P r i n c i p l e s o f A n i m a l P h y s i o l o g y M o y e s S c h u l t e 2 e Principles of Animal Physiology Christopher D. Moyes ISBN 978-1-29202-638-1 Patricia M. Schulte Second Edition 9 781292 026381 Principles of Animal Physiology Christopher D. Moyes Patricia M. Schulte Second Edition ISBN 10: 1-292-02638-3 ISBN 13: 978-1-292-02638-1 Pearson Education Limited Edinburgh Gate Harlow Essex CM20 2JE England and Associated Companies throughout the world Visit us on the World Wide Web at: www.pearsoned.co.uk © Pearson Education Limited 2014 All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, without either the prior written permission of the publisher or a licence permitting restricted copying in the United Kingdom issued by the Copyright Licensing Agency Ltd, Saffron House, 6–10 Kirby Street, London EC1N 8TS. All trademarks used herein are the property of their respective owners. The use of any trademark in this text does not vest in the author or publisher any trademark ownership rights in such trademarks, nor does the use of such trademarks imply any affi liation with or endorsement of this book by such owners. ISBN 10: 1-292-02638-3 ISBN 13: 978-1-292-02638-1 British Library Cataloguing-in-Publication Data A catalogue record for this book is available from the British Library Printed in the United States of America 11223345562416273740501646824460086 P E A R S O N C U S T O M L I B R AR Y Table of Contents Glossary Christopher D. Moyes/Patricia M. Schulte 1 1. Introduction to Physiological Principles Christopher D. Moyes/Patricia M. Schulte 26 2. Chemistry, Biochemistry, and Cell Physiology Christopher D. Moyes/Patricia M. Schulte 44 3. Cell Signaling and Endocrine Regulation Christopher D. Moyes/Patricia M. Schulte 116 4. Neuron Structure and Function Christopher D. Moyes/Patricia M. Schulte 168 5. Cellular Movement and Muscles Christopher D. Moyes/Patricia M. Schulte 222 6. Sensory Systems Christopher D. Moyes/Patricia M. Schulte 274 7. Functional Organization of Nervous Systems Christopher D. Moyes/Patricia M. Schulte 334 8. Circulatory Systems Christopher D. Moyes/Patricia M. Schulte 376 9. Respiratory Systems Christopher D. Moyes/Patricia M. Schulte 440 10. Ion and Water Balance Christopher D. Moyes/Patricia M. Schulte 500 11. Digestion Christopher D. Moyes/Patricia M. Schulte 558 12. Locomotion Christopher D. Moyes/Patricia M. Schulte 606 I 677777603344007903 13. Thermal Physiology Christopher D. Moyes/Patricia M. Schulte 660 14. Reproduction Christopher D. Moyes/Patricia M. Schulte 700 Appendix: The International System of Units Christopher D. Moyes/Patricia M. Schulte 737 Appendix: Logarithms Christopher D. Moyes/Patricia M. Schulte 739 Appendix: Linear, Exponential, Power, and Allometric Functions Christopher D. Moyes/Patricia M. Schulte 740 Index 743 II Glossary action potential A relatively large- adrenal cortex See adrenal gland. amplitude, rapid change in the adrenal gland A gland near the membrane potential of an excitable kidney, which in mammals is cell as a result of the opening and composed of an outermost layer (the closing of voltage-gated ion channels; adrenal cortex) and an inner layer A-band (or anisotropic band) The involved in transmitting signals (adrenal medulla). region of a muscle sarcomere where across long distances in the nervous adrenal medulla See adrenal gland. the thick filaments occur. system. adrenergic receptors Receptors for absolute refractory period The period activation energy (E ) The energetic the catecholamines norepinephrine during and immediately following an a barrier that must be reached before a and epinephrine. action potential in which an excitable reactant can be transformed into a adrenoreceptors See adrenergic cell cannot generate another action product. receptors. potential, no matter how strong the activation gate One of the two gates aerobic Occurring in, or depending on, stimulus. that open and close voltage-gated the presence of oxygen. absolute temperature A measure of sodium channels (see also aerobic scope The ratio of the temperature in kelvins, where 0 K inactivation gate). maximal aerobic metabolic rate to (absolute zero) is the temperature at active site A region of an enzyme that the basal metabolic rate, typically in which there is no atomic or binds the substrate and undergoes the range of 3–10. molecular movement. 1 unit on the conformational changes to catalyze afferent Leading toward a region of Kelvin scale equals 1°on the Celsius scale. 0 K (cid:2)(cid:3)273°C. the reaction. interest (see also efferent). active state The phase of a cross- afferent neuron A neuron that acclimation A persistent but reversible bridge cycle in which myosin is conducts a signal from the periphery change in a physiological function attached to actin and generating to an integrating center (see also that occurs as a result of an force. sensory neuron). alteration in an environmental active transport Protein-mediated affinity A measure of the degree of parameter, such as temperature or movement of a substance across a attraction between a ligand and a photoperiod. Acclimation usually membrane with the utilization of molecule that binds the ligand (see occurs as a result of an experimental some form of energy. Primary active also K ). manipulation (see also m transport uses ATP. Secondary active affinity constant (or K)Reciprocal of acclimatization). a transport uses an electrochemical the dissociation constant. acclimatization A reorganization of gradient (see also facilitated after-hyperpolarization A prolonged physiological functions that occurs as diffusion, passive transport). hyperpolarization following an action a result of complex environmental acuity, sensory The ability to resolve potential. changes, such as season or altitude fine detail of a stimulus. aglomerular kidney A derived form of (see also acclimation). acute response The rapid phase of kidney, with tubules that lack a accommodation The process by which response to an external or internal glomerulus, found in many lineages an eye changes its focal length. change in conditions, usually within of marine fish. Accommodation allows the eye to seconds to minutes. agonist A substance that binds to a produce a focused image of objects at adaptation Used in two contexts in receptor and initiates a signaling different distances. physiology: (1) a change in the event. May include both the natural acetyl CoA An activated form of genetic structure of a population as a endogenous ligand as well as acetate that serves as the entry point result of natural selection; (2) a pharmaceutical agents that mimic the for carbon into the TCA cycle. reversible change in a physiological natural substance. acetylcholine A neurotransmitter found parameter that provides a beneficial albumen A protein found in eggs that in most animal species in many types response to an environmental cushions the embryo. of neurons, including motor neurons change. Evolutionary and albumin A binding globulin (carrier and the autonomic ganglia of comparative physiologists prefer to protein) that is one of the primary vertebrates. use only the first definition. proteins of vertebrate plasma; makes acetylcholinesterase An enzyme that adaptation, sensory See receptor a major contribution to blood osmotic catalyzes the breakdown of adaptation. pressure. acetylcholine into choline and adenine A purine nitrogenous base aldosterone Mineralocorticoid acetate. component of nucleotides, including hormone secreted by the adrenal acid A chemical that donates a proton nucleic acids. cortex. Its main function is to alter (see also base). adenosine A nucleoside composed of the levels of Na(cid:6)and K(cid:6)in the urine, acidosis A decrease in pH arising adenine and the sugar deoxyribose, secondarily affecting water transport. through respiration (respiratory important as a signaling molecule. alkaloids A large group of compounds acidosis) or metabolism (metabolic adenosine diphosphate (ADP) A derived from plants that have acidosis). nucleotide composed of the pharmacological effects in animals. acrosomal reaction The exocytosis of nucleoside adenine with two alkalosis The condition of being the enzyme-laden acrosomal vesicle phosphate groups, with a single high- alkaline (see also metabolic alkalosis, of sperm in response to contact with energy phosphodiester bond. respiratory alkalosis). the ovum. adenosine triphosphate (ATP) A allantoic membrane One of four acrosome A vesicle in sperm that nucleotide composed of the membranes in an amniote egg. contains digestive enzymes that nucleoside adenine with three allantoin An intermediate in enable the sperm to penetrate the phosphate groups, with two high- nucleotide breakdown and uric acid outer layers of an ovum. energy phosphodiester bonds. synthesis; an important form of actin G-actin is a monomeric protein adenylate cyclase (adenylyl cyclase) nitrogenous waste for some animals. that can be polymerized to construct The enzyme that converts ATP to allatostatin A neuropeptide hormone filamentous actin (F-actin). Actin is cyclic AMP. in arthropods that inhibits the corpus the basis of both cytoskeletal adhesion plaque A membrane protein allatum from secreting juvenile microfilaments (composed of the (cid:4)- complex that anchors thin filaments hormone. actin isoform of G-actin) and skeletal to the membrane. allatotropin A neuropeptide hormone thin filaments (composed of the (cid:5)- adipose tissue A tissue composed of in arthropods that stimulates the actin isoform of G-actin) (see also fat cells (adipocytes) that produce corpus allatum to secrete juvenile myosin). and store lipid. hormone. actinomyosin The combination of actin ADP See adenosine diphosphate. alleles Different forms of the same and myosin, joined by a cross-bridge. protein that are encoded by the same From Principles of Animal Physiology, Second Edition. Christopher D. Moyes, Patricia M. Schulte. Copyright © 2008 by Pearson Education, Inc. Published by Pearson Benjamin Cummings. All rights reserved. 1 GLOSSARY gene but differ slightly in primary amylase An enzyme that breaks down anus The sphincter through which sequence. starch (amylose, amylopectin). feces exit the gastrointestinal tract. allometry (or allometric scaling) The anabolic pathways (or anabolism) aorta The major artery exiting the pattern seen when comparing Metabolic reactions or pathways that heart. structural or functional parameters in build complex molecules from aortic body A sensory structure relation to body size. simpler molecules. located in the vertebrate aorta that allosteric regulator A molecule that anadromous The life history strategy contains baroreceptors and binds an enzyme at a site distinct of an animal living most of its life in chemoreceptors. from the substrate binding site to the sea, then returning to apical The end of a structure opposite regulate activity. freshwater to reproduce (see also the base. allosteric site A region of an enzyme, catadromous). apical membrane The end of the cell distinct from the active site, that anaerobic Without oxygen. Pertains to furthest from the basolateral binds a molecule other than the an environment without oxygen, or a membrane; the membrane oriented substrate or product, triggering a pathway that occurs in the absence of away from the circulatory system. structural change that alters the oxygen (see also aerobic). apnea A period without breathing. catalytic properties of the enzyme. anaplerotic pathway (or anaplerosis) apocrine A type of secretion whereby allozyme An allelic variant of an A metabolic reaction that replenishes the cell sheds the apical region of enzyme. intermediates of pathways. plasma membrane as part of a (cid:2)adrenergic receptor A G-protein- anastomosis A convergence of two or signaling pathway. linked cell membrane receptor that more branches of a tubular structure; apoenzyme The proteinaceous part of binds norepinephrine preferentially, e.g., a direct connection between two an enzyme. with a lower affinity for epinephrine. arteries in the circulatory system. aquaporin A large tetrameric channel (cid:2)-helix A secondary structure of anatomical dead space The portion of that allows the passage of water protein or DNA in which the a respiratory structure that cannot through the plasma membrane. molecule twists in a characteristic participate in gas exchange (e.g., the arginine phosphate A major pattern, with structure stabilized by trachea and bronchi). phosphagen in invertebrates, which hydrogen bonds between adjacent androgens Steroid hormones performs the same role as creatine regions. structurally related to testosterone phosphate in vertebrates. alternative splicing One of the that control masculine features. aromatase See cytochrome P450 processes that can result in different anemia A condition in which the aromatase. mRNAs being coded by a single gene. number of erythrocytes or Arrhenius plot A curve relating Different exons of the gene are hemoglobin in the blood is lower temperature to activity, enabling the spliced out in each mRNA, resulting than normal. calculation of activation energy. in a number of possible angiogenesis Synthesis of new blood arteriole A small branch of the arterial combinations. vessels, often in response to local network immediately preceding a alveoli (singular: alveolus) The site of hypoxia. capillary bed (see venule). gas exchange in mammalian lungs. angiotensin A peptide hormone that artery A large blood vessel carrying ambient External or environmental controls blood pressure. Its precursor blood away from the heart. conditions, such as ambient is angiotensinogen, which is cleaved asexual reproduction Production of temperature. by renin to form angiotensin I. This offspring without the fertilization of amine A class of molecules based on decapeptide is cleaved to the final an ovum by a sperm (see also ammonia, with a side group form, angiotensin II, an octapeptide. automictic parthenogenesis). substituting for at least one N atom. angiotensin-converting enzyme (ACE) assimilation Conversion of dietary amino acid Organic molecules with at An enzyme that converts angiotensin nutrients into metabolizable fuels. least one amino group and at least I to angiotensin II. assimilation efficiency Proportion of one carboxyl group. The amino acids anion An ion with a negative charge. dietary nutrients successfully that are used to build proteins are (cid:4)- anoxic See anaerobic. assimilated. amino acids. antagonist A substance that binds to a astrocytes Vertebrate glial cells that ammonia A general term that includes receptor but does not stimulate a help to support and regulate the both NH and NH(cid:6)(ammonium), signaling event. Antagonists interfere action of neurons in the central 3 4 potent neurotoxins. with the binding of the natural nervous system. ammoniotele An animal with an ligand. asynchronous muscle A muscle in excretory strategy in which more than antagonistic controls For a given step which a single neuronal stimulation half of the nitrogen is excreted as or pathway, sets of controls that exert causes multiple cycles of contraction ammonia (see also ureotele, uricotele). opposing effects. and relaxation. amniote Vertebrates with an amnion, antagonistic muscle A muscle that ATP See adenosine triphosphate. namely reptiles, birds, and opposes the movement of another ATP-binding cassette A common mammals. muscle. structural motif found in diverse amphibolic pathway A metabolic anterior pituitary gland The anterior proteins that binds ATP. pathway that both synthesizes lobe of the pituitary gland of ATPase A class of proteins, including (catabolic) and degrades (anabolic) vertebrates, also called the enzymes and transporters, that metabolites. adenohypophysis; secretes tropic couples ATP hydrolysis to a amphipathic A molecule with both hormones. mechanical or chemical process. hydrophobic and hydrophilic parts. antidiuretic A substance that induces ATPS Standardized reference condition amplification An exponential increase a reduction in urine volume. for measuring gas volumes: ambient in activity from one step of a pathway antifreeze protein A protein that temperature, pressure, and saturated to the next; typically used in the disrupts the growth of ice crystals, with water. context of signal transduction allowing an organism to survive atresia The programmed cell death pathways. subzero temperatures. (apoptosis) of follicles other than the ampullae of Lorenzini Polymodal antigen A substance, usually a protein, dominant follicle that matures during receptors that detect both electrical that induces the formation of an the ovulatory cycle. and mechanical stimuli; found on the antibody that can bind the antigen. atrial natriuretic peptide (ANP) A nose of sharks. antiport (or exchanger) A transport peptide hormone produced in the amygdala A part of the limbic system protein that exchanges one ion (or heart that exerts effects on ion and of the vertebrate brain that is molecule) for another ion (or water balance that tend to reduce involved in emotional responses such molecule) on the opposite side of a blood pressure. It increases urine as fear and anger. membrane. volume and Na(cid:6)excretion. 2 GLOSSARY atrioventricular node (AV node) Part axoneme The microtubule-based taurine; assist in emulsification of of the conducting pathways of the structure that underlies flagella and lipid within the small intestine. mammalian heart; delays conduction cilia. binocular vision The ability to of the electrical signal between the axosomatic synapse A synapse formed compare the images coming from two atrium and ventricles. between the axon terminal of one eyes to produce three-dimensional atrium (plural: atria) One of the neuron and the soma (cell body) of perception. chambers of a heart. Blood moves another neuron. biogenic amine A class of from the atrium to the ventricle. neurotransmitters derived from atrophy Loss of tissue mass as a result baroreceptor A receptor that senses amino acids including the of dying cells; often seen with pressure (by sensing the resulting catecholamines and dopamine. locomotor muscle in response to stretch on the cell membrane). bioluminescence The production of prolonged periods of inactivity. basal lamina The extracellular matrix light by living organisms. August Krogh principle Principle that underlying a sheet of epithelial cells; bipolar neuron A neuron with two for every biological problem, there is part of the connective tissue formed main processes leading from the cell an organism on which it can most largely by fibroblasts. body, one of which conveys signals conveniently be studied. basal metabolic rate (BMR) The toward the cell body, and one of autocrine A type of cell signaling in metabolic rate of an homeothermic which conveys signals away from the which a single cell signals another animal at rest, at a thermal neutral cell body. cell of the same type, including itself. temperature, and post-absorptive blastocoel The cavity formed by the automictic parthenogenesis (see also resting metabolic rate, inpouching of the blastocyst, which Production of offspring by a female in standard metabolic rate). eventually forms the alimentary which the second polar body fuses basal nuclei Interconnected groups of canal. with the ovum to produce a diploid gray matter within the mammalian blastocyst The hollow sphere of cells offspring. brain. formed early in embryonic autonomic division (of the nervous base A molecule that accepts a proton, development. system) See autonomic nervous or otherwise causes a reduction in bleaching The fading of a system. proton concentration through effects photopigment following absorption of autonomic ganglia Ganglia of the on the dissociation of water. energy from photons. In the case of vertebrate peripheral nervous system. basement membrane See also basal the retinal-opsin complex, absorption autonomic nervous system Part of the lamina. of energy from light causes retinal to vertebrate peripheral nervous system basilar membrane The location of the dissociate from opsin. Opsin is not that controls largely involuntary auditory hair cells in the mammalian pigmented, and thus the functions such as heart rate. It is cochlea. photopigment loses its color. divided into three main branches: the basophil A type of white blood cell that blood The circulatory fluid in animals sympathetic, parasympathetic, and releases histamine; involved in the with closed circulatory systems. enteric nervous systems. vertebrate immune response. Generally contains proteins, ions, autotrophy An organism that batch reactor A chemical reactor in organic molecules, and various cell synthesizes its own nutrients from which nutrients enter and exit types. inorganic material, using the energy through the same opening; nutrients blood-brain barrier A specialized of the sun (photoautotroph) or are retained in the reactor and protective barrier made up of glial inorganic reactions digested; the undigested material is cells that separates the circulatory (chemoautotrophs). then expelled, and replaced by system and the central nervous Avogadro’s number The number of another batch of nutrients to be system in vertebrates. molecules in a mole (6.02252 ×1023). processed. blood vessels Tubes that carry blood axoaxonic synapse A synapse formed behavioral thermoregulation The use through an animal’s body. between the axon terminal of one of behavior to control the body blubber Subcutaneous lipid deposits of neuron and the axon of another temperature of a poikilotherm, or to marine mammals, which provide neuron (at any point along its length). reduce the costs of thermoregulation thermal insulation. axodendritic synapse A synapse for a homeotherm. Bohr effect A change in hemoglobin formed between the axon terminal of (cid:5)-oxidation Pathway of fatty acid oxygen affinity due to a change in one neuron and the dendrite of catabolism that produces acetyl CoA pH. another neuron. and reducing equivalents. bolus A volume of material introduced axon A projection of the cell body of a (cid:5)-sheet Protein folding pattern in into a flow-through system that neuron that is involved in carrying which stretches of amino acids are moves through the system as a unit, information, usually in the form of aligned along another amino acid with some dispersion along the way; action potentials, from the cell body stretch. This secondary structure is often used in the context of a bolus of to the axon terminal. stabilized by hydrogen bonds. food moving through the axon hillock The junction between the bilateral symmetry A body form in gastrointestinal tract. cell body and axon of a neuron. In which the body can be divided by a bombesin A hormone that regulates many neurons, the axon hillock is the single plane such that the right and release of gastrointestinal hormones site of action potential initiation, left sides are approximate mirror and control of gastrointestinal acting as the trigger zone for the images. motility in vertebrates. neuron. bile A thick, yellow-green fluid bond energy The energy required to axon terminal The distal end of an composed of salts, pigments, and form a chemical bond. axon that forms a synapse with an lipids produced by the liver and stored bone In vertebrates, a solid structure effector cell or neuron. by the gallbladder; when released into composed of mineralized axon varicosity A type of synapse in the small intestine it neutralizes extracellular matrix of osteocytes; which the presynaptic cell releases gastric acid and aids in the digestion with cartilage and tendon, it neurotransmitter at a series of of nutrients, particularly lipids. constitutes the skeleton. swellings along the axon. bile duct The connection between the book gills The respiratory surfaces of axonal transport Cytoskeletal- liver and the small intestine. water-breathing chelicerates such as mediated movement of organelles bile pigments Nondigestible horseshoe crabs. and vesicles along the length of an breakdown products of porphyrins, book lungs The respiratory surfaces of axon. including the hemes found in some air-breathing chelicerates such axonemal dyneins Motor proteins that hemoglobin and cytochromes. as spiders and scorpions. enable the sliding of microtubules in bile salts Cholic acid conjugated with boundary layer The region of a cilia and flagella. amino acids, primarily glycine and solution that is in direct contact or 3 GLOSSARY otherwise influenced by a surface; cable properties The electrical carotid rete A network of blood often called an unstirred layer. properties of axons. vessels that cools the brain. Bowman’s capsule A cup-shaped calcium-induced calcium release A carrier protein (or binding protein; expansion of the vertebrate kidney mode of muscle activation where binding globulin) Blood proteins tubule; surrounds the glomerulus. calcium crossing the sarcolemma that help to transport hydrophobic brackish water Water that is through a Ca2(cid:6)channel causes a Ca2(cid:6) molecules (such as steroid hormones) intermediate between freshwater and channel in the sarcoplasmic reticulum in the blood. seawater; typically found in estuaries, to open. carrier-mediated transport All forms salt marshes, or isolated ponds. caldesmon A calcium-binding protein of transport across membranes that bradycardia A heart rate that is important in the regulation of smooth require a protein. slower than normal. muscle contractility. cartilage In vertebrates, a semisolid brain A large grouping of ganglia that calmodulin A calcium-sensing protein structure composed of the act as a sophisticated integrating involved in many signal transduction extracellular matrix of chondrocytes: center. Typically located toward the pathways. the major component of the skeleton anterior end of the body in the caloric deficit The condition in which of chondrichthians but important in cephalic (head) region. energy derived from the diet is less other vertebrates as a cushion brainstem A portion of the vertebrate than energetic expenditure, resulting between joints. central nervous system that connects in net loss of energy by the animal. catabolic pathway (or catabolism) A the cerebrum of the brain to the calorie A unit of heat equal to 4.2 metabolic pathway that degrades spinal cord; contains the pons and joules; nutritional literature may macromolecules into smaller medulla, the sites of the respiratory refer to the unit Calorie, which is molecules. and cardiovascular control centers. equivalent to 1000 calories. The unit catadromous A life history strategy of branchial Relating to gills. of heat required to raise 1 g of water fish (e.g., eels) in which the adult bronchi (singular: bronchus) Airways at 1 atm by 1°C. migrates from freshwater to seawater of vertebrate lungs leading from the calorimetry The measurement of heat to breed (see also anadromous). trachea to the bronchioles. production as an index of metabolic catalysis The progression of a bronchioles The smallest branches of rate. chemical reaction that proceeds with the airways of mammalian lungs; calsequestrin A calcium-binding the help of a catalyst. lead to the terminal alveoli. protein that allows a muscle to catalyst A molecule that accelerates brood spot A well-vascularized, concentrate Ca2(cid:6)within the chemical reactions but is not changed featherless region on the underside of sarcoplasmic reticulum. in the process. birds that is important for warming cAMP (cyclic AMP) A second catalytic rate constant (k ) The cat developing eggs. messenger produced by adenylate number of reactions catalyzed by a brown adipose tissue Also known as cyclase; most important action is the single molecule of enzyme per second. brown fat, a thermogenic tissue stimulation of protein kinase A. catecholamines The biogenic amines found in many small mammals, often capacitation A maturation step epinephrine and norepinephrine. in the back or neck region. Abundant experienced by sperm after they cation An ion with a positive charge. mitochondria in the brown encounter fluids from the female caudal A location near the posterior of adipocytes possess thermogenin, a reproductive tract. an animal. protein that uncouples oxidative capillary The smallest of the blood cecum A blind-ended sac that carries phosphorylation to enhance heat vessels in a closed circulatory system; out digestive reactions in the production. the site of exchange of materials with gastrointestinal tract. brush border Abundant microvilli on the tissues. cell body See soma. epithelial cells in the gastrointestinal carbaminohemoglobin Hemoglobin cell membrane See plasma membrane. tract, giving the tissue a microscopic bound to carbon dioxide. cellular membranes A general term brushlike appearance. carbohydrate A group of organic that refers to the collection of BTPS Standardized reference molecules that share a membranes within a cell, including conditions for measuring gas preponderance of hydroxyl groups plasma membrane and organelle volumes: body temperature, (see also disaccharide, membranes. atmospheric pressure, and saturated monosaccharide, polysaccharide). cellulose A glucose polymer that with water. carbonic anhydrase (CA) An enzyme serves a structural role in plants; buccal cavity Mouth cavity. that catalyzes the conversion of indigestible by most animals without buffer Chemicals which, when placed carbon dioxide and water to the assistance of symbionts. in solution, confer on the solution an bicarbonate and protons. central chemoreceptors A group of ability to resist changes in pH when carboxyhemoglobin Hemoglobin chemoreceptors located in the acid or base is added. bound to carbon monoxide. medulla of vertebrate brains. bulbourethral gland A mucus- cardiac muscle A form of striated central lacteal A small, saclike vessel secreting accessory gland of the male muscle that occurs in the heart. in an intestinal villus; collects lipids reproductive tract. cardiac output The volume of blood that cross the intestinal epithelium. bulbus arteriosus The outflow pumped by the heart per unit time; central nervous system The portion of tract of the heart in bony fishes; the product of heart rate and stroke the nervous system containing the nonmuscular and elastic (see also volume. primary integrating centers. In conus arteriosus). cardiomyocyte A muscle cell found in vertebrates it consists of the brain bulk flow The movement of a fluid as a the heart. and spinal cord. In invertebrates, it result of a pressure or temperature cardiovascular control center A consists of the brain, the major gradient. region of the brain within the ganglia, and the connecting bulk phase (or bulk solution) The medulla oblongata that is involved in commissures. volume of solution that is beyond the regulating heart rate and blood central pattern generator A group of influence of the surfaces (see also pressure. neurons located in the central boundary layer). cardiovascular system An alternate nervous system that produce a bundle of His One of the conducting term for the circulatory system of rhythmic neural output. pathways of the mammalian heart. animals such as vertebrates. Consists cephalic Toward the anterior end of an burst exercise High-intensity of the heart, blood, and blood vessels. animal. exercise powered by glycolytic carotid body A structure located in the cephalization An evolutionary trend muscle fibers; can continue for only carotid artery leading to the head of toward the centralization of nervous short periods, until glycogen stores vertebrates; contains baroreceptors and sensory functions at the anterior are exhausted. and chemoreceptors. end of the body (in the head). 4 GLOSSARY cerebellum A part of the vertebrate cholesterol A steroid compound cloaca The distal portion of the hindbrain that is involved in produced from isoprene units; hindgut in some fishes, amphibians, maintaining balance and present in cellular membranes and birds, and reptiles; in these species coordinating voluntary muscle acts as a precursor for steroid both excretory and reproductive movement. hormones. products are emitted into the cloaca, cerebral cortex Outer surface of the cholinergic receptor A receptor that and leave the body via a single vertebrate brain. binds the signaling molecule opening. cerebral hemispheres Paired acetylcholine. Cholinergic receptors clonal reproduction A form of asexual structures of the cerebrum (part of can be divided into nicotinic and reproduction whereby an animal the vertebrate forebrain). The muscarinic receptors. produces a genotypically identical cerebral hemispheres are the most chondrocytes The cells that produce offspring (a clone). obvious structures of a mammalian cartilage. closed circulatory system A brain. chorion The outer protein layer of an circulatory system in which the blood cerebral ventricle See ventricle. insect egg; the outer membrane of a remains within a series of enclosed cerebrospinal fluid (CSF) A fluid vertebrate ovum. blood vessels throughout the contained within the meninges that chorionic gonadotropin (CG) A third circulation. surrounds the brain and spinal cord gonadotropin of vertebrates, cochlea Spiral structure in the inner of vertebrates. produced by the placenta but only in ear of mammals; contains the organs cerebrum The largest part of the primates. of hearing. Less elaborate, but mammalian forebrain. chromaffin cells Cells that secrete the present in birds as the cochlear duct. cGMP See cyclic GMP. hormone epinephrine (adrenaline). In Derived from the lagena of other cGMP phosphodiesterase An enzyme mammals they are located in the vertebrates. that cleaves cGMP, producing GMP. compact adrenal medulla, but in coelom The internal compartment of channel A transport protein that other vertebrates they are more coelomate animals that forms facilitates the movement of specific dispersed. between two layers of mesoderm. ions or molecules across a cellular chromophore A molecule that is able coenzymes Organic cofactors. membrane down an electrochemical to absorb light. In photoreception, coenzyme A A coenzyme derived from gradient. the chromophore absorbs the energy the vitamin pantothenic acid. chaperone protein See molecular from incoming photons and cofactors Nonprotein components of chaperone. undergoes a conformational change, enzymes, including metals, chemical energy The energy which sends a signal to an associated coenzymes, and prosthetic groups. associated with the reorganization G protein, in the first step of visual coitus Sexual intercourse. of the chemical structure of a phototransduction. collagen A trimeric protein found in molecule. chromosome A single, contiguous extracellular matrix. It interacts with chemical gradient An area across polymer of DNA found within the other collagen molecules to form which the concentration of a genome. rigid fibers or durable sheets. chemical differs, often across a chylomicron A large lipoprotein collecting duct The tube that receives membrane. complex that carries lipid from the the fluid from the distal tubules of the chemical synapse A junction between digestive tract through the nephron and empties into the minor a neuron and another cell in which circulation to processing and target calyx of the kidney. the signal is transmitted across the tissues. colligative properties Four properties synapse in the form of a cilia (singular: cilium) Microtubule- of a solute that are due solely to the neurotransmitter. based extensions from a cell that concentration of solutes, and not chemoautotroph An organism that move in a wavelike pattern. their chemical nature. uses inorganic chemical energy to ciliary body A part of the vertebrate colloidal osmotic pressure See oncotic convert organic sources of carbon eye that secretes the aqueous humor. pressure. and nitrogen into biosynthetic ciliary muscle The muscle that colon A region of the large intestine building blocks. controls the shape of the lens of the primarily responsible for water chemokinetic An increase in vertebrate eye; involved in producing resorption. nondirectional movement in response a focused image. compatible solute A solute that, at to the detection of a chemical. ciliary photoreceptors One of two high concentration, does not disrupt chemoreceptor Used to describe either types of animal photoreceptor cells. protein structure or enzyme kinetics. a cell containing chemoreceptive Vertebrate photoreceptors belong to competitive inhibition A mode of proteins, or the proteins themselves. this class (see also rhabdomeric enzyme inhibition in which a Chemicals such as hormones, photoreceptors). molecule competes with the substrate odorants, and tastants bind circadian rhythm Regular changes in for the active site on the enzyme; specifically to chemoreceptor gene expression, biochemistry, competitive inhibitors have the effect proteins, altering their conformation physiology, and behavior that cycle of reducing the apparent substrate and causing a signal within the with a period of approximately 24 affinity without affecting V . max chemoreceptor cell. hours. Endogenous circadian compliance A measure of the ability of chemotaxic Movement toward higher rhythms persist even in constant a hollow structure (e.g., blood vessel, concentrations of a chemical. darkness. lung) to stretch in response to an chief cell The secretory cells of the circulatory system A group of organs applied pressure. gastric epithelium that release and tissues involved in moving fluids compound eye A type of eye seen in pepsin. through the body; consists of one or arthropods; consists of many chitin A polymer of N-acetyl more pumping structures and a individual photoreceptive structures. glucosamine used by arthropods to series of tubes or other spaces conduction Transfer of heat from one construct the exoskeleton. through which fluid can move. object to another object or a fluid. chloride cell An ion-pumping cell of citric acid cycle See tricarboxylic acid cone A type of vertebrate fish gill epithelium (also called a cycle. photoreceptor cell (see also rod). mitochondria-rich cell). clathrin A triskelion-shaped (three- Cones are typically responsible for chloride shift The exchange of armed) protein that coats some types color vision in bright light. chloride and bicarbonate across the of vesicles; vesicle formation begins conformer A strategy whereby the erythrocyte membrane. with a clathrin-coated pit, which physicochemical properties of an chlorocruorin A type of hemoglobin enlarges to form a clathrin-coated animal (e.g., temperature and found in some annelids; known as vesicle. osmolarity) parallel those of the the green hemoglobins. clearance See renal clearance. environment. 5