottlenose Dolphins (Tursiops Ad Uncus Ehrenburg) , 1973 .

[34]  W. Keeton,et al.  Magnets interfere with pigeon homing. , 1971, Proceedings of the National Academy of Sciences of the United States of America.

[35]  J. Pettigrew,et al.  Ferromagnetic coupling to muscle receptors as a basis for geomagnetic field sensitivity in animals , 1980, Nature.

[36]  T. Poulter,et al.  Innervation of the vibrissae of the California sea lion, Zalophus californianus , 1973, The Anatomical record.

[37]  C H Loh,et al.  Multiple scale analysis of the spirally coiled cochlea. , 1983, The Journal of the Acoustical Society of America.

[38]  F. Fraser Sound Emitted by Dolphins , 1947, Nature.

[39]  T G Lang,et al.  Swimming Speed of a Pacific Bottlenose Porpoise , 1966, Science.

[40]  John K. B. Ford,et al.  Underwater acoustic signals of the narwhal (Monodon monoceros) , 1978 .

[41]  R. Schusterman,et al.  Visual Acuity of the Harbour Seal and the Steller Sea Lion under Water , 1970, Nature.

[42]  S H Ridgway,et al.  Auditory brainstem response in dolphins. , 1981, Proceedings of the National Academy of Sciences of the United States of America.

[43]  J. Kirschvink The horizontal magnetic dance of the honeybee is compatible with a single-domain ferromagnetic magnetoreceptor. , 1981, Bio Systems.

[44]  J. L. Gould,et al.  Pigeons have magnets. , 1979, Science.

[45]  William A. Watkins,et al.  Underwater Sounds of Cephalorhynchus heavisidii (Mammalia:Cetacea) , 1977 .

[46]  William A. Watkins,et al.  Listening to Hawaiian Spinner Porpoises, Stenella cf. Longirostris, with a Three-Dimensional Hydrophone Array , 1974 .

[47]  W. McFarland,et al.  Cetacean visual pigments. , 1971, Vision research.

[48]  K. Pryor,et al.  Hydrodynamic Performance of Porpoises (Stenella attenuata) , 1966, Science.

[49]  James H. W. Hain,et al.  Vocalizations and Feeding Behavior of the Killer Whale (Orcinus orca) , 1979 .

[50]  S. S. Stevens,et al.  Critical Band Width in Loudness Summation , 1957 .

[51]  C. S. Johnson Masked tonal thresholds in the bottlenosed porpoise. , 1968, The Journal of the Acoustical Society of America.

[52]  W. Watkins,et al.  Underwater Calls of Trichechus (Manatee) , 1965, Nature.

[53]  W. Watkins,et al.  Underwater Sounds of Pinnipeds , 1963, Science.

[54]  P. O. Thompson,et al.  Underwater sounds of migrating gray whales, Eschrichtius glaucus (Cope). , 1968, The Journal of the Acoustical Society of America.

[55]  M. C. Caldwell,et al.  STATISTICAL EVIDENCE FOR INDIVIDUAL SIGNATURE WHISTLES IN THE PACIFIC WHITESIDED DOLPHIN, LAGENORHYNCHUS OBLIQUIDENS , 1970 .

[56]  J. Ling The skin and hair of the southern elephant seal, Mirounga leonina (Linn.) I. The facial vibrissae , 1966 .

[57]  K. Ronald,et al.  Pinniped visual pigments. , 1975, Comparative biochemistry and physiology. B, Comparative biochemistry.

[58]  S. S. Easter Retinal Specialisations for Aquatic Vision: Theory and Facts , 1975 .

[59]  E. Hobson,et al.  Visual Orientation and Feeding in Seals and Sea Lions , 1966, Nature.

[60]  R. Schusterman Auditory sensitivity of a California sea lion to airborne sound. , 1974, The Journal of the Acoustical Society of America.

[61]  R. Payne,et al.  Songs of Humpback Whales , 1971, Science.

[62]  L. A. Rivamonte Eye model to account for comparable aerial and underwater acuities of the bottlenose dolphin , 1976 .

[63]  J. Thomas,et al.  Quantitative analysis of Weddell seal (Leptonychotes weddelli) underwater vocalizations at Mcmurdo Sound, Antarctica. , 1982, The Journal of the Acoustical Society of America.

[64]  K. Ronald,et al.  The harp seal, Pagophilus groenlandicus (Erxleben 1777). XXIII. Spectral sensitivity. , 1972, Canadian journal of zoology.

[65]  R. Brownell,et al.  Blind River Dolphin: First Side-Swimming Cetacean , 1969, Science.

[66]  W E Schevill,et al.  Underwater Listening to the White Porpoise (Delphinapterus leucas). , 1949, Science.

[67]  R. Dykes Afferent fibers from mystacial vibrissae of cats and seals. , 1975, Journal of neurophysiology.

[68]  L. Herman,et al.  Symbolic, identity, and probe delayed matching of sounds by the bottlenosed dolphin , 1982 .

[69]  L M Herman,et al.  Underwater frequency discrimination in the bottlenosed dolphin (1-140 kHz) and the human (1-8 kHz). , 1975, The Journal of the Acoustical Society of America.

[70]  J. L. Brown,et al.  Dark adaptation and spectral sensitivity in the cat. , 1970, Vision research.

[71]  S. Ridgway,et al.  Synchrony of dolphin eye movements and their power density spectra , 1981 .

[72]  A. Breathnach THE CETACEAN CENTRAL NERVOUS SYSTEM , 1960 .

[73]  R. Busnel,et al.  ÉTUDES PRÉLIMINAIRES DE SIGNAUX ACOUSTIQUES DU PONTOPORIA BLAINVILLEI GERVAIS ET D’ORBIGNY (CETACEA, PLATANISTIDAE) , 1974 .

[74]  S. Duntley Light in the Sea , 1963 .

[75]  R. Busnel,et al.  Animal Sonar Systems , 1980, NATO Advanced Study Institutes Series.

[76]  A. Dral Vision in Cetacea , 1975 .

[77]  W. Au,et al.  Target detection in reverberation by an echolocating Atlantic bottlenose dolphin (Tursiops truncatus). , 1983, The Journal of the Acoustical Society of America.

[78]  K. E. Moore,et al.  Radio tracking of finback (Balaenoptera physalus) and humpback (Megaptera novaeangliae) whales in Prince William Sound, Alaska☆ , 1981 .

[79]  D. K. Caldwell,et al.  Vocalization of Naive Captive Dolphins in Small Groups , 1968, Science.

[80]  L. Herman,et al.  Discrimination of auditory temporal differences by the bottlenose dolphin and by the human. , 1974, The Journal of the Acoustical Society of America.

[81]  D. K. Caldwell,et al.  STATISTICAL EVIDENCE FOR INDIVIDUAL SIGNATURE WHISTLES IN THE SPOTTED DOLPHIN, STENELLA PLAGIODON. , 1970 .

[82]  W. E. Schevill,et al.  Evidence for echolocation by cetaceans , 1956 .

[83]  R J Schusterman Low false-alarm rates in signal detection by marine mammals. , 1974, The Journal of the Acoustical Society of America.

[84]  W. Watkins,et al.  Underwater Playback of Their Own Sounds to Leptonychotes (Weddell Seals) , 1968 .

[85]  W. Watkins,et al.  Click Structure in the Porpoise, Phocoena Phocoena , 1969 .

[86]  B. Møhl,et al.  Echolocation: high-frequency component in the click of the harbour porpoise (Phocoena ph. L.). , 1973, The Journal of the Acoustical Society of America.

[87]  R. Schusterman,et al.  AERIAL AND UNDERWATER VISUAL ACUITY IN THE CALIFORNIA SEA LION (ZALOPHUS CALIFORNIANUS) AS A FUNCTION OF LUMINANCE * , 1971, Annals of the New York Academy of Sciences.

[88]  G. Oliver Navigation in Mazes By a Grey Seal, Halichoer Us Gr Yp Us (Fabricius) , 1978 .

[89]  J M Terhune,et al.  The harp seal, Pagophilus groenlandicus (Erxleben, 1777). 3. The underwater audiogram. , 1972, Canadian journal of zoology.

[90]  W. N. Kellogg,et al.  Echo ranging in the porpoise; perception of objects by reflected sound is demonstrated for the first time in marine animals. , 1958, Science.

[91]  W. E. Evans Echolocation by marine delphinids and one species of fresh‐water dolphin , 1973 .

[92]  J. Wallman A simple technique using an optomotor response for visual psychophysical measurements in animals , 1975, Vision Research.

[93]  Hal Whitehead,et al.  Identification of Humpback Whales by Fluke Photographs , 1979 .

[94]  H. E. Winn,et al.  The song of the humpback whale Megaptera novaeangliae in the West Indies , 1978 .

[95]  Evidence that seals may use echolocation , 1982, Nature.

[96]  V. SOKOLOV,et al.  Interaction of Dolphin Epidermis with Flow Boundary Layer , 1969, Nature.

[97]  Echo sounder records of ultrasonic sounds made by killer whales and dolphins , 1961 .

[98]  Peggy L. Edds,et al.  Vocalizations of the Blue Whale, Balaenoptera musculus, in the St. Lawrence River , 1982 .

[99]  C. S. Johnson,et al.  Auditory Thresholds of a Killer Whale Orcinus orca Linnaeus , 1972 .

[100]  Kenneth S. Norris,et al.  AN EXPERIMENTAL DEMONSTRATION OF ECHOLOCATION BEHAVIOR IN THE PORPOISE, TURSIOPS TRUNCATUS (MONTAGU) , 1961 .

[101]  P. Moore Underwater localization of click and pulsed pure-tone signals by the California sea lion (Zalophus californianus). , 1975, The Journal of the Acoustical Society of America.

[102]  M. J. White An evaluation of tagging, marking, and tattooing techniques for small delphinids , 1981 .

[103]  P. Morgane,et al.  The anatomy of the brain of the bottlenose dolphin (Tursiops truncates). Rhinic lobe (rhinencephalon). I. The paleocortex , 1971, The Journal of comparative neurology.

[104]  B. Lawrence,et al.  High‐Frequency Auditory Response of a Bottlenosed Porpoise, Tursiops truncatus (Montagu) , 1953 .

[105]  P. O. Thompson,et al.  Underwater Sounds from the Blue Whale, Balaenoptera musculus , 1971 .

[106]  J. D. Lorge,et al.  Operant Methods Assessing the Effects of ELF Electromagnetic Fields , 1974 .

[107]  W. Watkins,et al.  Characteristic Features of the Underwater Sounds of Cephalorhynchus commersonii , 1980 .

[108]  W. Montagna Comparative anatomy and physiology of the skin. , 1967, Archives of dermatology.

[109]  P. O. Thompson,et al.  Sound Production And Other Behavior Of Southern Right Whales Eubalaena-Glacialis , 1972 .

[110]  I. Stirling Vocalization in the Ringed Seal (Phoca hispida) , 1973 .

[111]  Donald B. Siniff,et al.  Underwater vocalizations of leopard seals (Hydrurga leptonyx) and crabeater seals (Lobodon carcinophagus) near the South Shetland Islands, Antarctica , 1979 .

[112]  J. L. Gould,et al.  Biogenic magnetite as a basis for magnetic field detection in animals. , 1981, Bio Systems.

[113]  J. Northrop,et al.  Underwater 20‐Hz Signals Recorded near Midway Island , 1971 .

[114]  W. Watkins,et al.  Right Whale Feeding and Baleen Rattle , 1976 .

[115]  J. Phillips,et al.  Directional and Discriminatory Responses of Salamanders to Weak Magnetic Fields , 1978 .

[116]  Delphinid sonar: measurement and analysis , 1973 .

[117]  H. Dartnall,et al.  Human Visual Purple , 1953, Nature.

[118]  R. Stewart,et al.  Influence of vessel noises on underwater vocal activity of harp seals , 1979 .

[119]  D. Renouf Fishing in Captive Harbour Seals (Phoca Vitulina Concolor): a Possible Role for Vibrissae , 1979 .

[120]  Distinctive characteristics of underwater calls of the harp seal, Phoca groenlandica, during the breeding season , 1979 .

[121]  R. Schusterman Behavioral Capabilities of Seals and Sea Lions: A Review of Their Hearing, Visual, Learning and Diving Skills , 1981 .

[122]  D. K. Caldwell,et al.  SIMULTANEOUS BUT DIFFERENT NARROW-BAND SOUND EMISSIONS BY A CAPTIVE EASTERN PACIFIC PILOT WHALE, GLOBICEPHALA SCAMMONI , 1969 .