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Cambridge University Press 978-0-521-76555-8 - Diving Physiology of Marine Mammals and Seabirds Paul J. Ponganis Index More information

Index

Adélie penguin aorta dive behavior, 18 aortic bulb aerobic dive limit anatomy, 122 ADL, 172 windkessel, 123 variability, 179 aortic windkessel ADL calculation, 175 coronary perfusion, 123 assumptions and limitations, 175 ascending aorta and arch diving metabolic rate, 175 windkessel in whales, 123 oxygen stores, 175 ascending aortic compliance behavioral ADL, 175 ventricular impedance, 123 calculated ADL, 175 descending aorta deﬁnition, 172 noncompliant structure, 124 diving lactate threshold, 176 Arnoux’s beaked whale measured ADL, 172 dive behavior, 9 Baikal seals, 173 arterialization

beluga whales, 173 effect on blood O2 store calculation, 53 bottlenose dolphins, 173 elephant seals – free dives, 92, 184 California sea lions, 173 emperor penguins – free dives, 107 emperor penguins, 172 arterio-venous (A-V) anastomoses Weddell seals, 172 brood patch, 128, 157 physiological basis duck, 100 muscle oxygen depletion – emperor penguin, feet and eyelids 177 birds, 128 African darter skin, 76, 121 dive behavior, 17 sympathetic innervation – bowhead whales, 77 African penguin arterio-venous (A-V) shunts dive behavior, 18 arterio-venous anastomoses, 76 air exhalation emperor penguins Antarctic fur seals, 189 free dives, 194 air sacs, 40 emperor penguins – free dives, 107 ama forced submersion – duck, 100 dive performance, 22 forced submersions, 76, 150 nitrogen levels, 27 in foreflipper of seals, 76 oxygen and carbon dioxide levels, 23 free dives – elephant seals, 76 American coot Atlantic puffin dive behavior, 20 dive behavior, 15 Antarctic fur seal Atlantic white-sided dolphin dive behavior, 7 dive behavior, 9 Antarctic prion atmospheres absolute dive behavior, 13 definition, 27 Antarctic shag Audubon’s shearwater dive behavior, 16 dive behavior, 13

321

© in this web service Cambridge University Press www.cambridge.org Cambridge University Press 978-0-521-76555-8 - Diving Physiology of Marine Mammals and Seabirds Paul J. Ponganis Index More information

322 Index

Australasian grebe blood volumes dive behavior, 19 avian divers, 53 Australian fur seal marine mammals, 53 dive behavior, 7 blue petrel Australian sea lion dive behavior, 13 dive behavior, 7 blue whale avian hemodynamics – forced submersion controlled exposure study, 218 central venous pressure, 131 dive behavior, 10 left ventricle distention, 131 blue-eyed shag pulmonary vascular resistance, 131 dive behavior, 16 right atrium and ventricle distention, 131 blue-footed booby ventricular contractility, 131 dive behavior, 17 body mass Baikal seal advantage of increased size dive behavior, 3 basal metabolic rate, 70 Baird’s beaked whale cost of locomotion, 70 dive behavior, 9 bottlenose dolphin Balearic shearwater dive behavior, 9 dive behavior, 13 bowhead whale bar-headed goose dive behavior, 10 hemoglobin, 46 Boyle’s Law, 25 Barolo shearwater bradycardia, 71 dive behavior, 13 duck – forced submersion, 98 barotrauma, 205 spontaneous dives – Baikal seals, 83 air-ﬁlled cavities, 207 break point cranial sinuses, 207 physiology, 23 middle ear cavities, 207 bronchiolar sphincters, 35 bearded seal Bryde’s whale dive behavior, 3 dive behavior, 10 beaver Bulwer’s petrel dive behavior, 12 dive behavior, 13 Beluga whale dive behavior, 9 California sea lion black-browed albatross dive behavior, 7 dive behavior, 13 canvasback duck black-vented shearwater dive behavior, 19 dive behavior, 13 cape cormorant Blainville’s beaked whale dive behavior, 16 controlled exposure study, 217 cape gannet dive behavior, 9 dive behavior, 17 blood lactate concentrations capybara forced submersions, 199 dive behavior, 12 free dives – emperor penguins, 199 carbon dioxide free dives – seals, 199 end tidal, 198 sleep apnea, 181 forced submersion – ducks, 197 blood pH forced submersion – seals, 196 forced submersion – ducks, 197 free dives – emperor penguins, 198 forced submersion – seals, 196 free dives – Weddell seal, 197 free dives – emperor penguins, 198 sleep apnea, 197 free dives – seals, 197 storage in humans, 24 post-dive in seals, 200 trained submersion – cetaceans, 197 stationary submersions – cetaceans, carbon monoxide 197 dissociation curve shift, 46 blood pressure hemoglobin afﬁnity, 45 forced submersion, 75 in seals, 53 forced submersion – duck, 98 mitochondrial biogenesis, 47 formula, 77 reperfusion injury, 46

Index 323

cardiac output bubble gas content, 214 tissue distribution at rest – humans, 73 dysbaric osteonecrosis – sperm whales, 213 Caspian seal fat emboli – necropsy, 213 dive behavior, 3 intravascular bubbles – necropsy, 213–14 Cassin’s auklet tissue bubbles – stranded dolphins, 214 dive behavior, 15 pathological mechanisms, 206 chinstrap penguin periarticular tissues, 206 dive behavior, 18 rectiﬁed diffusion, 206 chromium 51 label spinal cord white matter, 206 red cell volume technique, 52 supersaturation, 206

common diving petrel tissue N2 solubility, 214 dive behavior, 13 tribonucleation, 206 common dolphin digestion dive behavior, 9 delay in gray seals, 6, 147 common eider, 19 dive reflex, 71, 79 common goldeneye dive response, 71 dive behavior, 19 forced submersion – birds, 97 common loon forced submersion – seals, 71 dive behavior, 19 forced submersion vs. exercise, 74 common murre free dives – birds, 101 dive behavior, 15 free dives – marine mammals, 85 common pochard simulated dives – penguins, 100 dive behavior, 19 simulated dives – seals, 79 common scoter sleep apnea, 83 dive behavior, 19 surface swimming compression hyperoxia cardiac response – birds, 100 emperor penguins, 192–3 cardiac response – marine mammals, 83 countercurrent exchange trained submersion – seals, 80 flukes, flippers, and dorsal fins, 121 diving metabolism manatee fluke, 149 basal metabolic rate reproductive organs, 122, 150 allometric equations, 165 tongues of mysticetes, 122, 151 definition, 165 vascular anatomy, 121 diving metabolic rate wings and feet in birds, 127 emperor penguins, 170 crabeater seal gray seals, 169 dive behavior, 3 Humboldt penguins, 170 Crozet shag sea otters, 169 dive behavior, 16 Steller sea lions, 169 Cuvier’s beaked whale tufted ducks and lesser scaups, 170 controlled exposure study, 218 Weddell seals, 168–9 dive behavior, 9 diving metabolic rate determinants, 162 cytoglobin field metabolic rate function, 44 elephant seals, 168 forced submersion Dall’s porpoise duck muscle metabolic rate, 163

dive behavior, 9 O2 debt of seals, 162 decompression sickness, 205 seal muscle metabolic rate, 163 arterial gas emboli, 213 metabolic rate at rest asymptomatic bubbles, 205 marine mammals, 166 autochthonous bubble formation, 206 oxygen consumption measurements bubble formation, 206 doubly labeled water, 167 bubble formation thresholds, 205 Fick method, 166 disseminated intravascular coagulation, heart rate, 168 206 respirometry, 166 fat, 206 possible mechanisms of metabolic suppression humans, 26 acidosis, 203 marine mammals carbon monoxide, 204

324 Index

diving metabolism (cont.) flightless cormorant hydrogen sulfide, 164 dive behavior, 16 hypoxia-linked, 163 forced submersion arrest of protein synthesis, adrenal blood flow, 76 163 adrenal gland – duck, 99 ion channel arrest, 163 angiography and Doppler flow probes, 75 reverse Pasteur effect, 163 arterio-venous shunts, 76 nitric oxide, 164 arterio-venous shunts – duck, 100 standard metabolic rate, 165 blood pressure, 75 drift dives, 6 brain blood flow, 76 ducks brain blood flow – duck, 99 divers vs. dabblers, 21 brain temperature, 150 dugong catecholamine levels, 76 dive behavior, 11 catecholamine levels – duck, 99 dusky dolphin cortisol levels, 76 dive behavior, 9 dive reflex bradycardia and vasoconstriction, ear 74 ear squeeze, 31 duck, 97 mechanisms of hearing, 31 duck – muscle blood flow, 98 venous plexuses, 31 heat loss – A-V shunts, 76 emperor penguin lactate wash out, 75 dive behavior, 18 mesenteric blood flow, 75 Eurasian otter muscle blood flow – Baikal seal, 78 dive behavior, 12 myocardial blood flow, 78 European shag myocardial blood flow – duck, 99 dive behavior, 16 penguin, 97 Evan’s blue dye renal blood flow, 75 plasma volume technique, 52 tissue perfusion – microspheres, 76 exercise response various diving birds, 99 mechanisms, 74 free dives – birds muscle blood flow eiders – heart rate, 102 local mechanisms, 74 emperor penguins neuroregulation heart rate and stroke rate, 110 baroreceptor reflex, 115 heart rate at isolated dive hole, 106 cardiac sympathetic nerve, 115 heart rates at sea, 109 central command, 114 muscle temperature, 107 exercise pressor reflex, 115 myoglobin saturation, 107 renal sympathetic nerve, 115 venous Hb saturation, 107 extradural vein heart rate, 103–4 anatomy, 120 Humboldt penguins – heart rate, 104 direction and magnitude of flow, 120 king penguins negative inspiratory pressures, 121 heart rate, 104 eye muscle temperature, 107 cornea, 29 macaroni penguins – heart rate, 104 lens and pupil, 29 redhead ducks – heart rate, 101 photoreceptors, 30 rhinoceros auklets – heart rate, 103 tapetum lucidum, 30 South Georgian shags – heart rate, 103 tufted ducks false killer whale chest muscle blood flow, 102 dive behavior, 9 heart rate, 101 fin whale leg blood flow, 102 dive behavior, 10 organ blood flow, 102 finless porpoise free dives – marine mammals, 85 dive behavior, 9 California sea lions flesh-footed shearwater heart rate, 93

dive behavior, 13 venous O2 proﬁles, 94

Index 325

dolphins size and anatomy heart rate, 95 birds, 127 heart rate and stroke rate, 88 hematocrit, 52 elephant seals ﬂuctuations in phocid seals, 119 heart rate, 89 seals – effect of anesthesia, 52

venous O2 saturation, 92 splenic contraction, 52 gray seals – heart rate, 89 hemoglobin heart rate, 85 Bohr effect – marine mammals, 47 manatees – heart rate, 89 Bohr effect – penguins, 48 multiple pinnipeds – heart rate, 89 carboxyhemoglobin, 45 other mammals – heart rate, 96 concentrations in diving birds, 47 Steller sea lions – heart rate and ODBA, concentrations in marine mammals, 47 92 dissociation curve, 45 Weddell seals 2,3 diphosphoglycerate, 45 fetal heart rate, 86 Bohr effect, 45 heart rate and stroke rate, 88 carboxyhemoglobin, 46 heart rates, 85–6 myoinositol 1,3,4,5,6-pentophosphate, 46 hepatic blood flow, 85–6 pH, 45 Mb saturation profiles, 87 temperature, 45 muscle temperature, 88 fetal hemoglobin – not in Weddell seal, 47 renal blood flow, 85–6 Hill coefficient – marine mammals, 47 Hill coefficient – penguins, 48 Galapagos fur seal methemoglobin, 45 dive behavior, 7 oxygen binding capacity, 45

Galapagos penguin P50,45 dive behavior, 18 P50 – diving birds, 47 Galapagos sea lion P50 – marine mammals, 47 dive behavior, 7 structure, 45 Gentoo penguin hemoglobin concentrations dive behavior, 18 avian divers, 53 gray whale marine mammals, 53 dive behavior, 10 Henry’s Law, 23, 25 great cormorant hepatic blood ﬂow dive behavior, 16 free dives, 85–6 great crested grebe hepatic sinus dive behavior, 19 venous capacitance, 119 gray seal high-pressure nervous syndrome dive behavior, 3 human threshold, 27 gray-headed albatross symptoms, 205 dive behavior, 13 threshold, 206 Guadalupe fur seal hoary-headed grebe dive behavior, 7 dive behavior, 19 hooded seal harbor porpoise dive behavior, 3 dive behavior, 9 horned grebe harbor seal dive behavior, 19 dive behavior, 3 human breath-hold divers harp seal “black out” threshold, 23 dive behavior, 3 barotrauma, 25 heart bradycardia, 24 anatomy and size break point, 22 marine mammals, 118 decompression sickness, 26 ductus arteriosus, 118 diuresis, 28 foramen ovale, 118 dive behavior, 22 right ventricle dive response and metabolic rate, 24 birds, 127 ear squeeze, 25 mammals, 118 heart rate, arrhythmias and pressure, 26

326 Index

human breath-hold divers (cont.) possible hypoxia-linked protective mechanisms, heat loss, 28 203 high-pressure nervous syndrome, 27 hypoxia inducible factor, 203 intrathoracic blood volume, 25 regional hypothermia

lung packing, 26 Q10 effect,, 203 lung residual volume and chest squeeze, 25 seal kidney – ischemic tolerance, 200 meditation, 25 seal neurons – intrinsic hypoxic tolerance, muscle sympathetic nerve activity, 77 204 nitrogen narcosis, 27 shift in penguin dissociation curve, 201 oxygen stores, 23 hypoxic pulmonary vasoconstriction, 231 skin sympathetic nerve activity, 77 acidosis, 232 splenic contraction, 24 human pathology, 231 taravana, 27 hypercarbia, 232 Humboldt penguin mechanisms, 231 dive behavior, 18 sea lions humpback whale hypoxic pulmonary vasodilation, 232 dive behavior, 10 hydrogen sulﬁde, 232 hyperventilation vascular dilation risk in human divers, 23 blood nitrite – dolphins, 232 hypoxemic limits carbon monoxide, 233 bar-headed goose at altitude exhaled nitric oxide – dolphins, 233 inspiratory oxygen, 192 nitric oxide, 232 dog arterial oxygen, 181 intra-aortic balloon pump human analogue to aortic bulb, 130 shallow water black-out thresholds, 192 ischemic preconditioning human patients ATP depletion rate, 228 arterial oxygen, 182 calcium homeostasis, 228 humans – ambient air-Mt Everest mitochondrial aldehyde dehydrogenase-2, arterial oxygen, 184 229 seal mitochondrial Kþdependent ATPase, 228 arterial blood oxygen, 181 mitochondrial permeability transition pore, venous blood oxygen, 181 229 hypoxemic tolerance isolated dive hole anaerobic metabolism – seal neuron, 201 emperor penguins, 106, 170, 191 avian tissue capillary densities, 201 Weddell seals, 86, 169, 172 avoidance of reperfusion injury carbon monoxide, 204 Japanese cormorant glutathione, 204 dive behavior, 16 nitrite reductase of myoglobin, 204 Juan Fernandez fur seal peroxidase activity, 204 dive behavior, 7 blood buffering capacity in marine mammals and penguins, 202 killer whale brain capillary densities, 200 dive behavior, 9 glycogen concentrations – brain and heart, 202 king cormorant human comparisons, 200 dive behavior, 16 ischemic pre-conditioning, 204 king penguin mitochondrial distribution, 200 dive behavior, 18 mitochondrial volume densities, 201 muscle buffering capacity in seals and penguins, least grebe 202 dive behavior, 19 neuroglobin, 201 leopard seal cetacean neurons, 202 dive behavior, 3 seal glial cells, 201 lesser scaup oxygen diffusion distances, 200 dive behavior, 19 pekin ducks – “imminent cardiovascular collapse” light blood oxygen, 44, 48, 191 absorption by water, 28

Index 327

light-mantled albatross penguins at sea, 42 dive behavior, 14 marine flighted birds, 41 little auk penguins, 41 dive behavior, 15 diving lung volumes – marine mammals, little penguin 39 dive behavior, 18 lungs and air sacs – penguins-CT scan, 43 locomotion tidal volumes buoyancy – oxygen costs, 143 birds, 43 cormorants – webbed feet, 142 marine mammals, 39 cost of transport total lung capacity-marine mammals, 37 marine mammals, 144 seabirds, 144 macaroni penguin semi-aquatic mammals, 144 dive behavior, 18 drag Magellanic penguin drag equation, 141 dive behavior, 18 oxygen consumption, 141 mallard (pekin) duck surface drag, 141 dive behavior, 20 engulfment drag – rorqual whales, 142 manatee hydrodynamics, 140 dive behavior, 11 appendages, 140 marbled murrelet boundary layer, 140 dive behavior, 15 drag, 141 Mediterranean monk seal feathers and microbubbles, 141 dive behavior, 3 fineness ratio, 140 minke whale laminar flow, 140 dive behavior, 10 maximum body width, 140 Monteiro’s storm petrel Reynolds number, 140 dive behavior, 13 skin and fur, 141 muscle locomotory appendages and thrust, 141 buffering capacity, 137 prolonged glides, 143 histidine dipeptides, 137 sea lion foreflipper capillary density, 138 hydrofoil and paddle, 142 fatty acid oxidation, 138 seal hind flippers fiber types, 133 hydrofoils, 142 Antarctic fur seal, 135 stroke and glide patterns, 142 auklet, 135 buoyancy, 142 beaked whales, 134 stroke rate patterns, 143 California sea lion, 135 pregnant elephant seals, 143 dolphins, 135 tail fluke elephant seal, 135 hydrofoil, 142 emperor penguin, 135 subdermal connective tissue sheath, 142 fin whale, 135 water density and viscosity, 139 gray seal, 135 long-tailed duck narwhal, 135 dive behavior, 19 pilot whales, 134 lung anatomy pygmy sperm whale, 135 airway reinforcement, 34 sea otter, 135 airway vascular plexuses, 34 tufted duck, 135 avian air sacs, 40 Weddell seal, 135 avian airways and lung, 40 glycogen, 137 bird vs mammals, 32 glycolytic enzyme activites, 137 bronchiolar myoelastic sphincters, 35 lipid droplets, 138 “lung collapse”, 33, 207, 209, 220 locomotory muscle, 133 tracheobronchial biomechanics, 35 metabolic suppression, 139 lung volumes mitochondrial volume densities, 138 avian lung and air-sac volumes, 41 myoglobin content, 139 diving air volume marine mammals, 61 ducks, 41 seabirds, 63

328 Index

muscle (cont.) muscle activity, 233 oxidative enzyme activities, 138 muscle maturation, 233 oxygen consumption during ischemia, 139 myoglobin concentration phosphocreatine, 137 marine mammal neonates, 234 temperature, 139 penguin chick, 235 muscle blood flow myoglobin genes, 233 forced submersion, 75 Weddell seals Baikal seal, 78 hypoxia þ lipid, 234 duck, 98 fluctuations, 80 narwhal sleep apnea, 83 dive behavior, 9 surface swimming – ducks, 100 neuroglobin trained submersion, 80 amino acid sequence, 44 muskrat assay, 201 dive behavior, 12 in cetacean neurons, 202 myocardial oxygen supply/demand in seal glial cells, 201 afterload, 130 neuroregulation, 111 aortic bulb, 130 anterior ethmoidal nerve, 111 aortic bulb as intra-aortic balloon pump, 130 baroreceptor, 112 Law of LaPlace, 130 ducks, 113 cardiac preload, 129 carotid chemoreceptor Law of LaPlace, 129 dabbling ducks, 112 vena caval sphincter, 129 diving ducks, 113 heart rate, 128 seals, 112 myocardial contractility, 129 cortical input, 112 effect of vagal input, 129 diving rat model, 112 myoglobin glossopharyngeal nerve, 111 dissociation curve, 49 lung inflation, 112 pH, 49 nucleus ambiguus, 111 temperature, 49 parasympathetic nervous system

facilitation of O2 diffusion, 50 acetylcholine, 113 nitric oxide, 50 vagus nerve, 111 nitrite reductase activity, 204 pontine neurotaxic center, 111 oxygen binding capacity, 49 respiratory center, 111

P50,49 spinal trigeminal nucleus, 111 peroxidase activity, 204 stimuli, 111

reactive O2 species, 50 sympathetic nervous system, 111 structure, 48 neurotransmitters, 113 myoglobin – facilitated diffusion of oxygen, 237 sympathetic nervous system

equipoise PO2, 238 adrenoreceptors, 113 myoglobin diffusion coefficient, 237 sympathetic nervous system oxygen flux equation, 237 cardiac sympathetic fibers, 113 rat myocardium, 238 trigeminal nerve, 111 seal muscle, 238 New Zealand dabchick seal muscle myoglobin diffusion coefficient, 238 dive behavior, 19 translational diffusion New Zealand fur seal net surface charge, 239 dive behavior, 7 myoglobin concentrations New Zealand sea lion avian divers, 63 dive behavior, 7 marine mammals, 61 nitrogen narcosis, 205 myoglobin production threshold, 27, 206 chicks – non-shivering thermogenesis, 235 northern bottlenosed whale emperor penguins – myoglobin mRNA, 236 dive behavior, 9 hypoxia þ exercise, 234 northern elephant seal messenger RNA dive behavior, 3 calcineurin-NFAT, 233 northern fulmar regulation of transcription, 233 dive behavior, 13

Index 329

northern fur seal blood oxygen proﬁles, 184 dive behavior, 7 hepatic sinus Hb saturation, 184

northern gannet venous PO2, 184 dive behavior, 17 emperor penguins northern rockhopper penguin air-sac oxygen, 192 dive behavior, 18 air-sac oxygen depletion rate, 193 arterial oxygen, 193

olfactory adaptations, 29 muscle O2 consumption, 195 oxygen store calculations muscle oxygen, 194 assumptions and potential sources of error, 50 venous oxygen, 193

blood O2 store Weddell seal assumptions, 51 arterial oxygen, 187 blood volume – effects of large spleens, 52 lung, 187 determination of blood volume, 52 muscle oxygen, 187 intial and ﬁnal Hb saturations, 53 Weddell seals

muscle O2 store blood oxygen depletion rate, 187 assumptions, 58 muscle O2 depletion rates, 188 distribution of Mb, 59 simulated dives Mb determinations, 59 penguins muscle mass, 59 air-sac oxygen, 191

respiratory O2 store arterial oxygen, 191 diving air volumes, 50 sleep apnea

net extraction of O2,51 blood oxygen, 181, 187 oxygen store depletion blood oxygen depletion rate, 182 end tidal oxygen, 198 metabolic rate, 183 ﬂume swim muscle blood ﬂow, 181

emperor penguin – maximum muscle O2 muscle O2 depletion rate, 183 consumption, 195 muscle oxygen, 182 forced submersion stationary breath holds pekin duck dolphins blood oxygen, 191 ﬂuke blood oxygen, 190 pekin ducks trained dives blood oxygen depletion rate, 191 dolphins penguins lung, 190 air-sac oxygen, 191 arterial oxygen, 190 Paciﬁc white-sided dolphin

blood O2 depletion rate, 191 dive behavior, 9 muscle oxygen, 191 pericardial venous plexus seals anatomy, 125 blood, 180 blood ﬂow, 126 blood oxygen depletion rate, 181 brown fat, 126 lungs, 180 Peruvian booby muscle, 180 dive behavior, 17 muscle O2 depletion rate, 180 Peruvian diving petrel free dives dive behavior, 13 ama Peruvian pelican blood oxygen depletion rate, 187 dive behavior, 17 Antarctic fur seals pie-billed grebe air exhalation, 189 dive behavior, 19 California sea lion pigeon guillemot arterial oxygen, 189 dive behavior, 15 California sea lions pilot whale blood O2 depletion rate, 190 dive behavior, 9 venous oxygen, 190 platypus elephant seals dive behavior, 12

arterial PO2, 184 polar bear ascent – muscle blood ﬂow, 186 swim behavior, 12

330 Index

portal vein sphincters pressure tolerance – birds cetaceans, 125 barotrauma pressure tolerance bone pneumaticity, 221 barotrauma, 205 middle ear venous sinuses, 221 central pooling of blood, 207 barotrauma chest compliance, 207 rigid lung, 221 cranial sinuses, 207 blood nitrogen levels engorgement of avian pulmonary blood Adélie and gentoo penguins, 224 capillary – hypothesis, 222 emperor penguins, 224 lungs – relative thoracic volumes, 209 king penguins, 224 middle ear cavities, 207 lung relative retial chest volumes, 209 air capillaries, 222 retia mirabilia, 207 blood capillary engorgement, 222 terminal air space engorgement, 207 penguin lung morphometrics, 222 thorax contents, 209 pulmonary blood capillary rigidity, 222 tracheal compliance, 207 lung–air-sac volume ratio, 222 tracheal venous sinuses, 207 pulmonary vascular resistance blood nitrogen levels hypoxia, hypercarbia and acidosis, 130 dolphins, post-dive, 215 hypoxia-induced pulmonary vasodilation, penguins, free dive, 224 131 penguins, pressure chamber, 224 nitric oxide precursors, 131 seals, free dive, 215 seals, pressure chamber, 215 razorbill cellular adaptations dive behavior, 15 cardiac function, 26 red-footed booby enzyme kinetics, 216 dive behavior, 17 glycolytic ﬂux, 216 red-headed duck membrane composition, 216 dive behavior, 19 platelet function, 58, 216 red-necked grebe decompression sickness, 205 dive behavior, 19 lack of intravascular bubbles – live animals, red-tailed tropic bird 214 dive behavior, 17 models – nitrogen kinetics, 219 renal blood ﬂow

N2 solubilities, 214 forced submersion, 75 thoraco-spinal retia model, 213 free dives, 85–6 tissue bubbles – live stranded dolphins, 214 reperfusion injury, 226 depth of lung collapse reactive oxygen species (ROS), 226 dolphins, 210 thiobarbituric acid reactive substances (TBARS), seals, 209 227 sea lions, 212 xanthine oxidase pathway, 226 gas exchange reperfusion injury – protection heart rate at depth – penguins, 224 antioxidant enzymes intrapulmonary shunting, 223 penguins, 228 high-pressure nervous syndrome, 205 seals, 228 lung blood antioxidant capacity – penguins, 228 penguin lung morphometrics, 223 carbon monoxide, 46 lung collapse catalase, 226 arterial nitrogen levels – seals, 209 glutathione, 227

arterial PO2 proﬁles – sea lions, 211 seals, 228 arterial PO2 proﬁles – seals, 209 glutathione peroxidase, 226 graded pulmonary shunt, 211–12 glutathione-S-transferase, 227 hyperbaric CT scans, 211 hypoxanthine concentrations – seals, 227

muscle N2 levels – dolphins, 210 ischemic preconditioning, 228 surfactant, 212, see surfactant myoglobin nitrite reductase, 165 tracheal anatomy and compliance, 211 myoglobin peroxidase, 228 nitrogen narcosis, 205 seal brain cell hypoxic tolerance, 227 penguin air-sac and lung volumes, 223 seal kidney ischemic tolerance, 227

Index 331

superoxide dismutase, 226 heart rate, 79 seal hearts, 227 penguins – heart rate, 100 thiobarbituric acid reactive substances – seals, 227 sinus arrhythmias, 82 respiratory drive sleep apnea, 82 hypercapneic, 198 cardiac output, 83 hypoxic, 198 heart rate, 83 respiratory mechanics, 36 muscle blood flow, 83 expiratory flow rates – marine mammals, 36 muscle oxygenation, 83 flow volume curves – cetacean vs human, 37 slender-billed petrel relaxation volumes – marine mammals, 36 dive behavior, 13 retia mirabilia sonar-associated strandings, 216 anatomy, 124 controlled exposure studies, 217

narwhal and beluga whale, 124 models – N2 absorption and distribution, baroprotection, 207 219 penguins necropsy findings, 216 ophthalmic, 127 rectified diffusion, 217 possible functions, 124 sound resonance, 217 relative thoracic volumes, 209 sooty shearwater sperm whales, 124 dive behavior, 13 thoracic retia sound production kogiid whales, 124 mechanisms, 30 thoracic retia and spinal meningeal artery, 124 South African fur seal rheology dive behavior, 7 blood viscosity, 57 South American fur seal blood viscosity – red cell characteristics, 58 dive behavior, 7 blood viscosity – shear rate, 57 South Georgian diving petrel blood viscosity and systemic vascular resistance, dive behavior, 13 57 southern elephant seal risk of thrombosis, 58 dive behavior, 3 absence of Factor XII in cetaceans, 58 southern rockhopper penguin clot lysis in seals, 58 dive behavior, 18 clotting factors and platelets in seals, 58 southern sea lion platelet function, 58 dive behavior, 7 rhinoceros auklet specific heat dive behavior, 15 definition, 28 ribbon seal of water, 28 dive behavior, 3 sperm whale right whale dive behavior, 10 dive behavior, 10 spinner dolphin ringed seal dive behavior, 9 dive behavior, 3 splanchnic blood flow Risso’s dolphin forced submersion, 75 dive behavior, 9 spleen Ross seal contraction/relaxation times, 120 dive behavior, 3 hematocrit fluctuations, 119 royal penguin human divers – contraction, 24 dive behavior, 18 red cell storage, 119 size pinnipeds, 119 sea otter sympathetic innervation, 119 dive behavior, 12 spotted dolphin shallow water black out dive behavior, 9 physiology, 23 spotted seal short-tailed shearwater dive behavior, 3 dive behavior, 13 star-nosed mole shy albatross dive behavior, 12 dive behavior, 14 Steller sea lion simulated dives, 79 dive behavior, 7

332 Index

streaked shearwater muscle temperature, 161 dive behavior, 13 upper abdomen, 158 sub-Antarctic fur seal upper/lower abdominal, and stomach dive behavior, 7 temperatures, 158 supraventricular tachycardia macaroni penguins facial immersion treatment, 226 abdominal temperature, 158 surface swimming regional heterothermy cardiac output emperor penguins, 161 sea lions, 84 guillemot, 161 seals, 83 king penguins, 158 heart rate South Georgian shags dolphins, 84 abdominal temperatures, 158 sea lions, 84 guillemot, 161 seals, 83 tufted ducks wave-riding dolphins, 85 abdominal temperature, 158

O2 consumption temperature – marine mammals dolphins, 84 Antarctic fur seals pinnipeds, 84 intraabdominal temperature, 155 stroke volume – seals, 83 California sea lions surface swimming – birds vena caval temperature, 155 Adelie penguins – heart rate, 101 dolphins ducks stomach and abdominal temperatures, 152 cardiac output, 100 elephant seal heart rate, 100 aortic, hepatic sinus and extradural vein hepatic blood flow, 100 temperatures, 153 muscle blood flow, 100 elephant seals myocardial blood flow, 100 stomach temperature, 152 renal blood flow, 100 gray seals splanchnic blood flow, 100 stomach temperature, 153 emperor penguin manatee heart rate, 100 stomach temperature, 152

O2 consumption, 100 muskrat gentoo penguins – heart rate, 101 abdominal temperature, 152 surfacer vs. diver, 4 regional heterothermy surfactant dolphin, 151 antiadhesive function, 229 elephant seal, 151, 153 birds, 230 humans, 151 cholesterol, 229 sea otter composition, 229 rectal temperature, 152 human pathologies, 230 Weddell seals phospholipid, 229 arterial temperature, 152 phosphatidyl choline, 229 muscle temperature, 152 production whales leptin – seals, 230 post-mortem temperatures, 151 sea lions, 230 thermal conductance, 157 properties – seals and sea lions, 230 thermal conductivity surface tension, 229 deﬁnition, 28 surfactant proteins, 229 of water, 28 thermoregulation temperature – diving birds baleen whales emperor penguins corpus cavernosum maxillaris, 122 aortic, vena caval, peripheral venous, muscle, body mass and metabolic rate, 147 stomach, subcutaneous, and sub-feather body mass and stroke rate, 147 temperatures, 159 brown fat, 145 gentoo penguins cold prey ingestion, 151, 157 abdominal temperature, 158 convection and conduction, 145 king penguins dive response – forced submersion, 150

Index 333

feathers muscle blood flow and oxygenation, 80 air layer, 156 stroke volume, 79 cormorant feather structure, 156 translocation model critical water penetration pressure, 156 elephant seals, 184 structure and density, 156 tufted duck water repellency, 156 dive behavior, 19 forced submersion A-V shunt – cooling, 150 vasoconstriction brain temperature, 150 coronary artery innervation, 78 temperature decline, 146 extramuscular proximal arteries – seals, heat dissipation 77 skin hot spots, 149 forced submersion, 71 heat flux vascular nerve fiber distribution, 119 brood patch, 157 duck, 100 dorsal fin, 150 seals, 77 heat increment of feeding, 145, 147 vena caval sphincter heat loss, 145 anatomy and function, 125 appendages, 149 ventilation appendages – birds, 157 air-sac pressure oscillations – diving, 40 hypothermia avian air flow pattern, 40 neuroprotection, 146 max. oxygen consumption

Q10 effect, 146 marine mammals, 39 insulation, 145 penguins, 43 blubber, 148 respiratory rate blood flow, 148 penguins, 43 heat conductivity and thickness, 148 marine mammals, 39 lipid content, 149 tidal volumes temperature difference, 148 birds, 43 feathers, 156 marine mammals, 39 fur – thermal resistance, 148 vibrissae hair density and morphology, 148 sensory function, 30 metabolic efficiency, 145 muscle walrus mitochondrial leak, 147 dive behavior, 8 secondary heat production, 146 wandering albatross respiratory heat loss, 150 dive behavior, 14 shivering, 145 Weddell seal specific dynamic action, 145 dive behavior, 3 vasoconstriction, 145 wedge-tailed shearwater water dive behavior, 13 heat capacity, 145 western grebe heat conductance, 145 dive behavior, 19 thick-billed murre Westland petrel dive behavior, 15 dive behavior, 13

total body O2 stores white-chinned petrel distribution of O2 stores, 63 dive behavior, 13 marine mammals and seabirds, 63 trained submersions Xantu’s murrelet cardiac output, 80 dive behavior, 15 cerebral blood ﬂow, 80 contraction/relaxation of the spleen, 80 yellow-eyed penguin heart rates, 79 dive behavior, 18