Animal Physiology

by Hill, Richard W.; Wyse, Gordon A.; Anderson, Margaret

• ISBN13: 9780878933150
• ISBN10: 0878933158
• Edition: 1st
• Format: Hardcover
• Copyright: 2004-06-01
• Publisher: Sinauer Associates Inc

Table of Contents

PART I Fundamentals of Physiology	1	(90)
Animals and Environments: Function on the Ecological Stage	3	(26)
The Importance of Physiology	4	(1)
Mechanism and Origin: Physiology's Two Central Questions	5	(4)
The study of mechanism: How do modern-day animals carry out their functions?	5	(1)
The study of origin: Why do modern-day animals possess the mechanisms they do?	6	(1)
Natural selection is a key process of evolutionary origin	6	(1)
Mechanism and adaptive significance are distinct concepts that do not imply each other	7	(2)
This Book's Approach to Physiology	9	(1)
Animals	9	(7)
The structural property of an animal that persists through time is its organization	10	(1)
Most cells of an animal are exposed to the internal environment, not the external environment	10	(1)
The internal environment may be permitted to vary when the external environment changes, or it may be kept constant	11	(1)
Homeostasis in the lives of animals: Internal constancy is often critical for proper function	11	(1)
Negative Feedback	12	(1)
Time in the lives of animals: Physiology changes in five time frames	13	(2)
The Evolution of Phenotypic Plasticity	15	(1)
Size in the lives of animals: Body size is one of an animal's most important traits	16	(1)
Environments	16	(6)
Earth's major physical and chemical environments	16	(5)
The environment an animal occupies is often a microenvironment or microclimate	21	(1)
Animals often modify their own environments	22	(1)
Evolutionary Processes	22	(7)
Some processes of evolution are adaptive, others are not	23	(1)
A trait is not an adaptation merely because it exists	24	(1)
Adaptation is studied as an empirical science	24	(2)
Evolutionary potential can be high or low, depending on available genetic variation	26	(3)
Molecules and Cells in Animal Physiology	29	(36)
Cell Membranes and Intracellular Membranes	30	(7)
The lipids of membranes are structured, diverse, and fluid	31	(1)
Proteins endow membranes with numerous functional capacities	32	(2)
Carbohydrates play important roles in membranes	34	(1)
Protein Structure and the Bond That Maintain It	35	(2)
Epithelia	37	(3)
Elements of Metabolism	40	(1)
Enzyme Fundamentals	40	(8)
Enzyme-catalyzed reactions exhibit hyperbolic or sigmoid kinetics	42	(1)
Maximum reaction velocity is determined by the amount and catalytic effectiveness of an enzyme	43	(1)
Enzyme--substrate affinity affects reaction velocity at the substrate concentrations that are usual in cells	44	(1)
Enzymes have specific three-dimensional binding sites that often interact	44	(2)
Enzymes catalyze reversible reactions in both directions	46	(1)
Multiple molecular forms of enzymes occur at all levels of animal organization	46	(2)
Regulation of Cell Function by Enzymes	48	(5)
The types and amounts of enzymes present depend on gene expression and enzyme degradation	48	(1)
Modulation of existing enzyme molecules permits fast regulation of cell function	49	(1)
Gene-Expression Studies Using DNA Microarrays	50	(3)
Evolution of Enzymes	53	(2)
Enzymes Are Instruments of Change in All Time Frames	55	(1)
Cell Signaling: Signal Reception and Cell Signal Transduction	56	(9)
Extracellular signals initiate their effects by binding to receptor proteins	56	(3)
Cell signal transduction often entails sequences of amplifying effects	59	(3)
Several second-messenger systems participate in cell signal transduction	62	(3)
Transport of Solutes and Water	65	(26)
Passive Solute Transport by Simple Diffusion	67	(7)
Concentration gradients give rise to the most elementary form of simple solute diffusion	68	(1)
Electrical gradients often influence the diffusion of charged solutes at membranes	68	(1)
Biological aspects of diffusion across membranes: Some solutes dissolve in the membrane, others require channels	69	(1)
Diffusion of ions across cell membranes is determined by simultaneous concentration and electrical effects	70	(1)
Diffusion often creates challenges for cells and animals	71	(2)
Concentration gradients can create electrical gradients that alter concentration gradients	73	(1)
Passive Solute Transport by Facilitated Diffusion	74	(1)
Active Transport	74	(8)
Active transport and facilitated diffusion are types of carrier-mediated transport	74	(1)
Basic properties of active-transport mechanisms	75	(1)
Primary and secondary active transport differ in their cellular-molecular mechanisms	75	(4)
Active transport across an epithelium does not imply a specific transport mechanism	79	(1)
Cellular Mechanisms of Ion Pumping in Freshwater Fish Gills	80	(1)
Two epithelial ion-pumping mechanisms help freshwater fish maintain their blood composition	80	(2)
Modulation of Channels and Transporters	82	(1)
Osmotic Pressure and Other Colligative Properties of Aqueous Solutions	82	(3)
Physiologists usually express osmotic pressure in osmolar units	84	(1)
Osmotic pressures can be measured in several ways	84	(1)
Osmosis	85	(3)
Quantification and terminology	86	(1)
Hydrostatic pressures develop from osmotic pressures only when two or more solutions interact	86	(1)
Water may dissolve in membranes or pass through water channels during osmosis	87	(1)
Osmosis and solute physiology often interact	87	(1)
Looking Forward	88	(3)
PART II Food, Energy, and Temperature	91	(166)
Nutrition, Feeding, and Digestion	93	(32)
Nutrition	94	(6)
Proteins are ``foremost''	94	(2)
Lipids are required for all membranes and are the principal storage compounds of animals	96	(2)
Carbohydrates are low in abundance in many animals but highly abundant when they play structural roles	98	(1)
Vitamins are essential organic compounds required in small amounts	98	(1)
Elemental nutrition: Many minerals are essential nutrients	99	(1)
Feeding	100	(11)
Many animals feed on organisms that are individually attacked and ingested	101	(2)
Suspension feeding is common in aquatic animals	103	(2)
Symbioses with microbes often play key roles in animal feeding and nutrition	105	(6)
Digestion and Absorption	111	(7)
Vertebrates, arthropods, and molluscs represent three important digestive-absorptive plans	111	(3)
Digestion is carried out by specific enzymes operating in three spatial contexts	114	(2)
Absorption occurs by different mechanisms for hydrophilic and hydrophobic molecules	116	(2)
Responses to Eating	118	(2)
Changes in Digestion and Absorption in Additional Time Frames	120	(5)
The nutritional physiology of individuals responds chronically to changed conditions	120	(1)
Nutritional physiology is under the control of biological clocks	120	(1)
Nutritional physiology undergoes programmed changes during development	120	(1)
Pythons: Extreme Examples of Feast and Famine	121	(1)
Nutritional physiology undergoes evolutionary change	121	(4)
Energy Metabolism	125	(24)
Why Animals Need Energy: The Second Law of Thermodynamics	126	(1)
Fundamentals of Animal Energetics	127	(3)
The forms of energy vary in their capacity for physiological work	127	(1)
Transformations of high-grade energy are always inefficient	127	(1)
Animals use energy to perform three major functions	128	(2)
Metabolic Rate: Meaning and Measurement	130	(5)
Direct calorimetry: The metabolic rate of an animal can be measured directly	130	(1)
Views on Animal Heat Production	130	(1)
Box 5.2 Units of Measure for Energy and Metabolic Rates	131	(1)
Indirect calorimetry: Animal metabolic rates are usually measured indirectly	131	(1)
Direct Measurement versus Indirect Measurement	131	(3)
Respirometry	134	(1)
Factors That Affect Metabolic Rates	135	(1)
Ingestion of food causes metabolic rate to rise	136	(1)
Basal Metabolic Rate and Standard Metabolic Rate	136	(1)
The Relation between Metabolic Rate and Body Size	137	(7)
Resting metabolic rate is an allometric function of body weight in related species	137	(2)
The metabolic rate of active animals is often also an allometric function of body weight	139	(1)
The metabolic rate--body weight relation has important physiological and ecological implications	140	(2)
The explanation for allometric metabolism--weight relations remains unknown	142	(2)
Energetics of Food and Growth	144	(1)
Conclusion: Energy as the Common Currency of Life	145	(1)
Postscript: The Energy Cost of Mental Effort	145	(4)
Aerobic and Anaerobic Forms of Metabolism	149	(26)
Mechanisms of ATP Production and Their Implications	150	(8)
Aerobic catabolism consists of four major sets of reactions	150	(4)
The Biochemistry of Coupling and Uncoupling of Oxidative Phosphorylation: The Chemiosmotic Theory	154	(1)
O2 deficiency poses two biochemical challenges: Impaired ATP synthesis and possible redox imbalance	155	(1)
Certain tissues possess anaerobic catabolic pathways that synthesize ATP	155	(1)
Anaerobic glycolysis is the principal anaerobic catabolic pathway of vertebrates	155	(1)
What happens to catabolic end products?	156	(1)
The functional roles of ATP-producing mechanisms depend on whether they operate in steady state or nonsteady state	157	(1)
Phosphagens provide an additional mechanism of ATP production without O2	157	(1)
Internal O2 stores may be used to make ATP	158	(1)
Comparative Properties of Mechanisms of ATP Production	158	(2)
Question 1: What is each mechanism's total possible ATP yield per episode of use?	158	(1)
Question 2: How rapidly can ATP production be accelerated?	159	(1)
Question 3: What is each mechanism's peak rate of ATP production (peak power)?	159	(1)
Question 4: How rapidly can each mechanism be reinitialized?	159	(1)
Conclusion: All mechanisms have pros and cons	159	(1)
Genetic Engineering as a Tool to Test Physiological Hypotheses	160	(1)
Two Themes in Exercise Physiology: Fatigue and Muscle Fiber Types	160	(1)
Fatigue has many causes, most of which are poorly understood	160	(1)
The muscle fibers in the muscles used for locomotion are heterogeneous in functional properties	160	(1)
The Interplay of Aerobic and Anaerobic Catabolism during Exercise	161	(5)
Metabolic transitions occur at the start and end of vertebrate exercise	162	(2)
The ATP source for all-out exercise varies in a regular manner with exercise duration	164	(1)
Related species and individuals within one species are sometimes poised very differently for use of aerobic and anaerobic catabolism	165	(1)
Responses to Impaired O2 Influx from the Environment	166	(9)
Diving vertebrates exploit anaerobic glycolysis during protracted dives	167	(1)
Animals faced with reduced O2 availability in their usual environments may show conformity or regulation of aerobic ATP synthesis	168	(1)
Aquatic anaerobes: Some aquatic animals are capable of protracted life without O2	168	(1)
Human Peak O2 Consumption and Physical Performance at High Altitudes	169	(1)
Mechanisms of invertebrate anaerobiosis	169	(1)
Mechanisms of anaerobiosis in goldfish and Crucian carp	170	(5)
The Energetics of Aerobic Activity	175	(16)
How Active Animals Are Studied	176	(2)
The Energy Costs of Defined Exercise	178	(5)
The most advantageous speed depends on the function of exercise	179	(1)
The minimal cost of transport depends in regular ways on mode of locomotion and body size	180	(3)
The Maximal Rate of Oxygen Consumption	183	(4)
Finding Power for Human-Powered Aircraft	183	(1)
VO2max differs among phyletic groups and often from species to species within a phyletic group	184	(1)
VO2max varies among individuals within a species	185	(1)
VO2max responds to training	186	(1)
The Energetics of Routine and Extreme Daily Life	187	(1)
Ecological Energetics	187	(4)
Thermal Relations	191	(50)
Temperature and Heat	193	(1)
Heat Transfer between Animals and Their Environments	194	(4)
Conduction and convection: Convection is intrinsically faster	194	(1)
Evaporation: The change of water from liquid to gas carries much heat away	195	(1)
Thermal radiation permits widely spaced objects to exchange heat	196	(2)
Poikilothermy (Ectothermy)	198	(16)
Poikilotherms often exert behavioral control over their body temperatures	198	(1)
Poikilotherms must be able to function over a range of body temperatures	199	(1)
Poikilotherms respond physiologically to their environments in all three major time frames	200	(1)
Acute responses: Metabolic rate is an approximately exponential function of body temperature	200	(1)
Chronic responses: Acclimation often blunts metabolic responses to temperature	201	(3)
Evolutionary changes: Species are often specialized to live at their respective body temperatures	204	(1)
Temperature and heat matter to animals because they affect the rates of processes and the functional states of molecules	205	(4)
Poikilotherms at high temperatures: Heat-shock proteins help repair damage	209	(1)
Poikilotherms threatened with freezing: They may survive by preventing freezing or tolerating it	210	(4)
Homeothermy in Mammals and Birds	214	(18)
Metabolic rate rises in cold and hot environments because of the costs of homeothermy	214	(1)
The shape of the metabolism--temperature curve depends on fundamental heat-exchange principles	214	(1)
Thermoregulatory Control, Fever, and Behavioral Fever	215	(4)
Homeothermy is metabolically expensive	219	(1)
Insulation is modulated by adjustments of the pelage or plumage, blood flow, and posture	219	(1)
Heat production is increased below thermoneutrality byshivering and nonshivering thermogenesis	220	(1)
Regional heterothermy: In cold environments, allowing some tissues to cool can have advantages	221	(1)
Countercurrent heat exchange permits selective restriction of heat flow to appendages	222	(2)
Mammals and birds in hot environments: Their first lines of defense are often not evaporative	224	(1)
Active evaporative cooling is the ultimate line of defense against overheating	225	(2)
Mammals and birds acclimatize to winter and summer	227	(1)
Evolutionary changes: Species are often specialized to live in their respective climates	228	(1)
Mammals and birds sometimes escape the demands of homeothermy by hibernation, torpor, or related processes	228	(4)
Warm-Bodied Fish	232	(2)
Endothermy and Homeothermy in Insects	234	(7)
The insects that thermoregulate during flight require certain flight-muscle temperatures to fly	235	(1)
Solitary insects employ diverse mechanisms of thermoregulation	235	(1)
Colonies of social bees and wasps often display sophisticated thermoregulation	236	(5)
Food, Energy, and Temperature at Work: The Lives of Mammals in Frigid Places	241	(16)
Food, Nutrition, Energy Metabolism, and Thermoregulation in the Lives of Adult Reindeer	242	(2)
Newborn Reindeer	244	(3)
Knockout Mice Clarify the Function of Brown Fat	245	(2)
Lifetime Patterns of Thermoregulation and Thermogenesis in Small Mammals	247	(1)
The Effect of Body Size on Mammals' Lives in Cold Environments	248	(1)
Hibernation as a Winter Strategy: New Directions and Discoveries	249	(8)
Arctic ground squirrels supercool during hibernation and arouse periodically throughout their hibernation season	250	(1)
The composition of the lipids consumed before hibernation affects the dynamics of hibernation	251	(1)
Although periodic arousals detract from the energy savings of hibernation, their function is unknown	252	(1)
The intersection of sociobiology and physiology: Social hibernation may save energy	253	(4)
PART III Integrating Systems	257	(206)
Neural and Endocrine Control, Nervous Systems, and Biological Clocks	259	(22)
The Physiology of Control: Neurons and Endocrine Cells Compared	260	(3)
Neurons transmit electrical signals to target cells	260	(1)
Endocrine cells broadcast hormones	261	(1)
Nervous systems and endocrine systems tend to control different processes	262	(1)
The Organization and Evolution of Nervous Systems	263	(4)
Nervous systems organize neurons into functional circuits	263	(1)
Multicellular animals have evolved complex nervous systems	264	(3)
The Vertebrate Nervous System: A Guide to the General Organizational Features of Nervous Systems	267	(7)
Nervous systems have central and peripheral divisions	267	(1)
The central nervous system controls physiology and behavior	267	(2)
Four principles of functional organization apply to all mammalian and most vertebrate brains	269	(2)
The peripheral nervous system has somatic and autonomic divisions that control different parts of the body	271	(3)
Biological Clocks	274	(7)
Organisms have endogenous rhythms	275	(1)
Biological clocks generate endogenous rhythms	276	(1)
Control by biological clocks has adaptive advantages	277	(1)
Endogenous clocks correlate with natural history, and compensate for temperature	278	(1)
Clock mechanisms involve rhythms of gene expression	278	(1)
The loci of biological clock functions vary among animals	278	(1)
Circannual and circatidal clocks: Some endogenous clocks time annual or tidal rhythms	279	(1)
Interval or ``hourglass'' timers can time shorter intervals	279	(2)
Neurons	281	(32)
The Cellular Organization of Nervous Systems	282	(2)
Neurons are structurally adapted to transmit action potentials	282	(1)
Glial cells support neurons physically and metabolically	283	(1)
The Ionic Basis of Membrane Potentials	284	(8)
Cell membranes have passive electrical properties: Resistance and capacitance	284	(4)
Resting membrane potentials depend on selective permeability to ions: The Nernst equation	288	(1)
Ion concentration differences result from active ion transport and from passive diffusion	289	(2)
Membrane potentials depend on the permeabilities to and concentration gradients of several ion species: The Goldman equation	291	(1)
Electrogenic pumps also have a small direct effect on Vm	291	(1)
The Action Potential	292	(13)
Action potentials are voltage-dependent, all-or-none electrical signals	292	(1)
Action potentials result from changes in membrane permeabilities to ions	293	(5)
The molecular structure of the voltage-dependent ionchannels reveals their functional properties	298	(2)
There are variations in the ionic mechanisms of excitable cells	300	(1)
The Evolution of Voltage-Gated Channels	301	(4)
Propagation of Action Potentials	305	(8)
Local circuits of current propagate an action potential	305	(1)
Membrane refractory periods prevent bidirectional propagation	306	(1)
The conduction velocity of an action potential depends on axon diameter, myelination, and temperature	306	(2)
Giant Axons	308	(5)
Synapses	313	(34)
Synaptic Transmission Is Usually Chemical but Can Be Electrical	314	(3)
Electrical synapses transmit signals instantaneously	314	(2)
Chemical synapses can condition and amplify signals	316	(1)
Synaptic Potentials Control Neuronal Excitability	317	(2)
Synapses onto a spinal motor neuron exemplify functions of fast synaptic potentials	317	(1)
Synapses excite or inhibit a neuron by depolarization or hyperpolarization at the site of impulse initiation	318	(1)
Fast Chemical Synaptic Actions Depend on Increases in Permeability to lons	319	(4)
Chemical synapses work by releasing and responding to neurotransmitters	319	(1)
Postsynaptic potentials result from permeability changes that are neurotransmitter-dependent and voltage-independent	320	(2)
Neuronal EPSPs resemble neuromuscular EPSPs but are smaller	322	(1)
Fast IPSPs can result from an increase in permeability to chloride	323	(1)
Presynaptic Neurons Release Neurotransmitter Molecules in Quantal Packets	323	(5)
Acetylcholine is synthesized and stored in the presynaptic terminal	323	(1)
Neurotransmitter release is voltage-and Ca2+ -dependent	324	(1)
Neurotransmitter release is quantal and vesicular	324	(2)
Synaptic vesicles are cycled at nerve terminals in distinct steps	326	(1)
Several proteins play roles in vesicular release and recycling	326	(2)
Neurotransmitters Are of Two General Kinds	328	(4)
Neurons have a characteristic neurotransmitter, but they may have more than one neurotransmitter	328	(1)
An agent is identified as a neurotransmitter if it meets five criteria	329	(1)
Vertebrate neurotransmitters have several general modes of action	329	(1)
Neuropeptides and Pain: The Nervous System Produces Natural Opiates	330	(1)
Neurotransmitter systems have been conserved in evolution	331	(1)
Postsynaptic Receptors for Fast lonotropic Actions: Ligand-Gated Channels	332	(2)
ACh receptors are ligand-gated channels that function as ionotropic receptors	332	(2)
Most ligand-gated channel receptors have evolved from a common ancestor	334	(1)
Postsynaptic Receptors for Slow, Metabotropic Actions: G Protein--Coupled Receptors	334	(4)
Metabotropic receptors act via second messengers	334	(1)
The structure of G protein--coupled receptors	335	(1)
G proteins act via intracellular effectors	336	(1)
Permeability-decrease synapses involve G protein-coupled receptors	337	(1)
Synaptic Plasticity: Synapses Change Properties with Time and Activity	338	(9)
Neurotransmitter metabolism is regulated homeostatically	338	(1)
Learning and memory may be based on synaptic plasticity	338	(9)
Sensory Processes	347	(42)
Organization of Sensory Systems	348	(2)
Receptors are classified by sensory modality, location, or form of stimulus energy	348	(1)
Sensory reception comprises a series of discrete operations	349	(1)
Receptor Functions and Their Control	350	(5)
Receptor potentials result from the influx of Na+ ions	351	(1)
Receptors adapt to sustained stimulation	351	(2)
Receptors encode information about modality, intensity, location, and timing of a stimulus	353	(1)
Efferent control can adjust receptor sensitivity	354	(1)
Photoreception	355	(6)
Photoreceptors and eyes have distinct evolutionary relationships	355	(1)
The vertebrate eye focuses light onto retinal rods and cones	356	(1)
Retinal rods and cones transduce light into a hyperpolarizing receptor potential	357	(4)
Visual Sensory Processing	361	(7)
Retinal neurons respond to patterns of light and dark	361	(4)
The vertebrate brain integrates visual information through parallel pathways	365	(3)
Arthropod Visual Systems	368	(1)
Arthropod eyes are compound eyes	368	(1)
Arthropod photoreceptors are depolarized by light	369	(1)
Mechanoreception	369	(10)
Proprioceptors monitor the spatial relationship of the body	369	(1)
Equilibrium receptors detect gravity and acceleration to maintain balance	370	(3)
Auditory receptors transduce sound into electrical signals	373	(5)
Echolocation	378	(1)
Chemoreception	379	(7)
Insect contact chemoreceptors are localized in sensilla	379	(1)
Insect olfactory receptors sense pheromones and other chemicals	380	(1)
Vertebrate taste involves four or five taste qualities	381	(1)
Vertebrate olfaction includes over a thousand receptor types	382	(4)
Electroreception	386	(3)
Endocrine and Neuroendocrine Physiology	389	(34)
Introduction to Endocrine Principles	390	(6)
Hormones bind to receptor molecules expressed by target cells	390	(1)
Concentrations of hormones in the blood vary	390	(3)
Most hormones fall into three chemical classes	393	(2)
Hormone molecules exert their effects by producing biochemical changes in target cells	395	(1)
Synthesis, Storage, and Release of Hormones	396	(2)
Peptide hormones are synthesized at ribosomes, stored in vesicles, and secreted on demand	396	(1)
Steroid hormones are synthesized from cholesterol, are not stored, and are secreted by diffusion	397	(1)
Types of Endocrine Glands and Cells	398	(1)
Control of Endocrine Systems: The Vertebrate Pituitary Gland	399	(4)
The posterior pituitary illustrates neural control of neurosecretory cells	399	(1)
The anterior pituitary illustrates neurosecretory control of endocrine cells	399	(2)
Hormones and neural input modulate endocrine control pathways	401	(2)
The Mammalian Stress Response	403	(4)
The autonomic nervous system and HPA axis coordinate the stress response to an acute threat	404	(1)
The HPA axis modulates the immune system	405	(1)
Chronic stress causes deleterious effects	406	(1)
Plasma glucocorticoid concentrations show seasonal variations	406	(1)
Endocrine Control of Nutrient Metabolism in Mammals	407	(3)
Insulin regulates short-term changes in nutrient availability	408	(1)
Glucagon works together with insulin to ensure stable levels of glucose in the blood	408	(1)
Other hormones contribute to the regulation of nutrient metabolism	409	(1)
Endocrine Control of Salt and Water Balance in Vertebrates	410	(2)
Antidiuretic hormones conserve water	410	(1)
The renin-angiotensin-aldosterone system conserves sodium	411	(1)
Atrial natriuretic peptide promotes excretion of sodium and water	412	(1)
Hormones and Other Chemical Signals	412	(4)
Can Mating Cause True Commitment?	413	(1)
Hormones and neurohormones fall along a ``distance continuum'' of chemical signals	413	(2)
Paracrines and autocrines are local chemical signals distributed by diffusion	415	(1)
Pheromones and kairomones are used as chemical signals between animals	415	(1)
Insect Metamorphosis	416	(7)
Insects in Forensics and Medicine	416	(1)
Insect metamorphosis may be gradual or dramatic	416	(1)
Hormones and neurohormones control insect metamorphosis	417	(6)
Reproduction	423	(24)
Sexual and Asexual Reproduction	423	(3)
Mammalian Reproduction	426	(3)
Gametes are produced in ovaries and testes	426	(2)
``Male Menopause'' in a Marsupial Mouse	428	(1)
Oocytes mature periodically in menstrual cycles or estrous cycles	428	(1)
Hormonal Control of Female Reproduction	429	(6)
Dynamic cellular changes occur in the ovary over the course of a cycle	429	(2)
Hormones influence development of the follicle	431	(1)
Estrogen influences target tissues outside the ovary	432	(1)
Ovulation is controlled by LH	433	(1)
The corpus luteum is essential for establishing and maintaining pregnancy	433	(1)
Why do all mammals cyclically change the endometrium? Why do humans and a few other primates menstruate?	434	(1)
Hormonal Control of Male Reproduction	435	(3)
Sex Determination in Mammals	436	(2)
Fertilization, Pregnancy, and Birth in Eutherian Mammals	438	(5)
Fertilization is the union of sperm and oocyte	438	(1)
Development begins in the oviduct, and implantation establishes pregnancy	439	(2)
The embryonic trophoblast and the maternal endometrium form the placenta	441	(1)
Parturition requires cellular changes and orchestrated neural and hormonal signals	442	(1)
Lactation	443	(1)
Maximizing Reproductive Success	444	(3)
Integrating Systems at Work: Animal Navigation	447	(16)
The Adaptive Significance of Animal Navigation	448	(2)
Navigational abilities promote reproductive success	448	(1)
Navigational abilities facilitate food acquisition	449	(1)
Migrating animals need navigation	449	(1)
Navigational Strategies	450	(8)
Trail following is the most rudimentary form of animal navigation	450	(1)
Piloting animals follow a discontinuous series of learned cues	450	(1)
Path integration is a form of dead reckoning	451	(1)
Animals can derive compass information from environmental cues	452	(5)
Some animals appear to possess a map sense	457	(1)
Sea turtles exemplify the degree of our understanding of navigation	458	(1)
Innate and Learned Components of Navigation	458	(5)
Some forms of navigation have strong innate aspects	459	(1)
The hippocampus is a critical brain area for vertebrate spatial learning and memory	459	(4)
PART IV Muscle and Movement	463	(62)
Muscle	465	(24)
Vertebrate Skeletal Muscle Cells	466	(4)
Thick and thin filaments are polarized polymers of individual protein molecules	467	(1)
Muscles require ATP to contract	468	(1)
Calcium and the regulatory proteins tropomyosin and troponin control contractions	469	(1)
Excitation--Contraction Coupling	470	(3)
Whole Skeletal Muscles	473	(5)
Muscle contraction is the force generated by a muscle during cross-bridge activity	473	(1)
A twitch is the mechanical response of a muscle to a single action potential	473	(1)
The velocity of shortening decreases as the load increases	473	(1)
The frequency of action potentials determines the tension developed by a muscle	474	(1)
Sustained calcium in the cytoplasm permits summation and tetanus	474	(1)
The amount of tension developed by a muscle depends on the length of the muscle at the time it is stimulated	474	(2)
In general, the amount of work a muscle can do depends on its volume	476	(1)
Electric Fish Exploit Modified Skeletal Muscles to Generate Electric Shocks	477	(1)
Muscle Energetics	478	(4)
ATP is the immediate source of energy for powering muscle contraction	478	(1)
Vertebrate muscle fibers are classified into different types	479	(2)
Insect Flight	481	(1)
Neural Control of Skeletal Muscle	482	(2)
The vertebrate plan is based on muscles organized into motor units	482	(1)
The innervation of vertebrate tonic muscle is intermediate between the general vertebrate and arthropod plans	482	(1)
The arthropod plan is based on multiterminal innervation of each muscle fiber by more than one neuron	483	(1)
Vertebrate Smooth Muscle	484	(2)
Vertebrate Cardiac Muscle	486	(3)
Control of Movement: The Motor Bases of Animal Behavior	489	(22)
Behavioral Background: Reflexes and Fixed Action Patterns	489	(1)
Neural Circuits Mediating Reflexes and Fixed Acts	490	(6)
Crayfish escape behavior is a fixed act mediated by a giant interneuron	490	(1)
Gill withdrawal in Aplysia is a reflex act	491	(1)
Vertebrate spinal reflexes compensate for circumstances, as well as initiating movements	492	(2)
Motor neurons are activated primarily by central input rather than by spinal reflexes	494	(2)
Action Patterns: Neural Generation of Rhythmic Behavior	496	(6)
Locust flight results from an interplay of central and peripheral control	497	(1)
Command neurons turn on the pattern generator	498	(1)
There are different mechanisms of central pattern generation	498	(3)
Central pattern generators can underlie relatively complex behavior	501	(1)
Control and Coordination of Vertebrate Movement	502	(9)
Locomotion in cats involves a spinal central pattern generator	502	(1)
The generation of movement involves several areas in the mammalian brain	503	(8)
Muscle and Movement at Work: Muscle in Human Health and Disease	511	(14)
Exercise	512	(5)
Power output determines a muscle's contractile performance	512	(1)
Endurance training elicits changes in fiber type, capillary density, and mitochondrial density	513	(3)
Resistance training causes hypertrophy and changes in fiber type	516	(1)
Stretch and force production activate protein synthesis in rabbit muscles	517	(1)
Atrophy	517	(4)
Humans experience atrophy in microgravity	517	(1)
A variety of techniques are used to study disuse atrophy in small mammals	518	(1)
Disuse influences the fiber-type composition of muscles	519	(1)
Muscles atrophy with age	519	(2)
No Time to Lose	521	(1)
Some animals experience little or no disuse atrophy	521	(1)
Muscle Disease	521	(4)
DMD results from mutation of the DMD gene on the short arm of the X chromosome	522	(1)
Dystrophin Connects F-Action To The Sarcolemma	522	(3)
PART V Oxygen, Carbon Dioxide, and Internal Transport	525	(138)
Introduction to Oxygen and Carbon Dioxide Physiology	527	(14)
The Properties of Gases in Gas Phases and Aqueous Solutions	528	(2)
Gases in the gas phase	528	(1)
Gases in solution	529	(1)
Diffusion of Gases	530	(2)
Gases diffuse far more readily through gas phases than through aqueous solutions	531	(1)
Gas molecules that combine chemically with other molecules cease to contribute to the gas partial pressure	531	(1)
Over What Distance Can Diffusion Meet the O2 Requirements of Tissues?	532	(1)
Convective Transport of Gases	532	(3)
Induction of Internal Flow by Ambient Currents	533	(1)
Gas transport in animals often occurs by alternating convection and diffusion	534	(1)
The Oxygen Cascade	535	(1)
Expressing the Amounts and Partial Pressures of Gases in Other Units	536	(1)
The Contrasting Physical Properties of Air and Water	537	(1)
Respiratory Environments	537	(4)
External Respiration: The Physiology of Breathing	541	(36)
Fundamental Concepts of External Respiration	542	(2)
Principles of Gas Exchange by Active Ventilation	544	(2)
The O2 partial pressure in blood leaving a breathing organ depends on the relation between the flow of the blood and the flow of the air or water	544	(1)
The relative changes in the partial pressures of O2 and CO2 depend on whether air or water is breathed	545	(1)
Introduction to Vertebrate Breathing	546	(3)
Breathing by Fish	549	(4)
Gill ventilation is usually driven by buccal--opercular pumping	550	(2)
Many fish use ram ventilation on occasion and some use it all the time	552	(1)
Decreased O2 and exercise are the major stimuli for increased ventilation in fish	552	(1)
Several hundred species of bony fish are able to breathe air	552	(1)
Breathing by Amphibians	553	(2)
Gills, lungs, and skin are used in various combinations to achieve gas exchange	555	(1)
Breathing by Reptiles	555	(1)
Breathing by Mammals	556	(8)
The total lung volume is employed in different ways in different sorts of breathing	557	(1)
The gas in the final airways differs from atmospheric air in composition and is motionless	558	(1)
The power for ventilation is developed by the diaphragm and the intercostal and abdominal muscles	559	(1)
The control of ventilation	559	(2)
Mammals at High Altitude	561	(2)
In species of different sizes, lung volume tends to be a constant proportion of body size but breathing frequency varies allometrically	563	(1)
Pulmonary surfactant keeps the alveoli from collapsing	563	(1)
Breathing by Birds	564	(3)
Ventilation is by bellows action	566	(1)
Air flows unidirectionally through the parabronchi	566	(1)
The gas-exchange system is cross-current	566	(1)
Bird Development: Filling the Lungs with Air for Hatching	567	(1)
Breathing by Aquatic Invertebrates and Allied Groups	567	(3)
Molluscs exemplify an exceptional diversity of breathing organs built on a common plan	567	(2)
Decapod crustaceans include many important water breathers and some air breathers	569	(1)
Breathing by Insects and Other Tracheate Arthropods	570	(7)
Diffusion is important in gas transport through the tracheal system	571	(1)
Some insects employ conspicuous ventilation	572	(1)
Microscopic ventilation is far more common than believed even a decade ago	572	(1)
Control of breathing	573	(1)
Some insects have gills or lungs formed by tracheae	573	(1)
Many aquatic insects breathe when under water through spiracles using external gas spaces	573	(1)
The Book Lungs of Arachnids	574	(3)
Transport of Oxygen and Carbon Dioxide in Body Fluids (With an Introduction to Acid-Base Physiology)	577	(32)
Absorption Spectra of Respiratory Pigments	578	(1)
The Chemical Properties and Distributions of the Respiratory Pigments	579	(4)
Hemoglobins contain heme and are the most widespread respiratory pigments	579	(3)
Blood Cells and Their Production	582	(1)
Copper-based hemocyanins occur in many arthropods and molluscs	582	(1)
Chlorocruorins resemble hemoglobins and occur in certain annelids	583	(1)
Iron-based hemerythrins do not contain heme and occur in four phyla	583	(1)
The O2-Binding Characteristics of Respiratory Pigments	583	(10)
Human O2 transport provides an instructive case study	584	(3)
A set of general principles helps elucidate O2 transport by respiratory pigments	587	(1)
The shape of the oxygen equilibrium curve depends on O2-binding site cooperativity	587	(2)
Respiratory pigments exhibit a wide range in their affinities for O2	589	(1)
The Bohr effect: Oxygen affinity depends on the partial pressure of CO2 and the pH	589	(2)
The Root effect: In unusual cases, CO2 and pH affect the oxygen-carrying capacity of the respiratory pigment	591	(1)
Thermal effects: Oxygen affinity depends on tissue temperature	591	(1)
Organic modulators often exert chronic effects on oxygen affinity	591	(1)
Inorganic ions may also act as modulators of respiratory pigments	592	(1)
The Functions of Respiratory Pigments in Animals	593	(6)
Patterns of circulatory O2 transport: The mammalian model is common but not universal	594	(1)
Respiratory pigments often display differences in O2 affinity that aid successful O2 transport	595	(1)
Evolved differences in affinity among related species	596	(1)
The respiratory-pigment physiology of individuals undergoes acclimation and acclimatization	596	(1)
Blood and Circulation in Mammals at High Altitude	597	(1)
Icefish live without hemoglobin	598	(1)
Carbon Dioxide Transport	599	(4)
The extent of bicarbonate formation depends on blood buffers	599	(1)
Carbon dioxide transport is interpreted by use of carbon dioxide equilibrium curves	600	(1)
The Haldane effect: The carbon dioxide equilibrium curve depends on blood oxygenation	601	(1)
Critical details of vertebrate CO2 transport depend on carbonic anhydrase and anion transporters	602	(1)
Acid-Base Physiology	603	(6)
Acid-base regulation involves excretion or retention of chemical forms affecting H+ concentration	604	(1)
Disturbances of acid-base regulation fall into respiratory and metabolic categories	604	(5)
Circulation	609	(34)
Hearts	610	(7)
The heart as a pump: The action of a heart can be analyzed in terms of the physics of pumping	611	(1)
The circulation must deliver O2 to the myocardium	612	(1)
The electrical impulses for heart contraction may originate in muscle cells or neurons	613	(2)
A heart produces an electrical signature, the electrocardiogram	615	(1)
Heart action is modulated by hormonal, nervous, and intrinsic controls	616	(1)
Principles of Pressure, Resistance, and Flow in Vascular Systems	617	(3)
The rate of blood flow depends on differences in blood pressure and on vascular resistance	619	(1)
The dissipation of energy: Pressure and flow turn to heat during circulation of the blood	620	(1)
Circulation in Mammals and Birds	620	(6)
The circulatory system is closed	620	(1)
Each part of the systemic vascular system has distinctive anatomical and functional features	621	(2)
Mammals and birds have a high-pressure systemic circuit	623	(1)
Fluid undergoes complex patterns of exchange across the walls of systemic capillaries	624	(1)
The pulmonary circuit is a comparatively low-pressure system	625	(1)
During exercise, blood flow is increased by orchestrated changes in cardiac output and vascular resistance	625	(1)
Species have evolved differences in their circulatory physiology	626	(1)
Circulation in Fish	626	(5)
The circulatory plans of air-breathing organs (ABOs) in fish pose unresolved questions	628	(1)
Lungfish have specializations to promote separation of oxygenated and deoxygenated blood	629	(2)
An Incompletely Divided Central Circulation Can Be an Advantage for Intermittent Breathers	631	(1)
Circulation in Amphibians and Reptiles	631	(1)
Concluding Comments on Vertebrates	632	(1)
Invertebrates with Closed Circulatory Systems	633	(1)
Bearing the Burden of Athleticism, Sort Of: A Synthesis of Cephalopod O2 Transport	634	(1)
Invertebrates with Open Circulatory Systems	634	(9)
The crustacean circulatory system provides an example of an open system	634	(3)
Circulation and O2: Lessons from the Insect World	637	(1)
Open systems are functionally different from closed systems but may be equal in critical ways	637	(6)
Oxygen, Carbon Dioxide, and Internal Transport at Work: Diving by Marine Mammals	643	(20)
Diving Feats and Behavior	643	(3)
Types of Dives and the Importance of Method	646	(1)
Physiology: The Big Picture	646	(1)
The Oxygen Stores of Divers	647	(4)
The blood O2 store tends to be large in diving mammals	647	(1)
Diving mammals have high myoglobin concentrations and large myoglobin-bound O2 stores	648	(1)
Diving mammals vary in their use of the lungs as an O2 store	648	(1)
Total O2 stores never permit dives of maximal duration to be fully aerobic	649	(2)
Circulatory Adjustments during Dives	651	(3)
Regional vasoconstriction: Much of a diving mammal's body is cut off from blood flow during forced or protracted dives	651	(1)
Diving bradycardia matches cardiac output to the circulatory task	652	(1)
Cardiovascular responses are graded in freely diving animals	652	(1)
The Evolution of Vertebrate Cardiac and Vascular Responses to Asphyxia	653	(1)
Red blood cells are removed from the blood between dive sequences in some seals	654	(1)
Metabolism during Dives	654	(2)
The body becomes metabolically subdivided during forced or protracted dives	654	(1)
Metabolic limits on dive duration are determined by O2 supplies, rates of O2 use and lactic acid accumulation, and tissue tolerances	654	(2)
The Aerobic Dive Limit: Physiology's Benchmark for Understanding Diving Behavior	656	(2)
Decompression Sickness	658	(2)
Human decompression sickness is usually caused by N2 absorption from a compressed-air source	658	(1)
Breath-hold dives must be repeated many times to cause decompression sickness in humans	659	(1)
Marine mammals avoid decompression sickness during deep dives by alveolar collapse	659	(1)
Dives too shallow for alveolar collapse present special problems	659	(1)
A possible advantage for pulmonary O2 sequestration in deep dives	660	(3)
PART VI Water, Salts, and Excretion	663	(2)
Water and Salt Physiology: Introduction and Mechanisms	665	(20)
The Importance of Animal Body Fluids	666	(1)
The Relations among Body Fluids	667	(1)
The Types of Regulation and Conformity	668	(1)
Natural Aquatic Environments	669	(2)
Natural Terrestrial Environments	671	(2)
Organs of Blood Regulation	673	(2)
The effects of kidney function on osmotic regulation depend on the osmotic U/P ratio	674	(1)
The effects of kidney function on volume regulation depend on the amount of urine produced	675	(1)
The effects of kidney function on ionic regulation depend on ionic U/P ratios	675	(1)
Food and Drinking Water	675	(1)
Salty drinking water may not provide H2O	676	(1)
Plants and algae with salty tissue fluids pose challenges for herbivores	676	(1)
Air-dried foods contain water in variable amounts	676	(1)
Protein-rich foods can be dehydrating for terrestrial animals	676	(1)
Metabolic Water	676	(2)
Metabolic water matters most in animals that conserve water effectively	677	(1)
Net Metabolic Water Gain in Kangaroo Rats	678	(1)
The Water and Salt Physiology of Tissue Cells	678	(3)
Most animals employ organic solutes for cell-volume regulation	679	(1)
Organic solutes have played major roles in the adjustment of intracellular osmotic pressure over evolutionary time	680	(1)
From Osmolytes to Compatible Solutes: Terms and Concepts	681	(4)
Water and Salt Physiology of Animals in Their Environments	685	(36)
Animals in Freshwater	686	(5)
Passive water and ion exchanges: Freshwater animals tend to gain water by osmosis and lose major ions by diffusion	686	(2)
Most types of freshwater animals share similar regulatory mechanisms	688	(2)
A few types of freshwater animals exhibit exceptional patterns of regulation	690	(1)
Why do most freshwater animals make dilute urine?	691	(1)
Animals in the Ocean	691	(8)
Most marine invertebrates are isosmotic to seawater	691	(1)
Hagfish are the only vertebrates with blood inorganic ion concentrations that make them isosmotic to seawater	692	(1)
The marine teleost fish are markedly hyposmotic to seawater	692	(1)
Where Were Vertebrates at Their Start?	692	(3)
Epithelial NaCI Secretion	695	(1)
Some arthropods of saline waters are hyposmotic regulators	696	(1)
Marine reptiles, birds, and mammals are also hyposmotic regulators	696	(1)
Marine elasmobranch fish are hyperosmotic but hypoinonic to seawater	697	(1)
The Evolution of Urea Synthesis in Vertebrates	698	(1)
Animals that Face Changes in Salinity	699	(3)
The migratory fish are dramatic examples of hyperhyposmotic regulators	700	(1)
Animals undergo change in all time frames in their relations to ambient salinity	701	(1)
Responses to Drying of the Habitat in Aquatic Animals	702	(1)
Life as Nothing More than a Morphological State	703	(1)
Animals on Land: Fundamental Physiological Principles	703	(7)
A low integumentary permeability to water is a key to reducing evaporative water loss on land	704	(1)
Respiratory evaporative water loss depends on the function of the breathing organs and the rate of metabolism	704	(3)
The total rate of evaporative water loss depends on an animal's body size and phylogenetic group	707	(1)
Excretory water loss depends on the concentrating ability of the excretory organs and the amount of solute that needs to be excreted	707	(2)
Terrestrial animals sometimes enter dormancy or tolerate wide departures from homeostasis to cope with water stress	709	(1)
The total rates of water turnover of free-living terrestrial animals follow allometric patterns	709	(1)
Animals on Land: Case Studies	710	(5)
Amphibians occupy a diversity of habitats despite their meager physiological abilities to limit water losses	710	(2)
Xeric invertebrates: Because of exquisite water conservation, some insects and arachnids have only small water needs	712	(1)
Xeric vertebrates: Studies of lizards and small mammals help clarify the complexities of desert existence	712	(1)
The Study of Physiological Evolution by Artificial Selection	713	(2)
Control of Water and Salt Balance in Terrestrial Animals	715	(6)
Kidneys and Excretion (With Notes on Nitrogen Excretion)	721	(36)
Basic Mechanisms of Kidney Function	722	(3)
Primary urine is introduced into kidney tubules by ultrafiltration or secretion	722	(2)
Primary urine is usually modified extensively before it is excreted as definitive urine	724	(1)
Urine Formation in Amphibians	725	(5)
The proximal convoluted tubule reabsorbs much of the filtrate without changing osmotic pressure	726	(1)
Quantity versus Concentration	727	(1)
The distal convoluted tubule can differentially reabsorb water and solutes, thus controlling water excretion	727	(1)
Renal Clearance and Other Methods of Study of Kidney Function	728	(1)
ADH exerts an elaborate pattern of control over nephron function	729	(1)
The bladder functions in urine formation in amphibians	729	(1)
The amphibian excretory system has mechanisms to promote excretion of urea	730	(1)
Urine Formation in Mammals	730	(14)
The nephrons, singly and collectively, give the mammalian kidney a distinctive structure	730	(2)
Comparative anatomy points to a role for the loops of Henle in concentrating the urine	732	(1)
Countercurrent multiplication is the key to producing concentrated urine	733	(3)
Countercurrent Multipliers versus Countercurrent Exchangers	736	(4)
The kidney tubules carry out many processes to produce definitive urine	740	(4)
Urine Formation in Other Vertebrates	744	(1)
Freshwater and marine teleost fish differ in nephron structure and function	744	(1)
Reptiles have nephrons like those of amphibians, but birds have some mammalian-type nephrons	744	(1)
Urine Formation in Decapod Crustaceans	745	(1)
Urine Formation in Molluscs	746	(1)
Urine Formation in Insects	747	(3)
The Malpighian tubules form and sometimes modify the primary urine	747	(1)
The hindgut modulates urine composition, concentration, and volume	748	(2)
Nitrogen Disposition and Excretion	750	(7)
Ammonotelism is the primitive state	751	(1)
Urea is more costly to synthesize but less toxic than ammonia	751	(1)
Uric acid and related compounds remove nitrogen from solution	752	(1)
Why Are Mammals Not Uricotelic?	753	(4)
Water, Salts, and Excretion at Work: Mammals of Deserts and Dry Savannas	757	(1)
Desert and Dry-Savanna Environments	757	(1)
The Relations of Animals to Water	758	(7)
In terms of water costs, large body size is a physiological advantage	758	(1)
Coexisting species are diverse in their relations to drinking water	759	(2)
Water conflicts threaten animals and people	761	(2)
All species of large herbivores require considerable amounts of preformed water	763	(1)
Water and food resources in the deserts and dry savannas are often complex	764	(1)
The Dramatic Adaptations of Particular Species	765	(1)
Oryxes represent the pinnacle of desert survival	766	(2)
Grant's and Thomson's gazelles differ in their relations to water	768	(1)
The dromedary camel does not store water, but conserves it and tolerates profound dehydration	768
Appendix A References	1	(2)
Appendix B The Systeme International and Other Units of Measure	3	(1)
Appendix C Prefixes Indicating Orders of Magnitude	4	(1)
Appendix D Gases at Standard Temperature and Pressure	5	(1)
Appendix E Fitting Lines to Data	6	(2)
Appendix F Logarithms	8	(2)
Appendix G Exponential and Allometric Equations	10	(2)
Appendix H Mitosis and Meiosis	12	(3)
Appendix I The Standard Amino Acids	15	(1)
Appendix J Basic Physics Terms	16
Glossary	1	(1)
Photo Credits	1	(1)
Figure and Table Citations	1	(1)
Index	1