A LANGE medical book Ganong’s Review of Medical Physiology T W E N T Y  F O U R T H E D I T I O N Kim E. Barrett, PhD Scott Boitano, PhD Professor, Department of Medicine Associate Professor, Physiology Dean of Graduate Studies Arizona Respiratory Center University of California, San Diego Bio5 Collaborative Research Institute La Jolla, California University of Arizona Tucson, Arizona Susan M. Barman, PhD Heddwen L. Brooks, PhD Professor, Department of Pharmacology/ Toxicology Associate Professor, Physiology Michigan State University College of Medicine East Lansing, Michigan Bio5 Collaborative Research Institute University of Arizona Tucson, Arizona New York Chicago San Francisco Lisbon London Madrid Mexico City Milan New Delhi San Juan Seoul Singapore Sydney Toronto Ganong_FM_i-x.indd i 1/6/12 5:30:16 PM Copyright © 2012 by Th e McGraw-Hill Companies, Inc. All rights reserved. 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Neither McGraw-Hill nor its licensors shall be liable to you or anyone else for any inaccuracy, error or omission, regardless of cause, in the work or for any damages resulting therefrom. McGraw-Hill has no responsibility for the content of any information accessed through the work. Under no circumstances shall McGraw-Hill and/or its licensors be liable for any indirect, incidental, special, punitive, consequential or similar damages that result from the use of or inability to use the work, even if any of them has been advised of the possibility of such damages. This limitation of liability shall apply to any claim or cause whatsoever whether such claim or cause arises in contract, tort or otherwise. Dedication to William Francis Ganong W illiam Francis (“Fran”) Ganong was an outstanding marked on and admired whenever the book came up for dis- scientist, educator, and writer. He was completely cussion among physiologists. He was an excellent writer and dedicated to the fi eld of physiology and medical ed- far ahead of his time with his objective of distilling a complex ucation in general. Chairman of the Department of Physiology subject into a concise presentation. Like his good friend, Dr. at the University of California, San Francisco, for many years, Jack Lange, founder of the Lange series of books, Fran took he received numerous teaching awards and loved working with great pride in the many diff erent translations of the Review of medical students. Medical Physiology and was always delighted to receive a copy Over the course of 40 years and some 22 editions, he was of the new edition in any language. the sole author of the best selling Review of Medical Physiology, He was a model author, organized, dedicated, and enthusi- and a co-author of 5 editions of Pathophysiology of Disease: An astic. His book was his pride and joy and like other best-selling Introduction to Clinical Medicine. He was one of the “deans” authors, he would work on the next edition seemingly every of the Lange group of authors who produced concise medical day, updating references, rewriting as needed, and always ready text and review books that to this day remain extraordinarily and on time when the next edition was due to the publisher. He popular in print and now in digital formats. Dr. Ganong made did the same with his other book, Pathophysiology of Disease: a gigantic impact on the education of countless medical stu- An Introduction to Clinical Medicine, a book that he worked on dents and clinicians. meticulously in the years following his formal retirement and A general physiologist par excellence and a neuroendo- appointment as an emeritus professor at UCSF. crine physiologist by subspecialty, Fran developed and main- Fran Ganong will always have a seat at the head table of tained a rare understanding of the entire fi eld of physiology. the greats of the art of medical science education and commu- Th is allowed him to write each new edition (every 2 years!) nication. He died on December 23, 2007. All of us who knew of the Review of Medical Physiology as a sole author, a feat re- him and worked with him miss him greatly. Ganong_FM_i-x.indd iii 1/6/12 5:30:17 PM Key Features of the 24th Edition of Ganong’s Review of Medical Physiology (cid:115)(cid:0)(cid:48)(cid:0) (cid:82)(cid:79)(cid:86)(cid:73)(cid:68)(cid:69)(cid:83)(cid:0)(cid:67)(cid:79)(cid:78)(cid:67)(cid:73)(cid:83)(cid:69)(cid:0)(cid:67)(cid:79)(cid:86)(cid:69)(cid:82)(cid:65)(cid:71)(cid:69)(cid:0)(cid:79)(cid:70)(cid:0)(cid:69)(cid:86)(cid:69)(cid:82)(cid:89)(cid:0)(cid:73)(cid:77)(cid:80)(cid:79)(cid:82)(cid:84)(cid:65)(cid:78)(cid:84)(cid:0)(cid:84)(cid:79)(cid:80)(cid:73)(cid:67)(cid:0)(cid:87)(cid:73)(cid:84)(cid:72)(cid:79)(cid:85)(cid:84)(cid:0)(cid:83)(cid:65)(cid:67)(cid:82)(cid:73)(cid:108)(cid:67)(cid:73)(cid:78)(cid:71)(cid:0)(cid:67)(cid:79)(cid:77)(cid:80)(cid:82)(cid:69)(cid:72)(cid:69)(cid:78)(cid:83)(cid:73)(cid:86)(cid:69)(cid:78)(cid:69)(cid:83)(cid:83)(cid:0) (cid:79)(cid:82)(cid:0)(cid:82)(cid:69)(cid:65)(cid:68)(cid:65)(cid:66)(cid:73)(cid:76)(cid:73)(cid:84)(cid:89) 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(cid:36)(cid:69)(cid:84)(cid:65)(cid:73)(cid:76)(cid:69)(cid:68)(cid:0)(cid:69)(cid:88)(cid:80)(cid:76)(cid:65)(cid:78)(cid:65)(cid:84)(cid:73)(cid:79)(cid:78)(cid:83)(cid:0)(cid:79)(cid:70)(cid:0) (cid:73)(cid:78)(cid:67)(cid:79)(cid:82)(cid:82)(cid:69)(cid:67)(cid:84)(cid:0)(cid:65)(cid:78)(cid:83)(cid:87)(cid:69)(cid:82)(cid:0)(cid:67)(cid:72)(cid:79)(cid:73)(cid:67)(cid:69)(cid:83)(cid:0) (cid:115)(cid:0)(cid:0)NEW:(cid:0)(cid:45)(cid:79)(cid:82)(cid:69)(cid:0)(cid:67)(cid:76)(cid:73)(cid:78)(cid:73)(cid:67)(cid:65)(cid:76)(cid:0)(cid:67)(cid:65)(cid:83)(cid:69)(cid:83)(cid:0)(cid:65)(cid:78)(cid:68)(cid:0)(cid:109)(cid:79)(cid:87)(cid:0) CHAPTER 34 Introduction to Pulmonary Structure and Mechanics 625 (cid:67)(cid:72)(cid:65)(cid:82)(cid:84)(cid:83)(cid:0)(cid:84)(cid:72)(cid:65)(cid:78)(cid:0)(cid:69)(cid:86)(cid:69)(cid:82) Capillaries ma (cid:115)(cid:0)(cid:0)NEW:(cid:0)(cid:37)(cid:88)(cid:80)(cid:65)(cid:78)(cid:68)(cid:69)(cid:68)(cid:0)(cid:76)(cid:69)(cid:71)(cid:69)(cid:78)(cid:68)(cid:83)(cid:0)(cid:70)(cid:79)(cid:82)(cid:0)(cid:69)(cid:65)(cid:67)(cid:72)(cid:0) Respiratory bronchiole (cid:73)(cid:76)(cid:76)(cid:85)(cid:83)(cid:84)(cid:82)(cid:65)(cid:84)(cid:73)(cid:79)(cid:78)(cid:136)(cid:83)(cid:79)(cid:0)(cid:89)(cid:79)(cid:85)(cid:0)(cid:68)(cid:79)(cid:78)(cid:7)(cid:84)(cid:0)(cid:72)(cid:65)(cid:86)(cid:69)(cid:0)(cid:84)(cid:79)(cid:0) Alveolar duct Alveolus cf a (cid:82)(cid:69)(cid:70)(cid:69)(cid:82)(cid:0)(cid:66)(cid:65)(cid:67)(cid:75)(cid:0)(cid:84)(cid:79)(cid:0)(cid:84)(cid:72)(cid:69)(cid:0)(cid:84)(cid:69)(cid:88)(cid:84) Alveolus Apolvreeolus en epI a cap Alveolus cf A C Cenadpoiltlahreylium Alveolar air Type II cell Bmaesmebmraennet Air space SF TM N N Erythrocyte Interstitium Pinl acsampiallary LB Amlavecroolaprhage CB Type I cell Erythrocyte Type II Golgi Fatty acids cell N RER CGhlyocleinreol B Type I cell Alveolar air D AEmtcino acids More than 600 full-color FIGURE 343 Prominent cells in the adult human alveolus. surfactant. Lamellar bodies (LB) are formed in type II alveolar epithelial A) A cross-section of the respiratory zone shows the relationship cells and secreted by exocytosis into the fl uid lining the alveoli. illustrations between capillaries and the airway epithelium. Only 4 of the 18 alveoli The released lamellar body material is converted to tubular myelin are labeled. B) Enlargement of the boxed area from (A) displaying (TM), and the TM is the source of the phospholipid surface fi lm (SF). intimate relationship between capillaries, the interstitium, and the Surfactant is taken up by endocytosis into alveolar macrophages and alveolar epithelium. C) Electron micrograph displaying a typical area type II epithelial cells. N, nucleus; RER, rough endoplasmic reticulum; depicted in (B). The pulmonary capillary (cap) in the septum contains CB, composite body. (Reproduced with permission from (A, B) Widmaier EP, Raff plasma with red blood cells. Note the closely apposed endothelial H, Strang KT: Vander’s Human Physiology: The Mechanisms of Body Function, 11th ed. and pulmonary epithelial cell membranes separated at places by McGraw-Hill, 2008; (C) Burri PA: Development and growth of the human lung. In: additional connective tissue fi bers (cf); en, nucleus of endothelial Handbook of Physiology, Section 3, The Respiratory System. Fishman AP, Fisher AB cell; epl, nucleus of type I alveolar epithelial cell; a, alveolar space; ma, [editors]. American Physiological Society, 1985; and (D) Wright JR: Metabolism and alveolar macrophage. D) Type II cell formation and metabolism of turnover of lung surfactant. Am Rev Respir Dis 136:426, 1987.) 0.5 (μm apart ( Figure 34–3 ). Th e alveoli also contain other process inhaled antigens for immunologic attack, and they specialized cells, including pulmonary alveolar macrophages secrete substances that attract granulocytes to the lungs as (PAMs, or AMs), lymphocytes, plasma cells, neuroendocrine well as substances that stimulate granulocyte and monocyte cells, and mast cells. PAMs are an important component of formation in the bone marrow. PAM function can also be the pulmonary defense system. Like other macrophages, detrimental—when they ingest large amounts of the sub- these cells come originally from the bone marrow. PAMs are stances in cigarette smoke or other irritants, they may release actively phagocytic and ingest small particles that evade the lysosomal products into the extracellular space to cause mucociliary escalator and reach the alveoli. Th ey also help infl ammation. CHAPTER 12 Refl ex and Voluntary Control of Posture & Movement 253 vestibulospinal tracts) regulate proximal muscles and posture. 3. Withdrawal refl exes are not Th e lateral corticospinal and rubrospinal tracts control distal A. initiated by nociceptive stimuli. limb muscles for fi ne motor control and skilled voluntary B. prepotent. movements. C. prolonged if the stimulus is strong. ■ Decerebrate rigidity leads to hyperactivity in extensor D. an example of a fl exor refl ex. muscles in all four extremities; it is actually spasticity due to E. accompanied by the same response on both sides of the body. fuanccilailt ahteiornn ioatfi tohne dsutree ttcoh a r seufl perxa. tIetn rteosreimal blelessio wnh. Dat eics osreteinca wtei th 4. Wof htiinleg leixnegr cinis ihnegr, rai g4h2t- yleega ra-nodld a fne minaalbei lditeyv etolo cpoendt rsould mdeonv eomnseentt End-of-chapter review rigidity is fl exion of the upper extremities at the elbow and in that limb. A neurological exam showed a hyperactive knee extensor hyperactivity in the lower extremities. It occurs on the jerk refl ex and a positive Babinski sign. Which of the following hcaepmsuiplele. gic side aft er hemorrhage or thrombosis in the internal Ais .n Rote cfl heaxreasc ctaenri sbteic m oof da irfie efl de xb?y impulses from various parts of questions help you assess ■ Th e basal ganglia include the caudate nucleus, putamen, the CNS. globus pallidus, subthalamic nucleus, and substantia nigra. Th e B. Refl exes may involve simultaneous contraction of some connections between the parts of the basal ganglia include a muscles and relaxation of others. your comprehension dopaminergic nigrostriatal projection from the substantia nigra C. Refl exes are chronically suppressed aft er spinal cord to the striatum and a GABAergic projection from the striatum transection. to substantia nigra. D. Refl exes involve transmission across at least one synapse. ■ Parkinson disease is due to degeneration of the nigrostriatal E. Refl exes oft en occur without conscious perception. dopaminergic neurons and is characterized by akinesia, 5. Increased neural activity before a skilled voluntary movement is bradykinesia, cogwheel rigidity, and tremor at rest. Huntington fi rst seen in the disease is characterized by choreiform movements due to A. spinal motor neurons. the loss of the GABAergic inhibitory pathway to the globus B. precentral motor cortex. pallidus. C. midbrain. ■ Th e cerebellar cortex contains fi ve types of neurons: D. cerebellum. Purkinje, granule, basket, stellate, and Golgi cells. Th e two E. cortical association areas. main inputs to the cerebellar cortex are climbing fi bers 6. A 58-year-old woman was brought to the emergency room of and mossy fi bers. Purkinje cells are the only output from her local hospital because of a sudden change of consciousness. the cerebellar cortex, and they generally project to the All four limbs were extended, suggestive of decerebrate rigidity. deep nuclei. Damage to the cerebellum leads to several A brain CT showed a rostral pontine hemorrhage. Which of characteristic abnormalities, including hypotonia, ataxia, the following describes components of the central pathway and intention tremor. responsible for control of posture? A. d Tho res toelcattoersapli naarel ap aotfh twhea ys pteinrmal ivneantetrsa ol nh onrenu rtohnats iinnn tehrev ate 246 SECTION II Central and Peripheral Neurophysiology MULTIPLECHOICE QUESTIONS B. l Thim eb m meudsucllleasr.y reticulospinal pathway terminates on Fdoirre catlel dq.u estions, select the single best answer unless otherwise C. nt Thh eauet r piononnnse tirinnv eat htreee atvixceiunallto raosnpmdin epadrlio apxla iatmrhewaala omyf uttehsrceml esispn.i antaels voenn tnreaul rhoonrsn CLINICAL BOX 12–7 1. W hen dynamic γ-motor neurons are activated at the same time in the dorsomedial area of the spinal ventral horn that Basal Ganglia Diseases the characteristic yellow K ayser–Fleischer rings. The domi- 2. B ThAABCDEas ....... αet o ip c t tt tw p -sehhhhilrlcm nohnaeeeaaocnvce nd muonnmmeenuo truuw.rp oss unoαsmm trhesin w osdci enbbos nhrllniteeeesgisherk uy rrcaαenwi enhrbnoot loicdaIaiy.fflt ahnp rli iii s tg osmmtnsocrie pneotch ppooifla tanro uu clercemofdcllxgofl ssndu leueeneeeir sss xtas. a c ric liiffsiaolnnh ne ceni a,insstrter.ppeic g aniiirennstete ddsaiidnn slla eee cbrs dreIIpyeaa. i iaa nnaascdffffhet ldiieeevbrr. a ieeIttannei odttassnff. ii essor fsge mrtnhetaaeslt ltGeearrko telhgsai n 7. thcW AEADuoo. .. m3n hsda Ida Thi Th8psinotcxxc-oo i rthniyiiirree.naoaa e ess Slovllulmaoor ahbfoaarsvml t-eneanneltaevoh tcdedtderweelaeesaidd rsau papsllfia .lshisaofrrv lme Sevoo lylbea ml hexxapsoorbrsertiioeewfa tmmie muarbialiaibear glunaao ur scuohhlllgeofts alf tmmagi ac t ri vsdtuhtslr hiheuuop tle npbsoae yssa t.oasrecc w t hsi pttllehnbpeee henitwr ss dinrwee ne..uda a mxasaayeiltity alg e ehv ta gvnrnteeibnegenrssnorneom ogtomuntsrr faritea c nima nddlynd lade h euarwhtc cotoespeoeiescrortrsr rnlhomeo neeo b gnsbta n t rrhor oheanm aafnfsta tet ehseeaet ui i tue lpvinrrlanr reioonsf gon lotsnleoinaoteudsructrss rviiair svtc nialliy an ointb ?mtet fgreth ah. b eiesn . tGrtttmtswd TeohhhhihibAisle eeeeteeelBmhn e apaedhAi,tsnie saniaasneesed t dnnire1sia tigead dua0ipoaffnii s –mcn palfweet 1 h .epp tc h5dSithastarbo eeppsroeyti nthiet5 geadenir ew aonrecoeryecrendtaav usshaanc, ey ts bg ach eilbopvothfrl.ueoetaf eepie Lr nc 1dr .odtram odg0uh Antetm0a aeeihmsntt,mlr o0teel ,gyi iee0 s nnahl otn ao0igg soynntr btlepp uiluchsayut eecrereh esirihrostsn ee ka i ppsogf d isatonlahfne H r e laelisy ls lauonwuptiiypdstinwndmocuo euat t tmrert csipi, kllh nd dhj t.ieeete ng owobhsxrTnemtkprytehio tdeeye iyiidsnn sricec .ftne ci . iornH laaoafIaodterclutlmj lms iihaaeynsssstpc, pe e t iututnamiograoonreschfmeswrguo hieestytatveaea rh olirennootltiidneyss-t--ff aiccvoabnp“nwaloznfaaau ilolsdnraid dvntaarnt Aee eirgl bgoo issnn g-eng itfoboetnhii oahtdruett.ehn r hheiLharmrtego eoe tyabdrilrn an.l in a oti laladgseegisp tol-aimn”i isetser lpteet d rpsrgiarai tee cdomaatsimurnhorner awl doul goa.trc lsiTihlyrroovtie h a ehomoge n,eo r r y nsbu rmf da pte euctirsyfseohiultota ser neerdirtmkutosleure yiesii,lanmgs xe .trnd e aeiecp.Mser srnafTtu ,deniu ieoah dgcritsvrt.re ys uaeeode Tsmtdndr mrdaih uo c adbrytioogsnoytsyit nhs sk o pdoatmt irsrunrrisfi suoo a eredetabcldd ,shlia hesauiudiesatcd nnceiau asi ecp sssirelne i t bubintapevasratsidha eotoanerideealmnco cuetnoyhcegmned oflegse utn ttemaanihidh,nnnri iitieeayct----, C. is a disynaptic refl ex with a single interneuron inserted B. Th e excitatory input from the reticulospinal pathway activates ed as an autosomal dominant disorder, and its onset is usually movements of the face and tongue and cogwheel rigidity. The between the aff erent and eff erent limbs. γ-motor neurons which indirectly activate α-motor neurons. between the ages of 30 and 50. The abnormal gene respon- neuroleptic drugs act via blockade of dopaminergic transmis- ED.. ob u isres gaetw asp netoy.e lpnyes ty hInIe aa apffff t eiecrre renentflt fi e axbn ewdr siet fffh re omrmean ntthy l eiim nGtbeosrl.ng ie uternodnos nin serted CDE... Is I Lttto rirewset saececrhm t eurxbeatflllre leeysxm wa. ithtyiaepts e ea norfse u sepexsat seatnfti cdeiret ydu dnwuciaethl t hotoe irennsh ipaibtoiiiotninote.n d o ifn twhaer d. scgwihubitralhoen m fiHnoouers n to(hCtmieAn edgG it)4so re. neIat ps dneeioa sisertms al,o seacelaa,l yctthe hcd oics onn ndteauianimrng tsbh f1eoe1r re –gin3sl du4i n tocacfymr tteohiansesie ens.hd eIno -ta ropdt aea4trn2imei–nn 8oet6s-f aesimnocnien. seP rorgonil com nroegtceoedrp ctdoornrustg ro aunl.s de alena dims tboa lhaynpcee rsine nnsiitgirvoitsyt roiaft aDl3 idnofl pu-- or more copies, and the greater the number of repeats, the ethaerl ideirs tehaes ea.g Teh oef g oennsee tc aondde st hfoe rm hournet rianpgitdi nth, ea pprroogtereins soiof nu no-f THERAPEUTIC HIGHLIGHTS known function. Poorly soluble protein aggregates, which are Treatment for Huntington disease is directed at treat- toxic, form in cell nuclei and elsewhere. However, the correla- ing the symptoms and maintaining quality of life as tion between aggregates and symptoms is less than perfect. there is no cure. In general, drugs used to treat the It appears that a loss of the function of huntingtin occurs that symptoms of this disease have side eff ects such as fa- is proportional to the size of the CAG insert. In animal mod- tigue, nausea, and restlessness. In August 2008, the els of the disease, intrastriatal grafting of fetal striatal tissue U.S. Food and Drug Administration approved the use improves cognitive performance. In addition, tissue caspase-1 of tetrabenazine to reduce choreiform movements activity is increased in the brains of humans and animals with that characterize the disease. This drug binds revers- the disease, and in mice in which the gene for this apoptosis- ibly to vesicular monoamine transporters (VMAT) and regulating enzyme has been knocked out, progression of the thus inhibits the uptake of monoamines into synaptic Clinical cases add disease is slowed. vesicles. It also acts as a dopamine receptor antago- Another basal ganglia disorder is W ilson disease (or he- nist. Tetrabenazine is the fi rst drug to receive approval patolenticular degeneration ), which is a rare disorder of for individuals with Huntington disease. It is also used real-world relevance cyoeaprpse or fm agetea, baoff leiscmti nwgh aicbho uhta fso uanr toimnseest abs emtwaeneyn f e6m aanleds 2a5s taos ttraeradti voet hdery shkyinpeesrkiai.n e Ctihce mlaotivnegm eangte nditsso r(deegr,s psuecnh- males. Wilson disease aff ects about 30,000 people worldwide. icillamine , t rienthine ) are used to reduce the copper to the text Iottrn ais nt ashp ego elrontinentggi c a AarTmuPt oaossfeo cmghearonl rmee co( AesTsosPmi7vBee) d1ini3sq ot.rh dIete arli ffvd eeurce, t lsteo at hade imn cguo tptaopt ieoarnn- idinny stphkaietn ieebsnoitads y hw ainist hpin rpdosvivyeicndh utiaoatl sbri ecw dditiihffis o Wcrudillestr otson i stdr oeisafettae. snTer e.d aTiratemrcdetievndet accumulation of copper in the liver and resultant progressive at prescribing a neuroleptic with less likelihood of caus- liver damage. About 1% of the population carries a single ing the disorder. Clozapine is an example of an atypical abnormal copy of this gene but does not develop any symp- neuroleptic drug that has been an eff ective substitute toms. A child who inherits the gene from both parents may for traditional neuroleptic drugs but with less risk for develop the disease. In aff ected individuals, copper accumu- development of tardive dyskinesia. lates in the periphery of the cornea in the eye accounting for Also available on the iPad through Inkling (cid:115)(cid:0)(cid:0)(cid:54)(cid:73)(cid:86)(cid:73)(cid:68)(cid:0)(cid:70)(cid:85)(cid:76)(cid:76)(cid:13)(cid:67)(cid:79)(cid:76)(cid:79)(cid:82)(cid:0)(cid:65)(cid:78)(cid:68)(cid:0) (cid:65)(cid:78)(cid:78)(cid:79)(cid:84)(cid:65)(cid:84)(cid:69)(cid:68)(cid:0)(cid:68)(cid:73)(cid:65)(cid:71)(cid:82)(cid:65)(cid:77)(cid:83) (cid:115)(cid:0)(cid:0)(cid:41)(cid:78)(cid:84)(cid:69)(cid:82)(cid:65)(cid:67)(cid:84)(cid:73)(cid:86)(cid:69)(cid:0)(cid:81)(cid:85)(cid:73)(cid:90)(cid:90)(cid:69)(cid:83)(cid:136)(cid:87)(cid:73)(cid:84)(cid:72)(cid:0) (cid:67)(cid:79)(cid:78)(cid:83)(cid:84)(cid:82)(cid:85)(cid:67)(cid:84)(cid:73)(cid:86)(cid:69)(cid:0)(cid:70)(cid:69)(cid:69)(cid:68)(cid:66)(cid:65)(cid:67)(cid:75) (cid:115)(cid:0)(cid:0)(cid:54)(cid:73)(cid:68)(cid:69)(cid:79)(cid:83)(cid:0)(cid:65)(cid:78)(cid:68)(cid:0)(cid:65)(cid:78)(cid:73)(cid:77)(cid:65)(cid:84)(cid:73)(cid:79)(cid:78)(cid:83)(cid:0) (cid:68)(cid:69)(cid:77)(cid:79)(cid:78)(cid:83)(cid:84)(cid:82)(cid:65)(cid:84)(cid:73)(cid:78)(cid:71)(cid:0)(cid:84)(cid:72)(cid:69)(cid:0)(cid:70)(cid:85)(cid:78)(cid:67)(cid:84)(cid:73)(cid:79)(cid:78)(cid:0) (cid:79)(cid:70)(cid:0)(cid:72)(cid:85)(cid:77)(cid:65)(cid:78)(cid:0)(cid:83)(cid:89)(cid:83)(cid:84)(cid:69)(cid:77)(cid:83) About the Authors KIM E. BARRETT MSU Faculty Professional Women’s Association and an MSU College of Human Medicine Distinguished Faculty Award. Kim Barrett received her PhD in biological She has been very active in the American Physiological Soci- chemistry from University College London ety (APS) and was recently elected to serve as its 85th Presi- in 1982. Following postdoctoral training at dent. She has also served as a Councillor as well as Chair of the the National Institutes of Health, she joined Central Nervous System Section of APS, Women in Physiol- the faculty at the University of California, ogy Committee and Section Advisory Committee of APS. In San Diego, School of Medicine in 1985, her spare time, she enjoys daily walks, aerobic exercising, and rising to her current rank of Professor of mind-challenging activities like puzzles of various sorts. Medicine in 1996. Since 2006, she has also served the University as Dean of Gradu- ate Studies. Her research interests focus on the physiology SCOTT BOITANO and pathophysiology of the intestinal epithelium, and how its function is altered by commensal, probiotics, and pathogenic Scott Boitano received his PhD bacteria as well as in specifi c disease states, such as infl amma- in genetics and cell biology from tory bowel diseases. She has published more than 200 articles, Washington State University in chapters, and reviews, and has received several honors for Pullman, Washington, where he her research accomplishments including the Bowditch and acquired an interest in cellular sig- Davenport Lectureships from the American Physiological So- naling. He fostered this interest at ciety and the degree of Doctor of Medical Sciences, honoris University of California, Los Ange- causa, from Queens University, Belfast. She has also been very les, where he focused his research active in scholarly editing, serving currently as the Deputy on second messengers and cellular physiology of the lung Editor-in-Chief of the Journal of Physiology. She is also a dedi- epithelium. He continued to foster these research interests at cated and award-winning instructor of medical, pharmacy, and the University of Wyoming and at his current positions with graduate students, and has taught various topics in medical and the Department of Physiology and the Arizona Respiratory systems physiology to these groups for more than 20 years. Her Center, both at the University of Arizona. eff orts as a teacher and mentor will be recognized with the Bodil M. Schmidt-Nielson Distinguished Mentor and Scientist HEDDWEN L. BROOKS Award from the American Physiological Society in 2012. Her teaching experiences led her to author a prior volume (Gas- Heddwen Brooks received her PhD trointestinal Physiology, McGraw-Hill, 2005) and she was hon- from Imperial College, University of ored to have been invited to take over the helm of Ganong in London and is an Associate Professor 2007 for the 23rd edition, and to have guided this new edition. in the Department of Physiology at the University of Arizona (UA). Dr Brooks SUSAN M. BARMAN is a renal physiologist and is best known for her development of microarray Susan Barman received her PhD in physi- technology to address in vivo signal- ology from Loyola University School of ing pathways involved in the hormonal Medicine in Maywood, Illinois. Aft erward regulation of renal function. Dr Brooks’ many awards include she went to Michigan State University the American Physiological Society (APS) Lazaro J. Mandel (MSU) where she is currently a Profes- Young Investigator Award, which is for an individual demon- sor in the Department of Pharmacology/ strating outstanding promise in epithelial or renal physiology. Toxicology and the Neuroscience Pro- In 2009, she received the APS Renal Young Investigator Award gram. Dr Barman has had a career-long at the annual meeting of the Federation of American Societ- interest in neural control of cardiorespi- ies for Experimental Biology. Dr Brooks is currently Chair of ratory function with an emphasis on the the APS Renal Section Steering Committee. She serves on the characterization and origin of the naturally occurring dis- Editorial Board of the American Journal of Physiology-Renal charges of sympathetic and phrenic nerves. She was a recipient Physiology (since 2001), and has served on study sections of of a prestigious National Institutes of Health MERIT (Method the National Institutes of Health and the American Heart As- to Extend Research in Time) Award. She is also a recipient of sociation. She is a current member of the Merit Review Board an Outstanding University Woman Faculty Award from the for the Department of Veterans’ Aff airs. vi Ganong_FM_i-x.indd vi 1/23/12 2:41:57 PM Contents Preface ix 13 Autonomic Nervous System 255 S E C T I O N Cellular and Molecular I Basis for Medical 14 Electrical Activity of the Brain, Sleep– Physiology 1 Wake States, & Circadian Rhythms 269 15 Learning, Memory, Language, 1 General Principles & Energy Production & Speech 283 in Medical Physiology 3 2 Overview of Cellular Physiology in Medical Physiology 35 S E C T I O N Endocrine and III Reproductive 3 Immunity, Infection, & Infl ammation 67 Physiology 297 4 Excitable Tissue: Nerve 83 16 Basic Concepts of Endocrine 5 Excitable Tissue: Muscle 97 Regulation 299 6 Synaptic & Junctional Transmission 119 17 Hypothalamic Regulation of Hormonal Functions 307 7 Neurotransmitters & Neuromodulators 135 18 The Pituitary Gland 323 19 The Thyroid Gland 339 S E C T I O N Central and Peripheral II Neurophysiology 155 20 The Adrenal Medulla & Adrenal Cortex 353 21 Hormonal Control of Calcium & Phosphate Metabolism & the Physiology of Bone 377 8 Somatosensory Neurotransmission: Touch, Pain, and Temperature 157 22 Reproductive Development & Function of the Female Reproductive System 391 9 Vision 177 23 Function of the Male Reproductive 10 Hearing & Equilibrium 199 System 419 11 Smell & Taste 217 24 Endocrine Functions of the Pancreas 12 Refl ex and Voluntary Control of & Regulation of Carbohydrate Posture & Movement 227 Metabolism 431 vii Ganong_FM_i-x.indd vii 1/6/12 5:30:17 PM viii CONTENTS S E C T I O N S E C T I O N Gastrointestinal IV VI Respiratory Physiology 619 Physiology 453 25 Overview of Gastrointestinal Function 34 Introduction to Pulmonary Structure & Regulation 455 and Mechanics 621 26 Digestion, Absorption, & Nutritional 35 Gas Transport & pH 641 Principles 477 36 Regulation of Respiration 657 27 Gastrointestinal Motility 497 28 Transport & Metabolic Functions of the Liver 509 S E C T I O N VII Renal Physiology 671 S E C T I O N Cardiovascular V Physiology 519 37 Renal Function & Micturition 673 38 Regulation of Extracellular Fluid Composition & Volume 697 29 Origin of the Heartbeat & the Electrical Activity of the Heart 521 39 Acidifi cation of the Urine & Bicarbonate Excretion 711 30 The Heart as a Pump 539 31 Blood as a Circulatory Fluid & the Dynamics Answers to Multiple Choice Questions 721 of Blood & Lymph Flow 555 Index 723 32 Cardiovascular Regulatory Mechanisms 587 33 Circulation Through Special Regions 601 Ganong_FM_i-x.indd viii 1/6/12 5:30:17 PM Preface FROM THE AUTHORS NEWGANONG’S REVIEW OF MEDICAL PHYSIOLOGY 24/E COMES We are very pleased to launch the 24th edition of Ganong’s Re- view of Medical Physiology. Th e current authors have attempt- TO LIFE WITH THE GANONG IPAD ed to maintain the highest standards of excellence, accuracy, DIGITAL VERSION! and pedagogy developed by Fran Ganong over the 46 years in which he educated countless students worldwide with this • Integrated assessment textbook. • Engaging interactivity and gorgeous, high resolution We were pleased with the reaction to the 23rd edition, illustrations. our fi rst at the helm. However, recognizing that improvement • Concepts are brought to life with movies, and integrates is always possible, and that medical knowledge is constantly them right into the book evolving, we convened panels both of expert colleagues and of students to give us feedback on style, content, level and or- • Opportunity to buy individual chapters ganizational issues. Based on this input, we have thoroughly reorganized the text and redoubled our eff orts to ensure that NEW! GANONG ONLINE the book presents state of the art knowledge. We have also in- creased clinical content, particularly related to the burden of LEARNING CENTER disease states that arise from abnormal physiology of the sys- WWW.LANGETEXTBOOKS.COM/BARRETT tems we discuss. We remain grateful to many colleagues and students who Th is dedicated Ganong website will include the following: contact us with suggestions for clarifi cations and new mate- rial. Th is input helps us to ensure that the text is as useful as • Movies and Animations for both students and professors possible. We hope that you enjoy the fruits of our labors, and to access. See concepts come to life! the new material in the 24th Edition. • PowerPoint of all images and tables for the instructor Th is edition is a revision of the original works of Dr. Francis • Review Questions for students to test themselves Ganong. NEW TO THIS EDITION! NEW THERAPEUTIC HIGHLIGHTS Each section will now have an introduction • Recognizing the critical links between physiology and therapeutics, the boxed clinical cases, also now include • Information on the burden of disease associated with succinct summaries of modern pharmacological each organ system approaches to the treatment or management of the • New introductory materials covering overarching condition discussed. principles of endocrine regulation in physiology • Answers to the review questions in the book, with additional explanations to incorrect questions will now be included • Additional Flow Charts—Students expressed how helpful fl ow charts are in tying concepts together and seeing the big picture! • Chapter summaries tied to chapter objectives • Expanded legends—Th is will help students understand fi gures, without necessarily referring back to the text • Increased number of clinical cases ix Ganong_FM_i-x.indd ix 1/23/12 2:42:22 PM