Ganong’s Review of Medical Physiology

763 Pages·2016·125.49 MB·English

A LANGE medical book
Ganong’s Review
of
Medical Physiology
T W E N T Y - F I F T H E D I T I O N
Kim E. Barrett, PhD Scott Boitano, PhD
Distinguished Professor, Department of Medicine Professor, Physiology and Cellular and Molecular
Dean of the Graduate Division Medicine
University of California, San Diego Arizona Respiratory Center
La Jolla, California Bio5 Collaborative Research Institute
University of Arizona
Tucson, Arizona
Susan M. Barman, PhD
Professor, Department of Pharmacology/
Heddwen L. Brooks, PhD
Toxicology
Michigan State University Professor, Physiology and Pharmacology
East Lansing, Michigan College of Medicine
University of Arizona
Tucson, Arizona
New York Chicago San Francisco Athens London Madrid Mexico City
Milan New Delhi Singapore Sydney Toronto
Barrett_FM_i-xii_P1.indd 1 6/30/15 1:20 PM
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Dedication to
William Francis Ganong
W
illiam Francis (“Fran”) Ganong was an outstanding remarked 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 field 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 different 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 field of physiology. the greats of the art of medical science education and commu-
This 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 him and worked with him miss him greatly.
Barrett_FM_i-xii_P1.indd 3 6/30/15 1:20 PM
Key Features of the Twenty-Fifth Edition of
CHAPTER 11 Smell & Taste 225
Ganong’s Review of Medical Physiology that Taraes tsei mexilhairb iints saoftmeer wreaaycst itoon vsi saunadl acftoenrt irmasat gpesh eannodm ceonna- 2. AO n3e7 -oyfe tahre- oplodt feenmtiaalle c wonasse dqiuaegnncoesse do fw thitihs dmisuoltridpeler sisc lerosis.
trasts. Some of these are chemical “tricks,” but others may be diminished taste sensitivity. Taste receptors
true central phenomena. A taste-modifier protein, miraculin, A. for sweet, sour, bitter, salt, and umami are spatially separated
has been discovered in a plant. When applied to the tongue, on the surface of the tongue.
this protein makes acids taste sweet. B. are synonymous with taste buds.
A concise, up-to-date and clinically relevant review anovfe esarsvs.Ao iThoindniiemsn sgtauo lprs nv,o ioivisvnaoeclnl lv sufa.odlouindeg so fihf s uuemacthian nagvs ,e trhfsoeio rfmnoso idps aiasrp tfpiocaulrlloeanwrtle yid n s btteyrro minllgs- 3. CDEW... haagAicrrlloeehl s oaison foft n ytpthephhreeeav aroafybotfen olcldgvoh eewbea.myiln anogffer derevcroeeeenpss.tt nso ori.nt itnhcer efaacsiea tl,h ter iagbeimlitiyn atol, and
discriminate many different odors?
CHAPTER SUMMARY A. Many different receptors
B. Pattern of olfactory receptors activated by a given
of human physiology ■ Oawniltdfha bcintao strahyle ss uteepnmpso ecrrey pl lnos erautrieroo nlno osc,fa stteuhdpe p innoa rtsthainel cgoa l(vfsaiutcyst.toernyt aecpuitlhare)li cuemlls , C. oPthdreoo jbrearcnatiitnon of different mitral cell axons to different parts of End-of-chapter review
■ The cilia located on the dendritic knob of the olfactory D. High β-arrestin content in olfactory neurons
■ sGdfIonee-rnfnpomsdrroo rmoritytleaef iatnsnic eosotu.nof Ar rmfoyrxno iogt mrcnloaos lmtn ohatefnea r oidounl lflt fiaoua.cdftctoteoordrray ycn setbel rlunsel scbione rtp rtytha onveree solu stlrf hvaoaicnatt s oat hcrreyoe n cblaotuatulecbprts alte lht doe t o 4. EAdaisna. s hmd aSe ean rmngeiffietsoou tsnrlothtg ltioinhkf aeea llnb ycr oataourint teio cximenpsceo.l rbuWiiedlnhein ciacgech? ct hoidef e tpnhete,r iafao m1ll0oy-wgydeinaarglo- soideldn, psboiorriyyf o sdruemfffie,c rietsd qyouuers tcioomnsp hreehlpe nysoiuo nassess
• Provides succinct coverage of every important topic without sacrificing otcholfera tcaetnxot,r eayrmi sotyrrg iodal afdaliacr,te oacrntyldy n etuonc ttloherueh soi,n lofaallfc actcootrrotyer yxc o.trutbeexr, cilnec, lpuidriifnogr m CAB... HAVuiysdpuieatrloo drsyims ptuiraorbbalenmces
comprehensiveness or readability ■ Taste buds are the specialized sense organs for taste and are D. Taste and odor abnormalities
composed of basal stem cells and three types of taste cells (dark, E. No major sensory deficits
lriegphrte, saenndt vinatreiormuse sdtiaagtee)s. oThf dei fftherreeen ttiyaptieosn o of ft adsetve ecleolplsi nmga tya ste 5. Which of the following are incorrectly paired?
• Reflects the latest research and developments in the areas of chronic pain, cloecllast,e wdi tinh tthhee mliguhcto cseal los fb tehine ge pthigel omttoiss,t p malaattuer, ea.n Tda spthe abruydnsx aarned AB.. GENusatCd u: cSionu :r B tiatsteter taste
in the walls of papillae of the tongue. C. T1R3 family of GPCRs : Sweet taste
reproductive physiology, and acid-base homeostasis ■ Thumearme ai.r eS itganstael trreacnepsdtourcst ifoonr smweeceht,a snoisumr, sb iinttcelru, dsael tp, aasnsadg e DE.. HEbenscehr egll asunldcsu :s T: aSsmtee allcuity
through ion channels, binding to and blocking ion channels, 6. A 9-year-old boy had frequent episodes of uncontrollable nose
and GPCRs requiring second messenger systems. bleeds. At the advice of his clinician, he underwent surgery
• Ipnhcyosripoolorgatice sc oenxcaempptsles from clinical medicine to illustrate important ■ Thtimonhf e etted hh saieeeaff v ltNe ehlrneTaemtlnSha.tnm, s Fin sufrircosnou,mt msah nt,t othdaa en tsorhdtenee , tt beovaunxe tndtohhtsner s ciaan rlana spttncheoiereasni tlote donrr n evoignrimavus eueetshd lt aeitr ao aail vpn snesydulin lfvcaarlitpoeaeuns reast a l l tsraAooub. lr oclgAsuoe rstnr hroy eoe,lc f lhwafta ecaac t tstopo orbrlrydoay b kt hsrlieineasmnng mss iominonr ti ythhsh sneieisroe o nunhvar?eeos ncanol. eusWexldpph trinuecomshst e.os sAfm at ehf welelw itfd hodele al roycawisnn iagnnfteag emo rifs ot htnre u e
operculum in the ipsilateral cerebral cortex. odorant receptors.
B. Lateral inhibition within the olfactory glomeruli reduces
the ability to distinguish between different types of odorant
• Section introductions help you build a solid foundation on the given topic MULTIPLE-CHOICE QUESTIONS receptors.
C. Conscious discrimination of odors is dependent on the
For all questions, select the single best answer unless otherwise pathway to the orbitofrontal cortex.
directed. D. Olfaction is closely related to gustation because odorant and
• Includes both end-of-chapter and board-style review questions 1. A young boy was diagnosed with congenital anosmia, a rare gustatory receptors use the same central pathways.
disorder in which an individual is born without the ability to E. All of the above.
smell. Odorant receptors are 7. A 31-year-old female is a smoker who has had poor oral
A. located in the olfactory bulb. hygiene for most of her life. In the past few years she has
• Chapter summaries ensure B. located on dendrites of mitral and tufted cells. noticed a reduced sensitivity to the flavors in various foods
C. located on neurons that project directly to the olfactory which she used to enjoy eating. Which of the following is not
cortex. true about gustatory sensation?
retention of key concepts D. located on neurons in the olfactory epithelium that project A. The sensory nerve fibers from the taste buds on the anterior
to mitral cells and from there directly to the olfactory cortex. two-thirds of the tongue travel in the chorda tympani
E. located on sustentacular cells that project to the olfactory bulb. branch of the facial nerve.
CHAPTER 37 Renal Function & Micturition 673
• More clinical cases and flow charts
than ever, along with modern A Proximal tubuleCapsule B Podocyte
Red blood cells
approaches to therapy Mesangial Glalmominearular basal Barrett_CH11_p217-226.indd 225 5/26/15 2:00 PM 132 SECTION I Cellular and Molecular Basis for Medical Physiology
cell Bowman’s space Capillary
• Expanded legends for each Capillary CLINICAL BOX 6–2
Granular cells
Myasthenia Gravis arthritis, systemic lupus erythematosus, and polymyositis.
irlelufestrr bataicokn t—o stoh ey otuex dton’t have to PproodcoEaecrfstfyeseterreiesonlet NerAavrfetfee frribieoenlerts Capillary Capillary PpBlaroamodsciaonelcasyste Mwtaoonchy cy1iau c2ashr5grt eh seonke febcn eleuiesavtt e asgirnle rym ea i1mnvu idsmssci vtiliseolild si aouh ansaare lvespre iweo inaoue psabt lkhaiem ne awidonr odds2rao l0tldmi srdwe ei( sitmedtiamreaisb ieianlusynl .tyf diI aot wt ncaoao,lc nmwdc uioiestrechnsca )ip usnaere n 2aaidn5kt Anspau arbregloo dgrlueeiest lsp ain3tto 0i tvsth%hietaei towo ipnnfi t adphttoiha va toiaidnegu uneataotnusliset m owswiimsmiitt huohm nf mm uethny ydeaaei ss sdtdtehhiiasseeesonnearii.dsaa eTe g ghrbr.re ayaT v vhtsihiseusy s spmhehp aaualvyrsseei sn maoag cg a imheyae ntapileotpletnaeirycsr-
60s (mainly men). It is caused by the formation of circulating T cells sensitized against thymic proteins that cross-react with
• Introductory materials cover key Distal tubuleMacula denSsamooth muscle Mesangial cell Cenydtoopthlaeslimal of taonrtsib. oThdeiesse taon tthibeo mdiuessc dlee sttyrpoey osof mniec ootfi nthice crehcoelpinteorrsg iacn rde bceinpd- apclaesttyilcc; haonldin 1e0 r–e1c5e%pt ohrasv. eIn a mthoysmt opmatiae.nts, the thymus is hyper-
cell others to neighboring receptors, triggering their removal by
principles of endocrine regulation endocytosis. Normally, the number of quanta released from THERAPEUTIC HIGHLIGHTS
in physiology CPodoEcnydteothBealisuaml lamina Basal lamina Endothelium DFenestrations Fofo poFoti ltdprsBorapooctaicwyoectnmees sssaelnits’s tsacoItncTasplehinhhtylantphnreaoii n eennmevepsdotsr ahsei et epnscbmu.sampp e eaaelyaelnaItseciorl nn aci. etadfgo actojefoaI noo tenrnsl pmreecaocrrai edmnlmtd aewtevei sn7 tuc ,ffet ayeoa 0 eow trrtlrmffacrsef–urehre e stvme t9svce eiithaoepncve0tch m,k ufaeti a l%aet tets tnxrryasbteh nhdas the .onrre nerdile Tmasea fadmr,o e ,ah n eo mq sgicrdbreuan ehcmeurriinrlo ssuaeaasslhaec oatelvapllan aalolmlraaisoedi sstrmrrtegrsswa , eua ee ema .eelna tcsn,i , Po nntn lcrreluadiiwmeearle ntdunseeytdltyroecepriudhs na ueofls a,lise eal cmann cmaisci zisywtenteb lln eaeu suemda.tcinj ud. tr os orTl woehyfumre cS hrase aar ui td akttimpssfbuslslihnioseupatgnr edoah drllircougrememim lece lthnetad yoernsyaastas dae sfsh ilrwwev nas osasiame rleh i tiisftytevtdt gh omhoo hrtucedear h hwe ec saaiiff teenssaracopcv heesuu plniteritedeaipcsh soomh ta ietteyarwsgnmriebansasltater cidsi e tsfnagaathptteao.auijvs ehhmvnooieIikeeliddneeesstrr--,.. Matpmfi(sittnianmnoehhyufniytggadureyelpoper c ,ttm usngip trhrhacdrch ioprrebaoloeaeaoeeevrp o m lvsns seergpie ls intwo di4sarera simae.aerg5 ner mtimEvnsiem%smaiuovits d tskassdeoeii oi. sobnucl nrn f ssndlTfoo aeseeit aeipophesfsdin .mc s, enceaysyr e ttlec t m uaithdrpinCistieyezlnneoutareehlnna o dchsetc ootg3itihed ntioshrli5tb ootui.n wnhniro%ailmoc e.ti efti ntltopi thsmeIinyheaeomrtrea rone iny is sanmsdursdaao ena aeuetds dmisud n,tncivtme mhnhn hdeedo ceyie ol pianrohcnpmua sriniapaniaoscsura tovom ttyhvtep riemnhtceeiagdperb luesntbar aonrii aosenttoe,etss o svsts senypsc ohiprmtnssomw soifyeo ig m ivrmismipcfttmmih etihnptpaaeey oo pieenlrrcadnmlwmesnry)t sr oevao dtsnyuschei vcam auf ageneot etisrnnoeaysstr----
Capillary lumen Clinical cases add CLINICAL BOX 6–3
Basal lamina
Lambert–Eaton Syndrome prostate, bladder, kidney, or gallbladder. Clinical signs usually
atgFonlIo dtGmh peUeo rcduRaolpaEcirly lp3atero7ysly –etaon2n dgi ooloSntmht.reuelricuutmluarr.a Dcla )d pEeinltllaaarirliges seo.m fC ge) nlDotem otafe itrlh uoelfu rtesh.ce At aw)n Sagyel cept oiinod nCo t cthyort osehus ogfowhr mvtha sfiec lpturolaadtri oopcnoy lstelei,t spsh rooonwc etihsnsege bsc.aa Tsphailell al afrumyz lzionyoa m,p asan.t Bder) ti Rahlee ol arnet iltoahtneio oirnf s moufre ftashacene lsga iimas li nceal ls rteoa tlh-we oterlxdt relevance Itatcnihhuo eatnaonn. iniTmrceeh mllsiassy tu nidivnndeee crl tyrohae metarta saernec ese(k Lr tcvEhaoeMegn adSenin)inot,d isrommti nnuaog lsncs cC aelalea lte2 o +wdf t i hentLaeflhak uemnn xee vbtsuohesrl aortitatsm– gccEueaaas-uugtcsosuaeenltsade rad m bc juyeyC tnaaaycs2nl+--- pcaalrsenoTc oeiHmcdceEpu aRtrih raA eCf tPade2Eri+a tUcghhneTao IunsCsnise e H olo ffIfu GcanamHcntciiLnoeInorG. .gAHly scTyonSsdidroem aen tsiibmioilatirc sto, wLhEMichS
More than agbivoeunt t1i2m me 2i.s Th30e– 4vo0 lmumL.e of blood in the renal capillaries at any gplroomloenrgu alanru firlitar ainti oancu rtaet ek (idGnFeRy) i annjudr iys (cAlaKimI)e.d to enhance and cadhbisooeluaints ee1 trchealaestae as epp.pe Tre ha1er0s 0i nt,o0c0 idh0ea nvpece eoa p oslfie mL; EiiltMa irsS o uicnsc uutahrlreley nU acnnei tiaendd mu Sltte-anot enassn eidst Sctheiner,c teihn tedh iefivrirdes tui sat rale haailtgsmoh e chnoatm ss tocrarabtneicdgeiytry ia swn tidoth, d isfem tseaorl,ml ctioenl le tl ruwenahgte ctthhaneart-
women. Proximal muscles of the lower extremities are primar- appropriately. In patients without cancer, immunother-
600 full-color LYMPHATICS INNERVATION OF itlhye a affremcste. dR,e pperotidtiuvcei nstgi mau wlaatidodnl inofg thgea itm aontodr dnieffirvcue ltfayc rilaitisaitnegs apphye reiss isin, iatinadte idn. trParveednnoisuosn iem madumnoingislotrbatuiolinn, aprela ssommae-
The kidneys have an abundant lymphatic supply that drains THE RENAL VESSELS accumulation of Ca2+ in the nerve terminal and increases ace- examples of effective therapies for LEMS. Also, the use of
via the thoracic duct into the venous circulation in the thorax. tylcholine release, leading to an increase in muscle strength. aminopyridines facilitates the release of acetylcholine
illustrations The renal nerves travel along the renal blood vessels as they This is in contrast to myasthenia gravis in which symptoms are in the neuromuscular junction and can improve muscle
CAPSULE enter the kidney. They contain many postganglionic sym- exacerbated by repetitive stimulation. About 40% of patients strength in LEMS patients. This class of drugs causes
pathetic efferent fibers and a few afferent fibers. There also with LEMS also have cancer, especially small cell cancer of the blockade of presynaptic K+ channels and promote activa-
Thepdreeesm sruaertneoa u(l rsc,e antphasleu ilnceta episrs sutthlietii nal ilbm puritte sst ostuuhrgee h)s. wrIifes eltlsih.n eThg ,k iiasdn nddee cytr hebeae scetoissm stuheees abopunptic ei atisrn sn ftueorn vbcaetti ioao ncn h icoso liumnneecrseg rpictra iiimnnn.a erThrilvyea tfirsooynmm vp itaaht ehth eleoti wvc aepgru rteshg noaernargcvlieic-, latmuotnta LalgiEcg.kM On taSnhn.e eLt t EtchhMaeynoSmc rheyoar im ssc taeah;ll slaasot n mabdne actyeaib nnao clasdesoirse ooasct ftt ihtaahtaceetkd hbC awraev2iaet+hs bct ,lhe yseamtnnopn mpheralooscsd,h alu,er cccaeooddlmion tnago;, tisniyohmnib poittfo ovmrosl stc aoagfn eL -EbgMea tSue.sde dC ab2+u ct hoafntenne lds.o A cneotty lacmhoelliinoerasttee rtahsee
Barrett_CH37_p669-694.indd 673 6/18/15 4:54 PM Barrett_CH06_p121-136.indd 132 5/15/15 4:18 PM
Barrett_FM_i-xii_P1.indd 4 6/30/15 1:20 PM
Key Features of the Twenty-Fifth Edition of
CHAPTER 11 Smell & Taste 225
Ganong’s Review of Medical Physiology that Taraes tsei mexilhairb iints saoftmeer wreaaycst itoon vsi saunadl acftoenrt irmasat gpesh eannodm ceonna- 2. AO n3e7 -oyfe tahre- oplodt feenmtiaalle c wonasse dqiuaegnncoesse do fw thitihs dmisuoltridpeler sisc lerosis.
trasts. Some of these are chemical “tricks,” but others may be diminished taste sensitivity. Taste receptors
true central phenomena. A taste-modifier protein, miraculin, A. for sweet, sour, bitter, salt, and umami are spatially separated
has been discovered in a plant. When applied to the tongue, on the surface of the tongue.
this protein makes acids taste sweet. B. are synonymous with taste buds.
A concise, up-to-date and clinically relevant review noavfe esarsvs.Ao iThoindniiemsn sgtauo lprs nv,o ioivisvnaoeclnl lv sufa.odlouindeg so fihf s uuemacthian nagvs ,e trhfsoeio rfmnoso idps aiasrp tfpiocaulrlloeanwrtle yid n s btteyrro minllgs- 3. CDEW... haagAicrrlloeehl s oaison foft n ytpthephhreeeav aroafybotfen olcldgvoh eewbea.myiln anogffer derevcroeeeenpss.tt nso ori.nt itnhcer efaacsiea tl,h ter iagbeimlitiyn atol, and
discriminate many different odors?
CHAPTER SUMMARY A. Many different receptors
B. Pattern of olfactory receptors activated by a given
of human physiology ■ Oawniltdfha bcintao strahyle ss uteepnmpso ecrrey pl lnos erautrieroo nlno osc,fa stteuhdpe p innoa rtsthainel cgoa l(vfsaiutcyst.toernyt aecpuitlhare)li cuemlls , C. oPthdreoo jbrearcnatiitnon of different mitral cell axons to different parts of End-of-chapter review
■ The cilia located on the dendritic knob of the olfactory D. High β-arrestin content in olfactory neurons
■ sGdfIonee-rnfnpomsdrroo rmoritytleaef iatnsnic eosotu.nof Ar rmfoyrxno iogt mrcnloaos lmtn ohatefnea r oidounl lflt fiaoua.cdftctoteoordrray ycn setbel rlunsel scbione rtp rtytha onveree solu stlrf hvaoaicnatt s oat hcrreyoe n cblaotuatulecbprts alte lht doe t o 4. EAdaisna. s hmd aSe ean rmngeiffietsoou tsnrlothtg ltioinhkf aeea llnb ycr oataourint teio cximenpsceo.l rbuWiiedlnhein ciacgech? ct hoidef e tpnhete,r iafao m1ll0oy-wgydeinaarglo- soideldn, psboiorriyyf o sdruemfffie,c rietsd qyouuers tcioomnsp hreehlpe nysoiuo nassess
• Provides succinct coverage of every important topic without sacrificing otcholfera tcaetnxot,r eayrmi sotyrrg iodal afdaliacr,te oacrntyldy n etuonc ttloherueh soi,n lofaallfc actcootrrotyer yxc o.trutbeexr, cilnec, lpuidriifnogr m ABC... VHAuiysdpuieatrloo drsyims ptuiraorbbalenmces
comprehensiveness or readability ■ Taste buds are the specialized sense organs for taste and are D. Taste and odor abnormalities
composed of basal stem cells and three types of taste cells (dark, E. No major sensory deficits
lriegphrte, saenndt vinatreiormuse sdtiaagtee)s. oThf dei fftherreeen ttiyaptieosn o of ft adsetve ecleolplsi nmga tya ste 5. Which of the following are incorrectly paired?
• Reflects the latest research and developments in the areas of chronic pain, cloecllast,e wdi tinh tthhee mliguhcto cseal los fb tehine ge pthigel omttoiss,t p malaattuer, ea.n Tda spthe abruydnsx aarned AB.. EGNusatCd u: cSionu :r B tiatsteter taste
in the walls of papillae of the tongue. C. T1R3 family of GPCRs : Sweet taste
reproductive physiology, and acid-base homeostasis ■ Thumearme ai.r eS itganstael trreacnepsdtourcst ifoonr smweeceht,a snoisumr, sb iinttcelru, dsael tp, aasnsadg e DE.. HEbenscehr egll asunldcsu :s T: aSsmtee allcuity
through ion channels, binding to and blocking ion channels, 6. A 9-year-old boy had frequent episodes of uncontrollable nose
and GPCRs requiring second messenger systems. bleeds. At the advice of his clinician, he underwent surgery
• Ipnhcyosripoolorgatice sc oenxcaempptsles from clinical medicine to illustrate important ■ Thtimonhf e etted hh saieeeaff v ltNe ehlrneTaemtlnSha.tnm, s Fin sufrircosnou,mt msah nt,t othdaa en tsorhdtneee , tt beovaunxe tndtohhtsner s ciaan rlana spttncheoiereasni tlote donrr n evoignrimavus eueetshd lt aeitr ao aail vpn snesydulin lfvcaarlitpoeaeuns reast a l l tsraAooub. lr oclgAsuoe rstnr hroy eoe,lc f lhwafta ecaac t tstopo orbrlrydoay b kt hsrlieineasmnng mss iominonr ti ythhsh sneieisroe o nunhvar?eeos ncanol. eusWexldpph trinuecomshst e.os sAfm at ehf welelw itfd hodele al roycawisnn iagnnfteag emo rifs ot htnre u e
operculum in the ipsilateral cerebral cortex. odorant receptors.
B. Lateral inhibition within the olfactory glomeruli reduces
the ability to distinguish between different types of odorant
• Section introductions help you build a solid foundation on the given topic MULTIPLE-CHOICE QUESTIONS receptors.
C. Conscious discrimination of odors is dependent on the
For all questions, select the single best answer unless otherwise pathway to the orbitofrontal cortex.
directed. D. Olfaction is closely related to gustation because odorant and
• Includes both end-of-chapter and board-style review questions 1. A young boy was diagnosed with congenital anosmia, a rare gustatory receptors use the same central pathways.
disorder in which an individual is born without the ability to E. All of the above.
smell. Odorant receptors are 7. A 31-year-old female is a smoker who has had poor oral
A. located in the olfactory bulb. hygiene for most of her life. In the past few years she has
• Chapter summaries ensure B. located on dendrites of mitral and tufted cells. noticed a reduced sensitivity to the flavors in various foods
C. located on neurons that project directly to the olfactory which she used to enjoy eating. Which of the following is not
cortex. true about gustatory sensation?
retention of key concepts D. located on neurons in the olfactory epithelium that project A. The sensory nerve fibers from the taste buds on the anterior
to mitral cells and from there directly to the olfactory cortex. two-thirds of the tongue travel in the chorda tympani
E. located on sustentacular cells that project to the olfactory bulb. branch of the facial nerve.
CHAPTER 37 Renal Function & Micturition 673
• More clinical cases and flow charts
than ever, along with modern A Proximal tubuleCapsule B Podocyte
Red blood cells
approaches to therapy Mesangial Glalmominearular basal Barrett_CH11_p217-226.indd 225 5/26/15 2:00 PM 132 SECTION I Cellular and Molecular Basis for Medical Physiology
cell Bowman’s space Capillary
• Expanded legends for each Capillary CLINICAL BOX 6–2
Granular cells
Myasthenia Gravis arthritis, systemic lupus erythematosus, and polymyositis.
irlelufestrr bataicokn t—o stoh ey otuex dton’t have to PproodcoEaecrfstfyeseterreiesonlet NerAavrfetfee frribieoenlerts Capillary Capillary PpBlaroamodsciaonelcasyste Mwtaoonchy cy1iau c2ashr5grt eh seonke febcn eleuiesavtt e asgirnle rym ea i1mnvu idsmssci vtiliseolild si aouh ansaare lvespre iweo inaoue psabt lkhaiem ne awidonr odds2rao l0tldmi srdwe ei( sitmedtiamreaisb ieianlusynl .tyf diI aot wt ncaoao,lc nmwdc uioiestrechnsca )i pusnaere n 2aaidn5kt Anspau arbregloo dgrlueeiest lsp ain3tto 0i tvsth%hietaei towo ipnnfi t adphttoiha va toiaidnegu uneataotnusliset m owswiimsmiitt huohm nf mm uethny ydeaaei ss sdtdtehhiiasseeesonnearii.dsaa eTe g ghrbr.re ayaT v vhtsihiseusy s spmhehp aaualvyrsseei sn maoag cg a imheyae ntapileotpletnaeirycsr-
60s (mainly men). It is caused by the formation of circulating T cells sensitized against thymic proteins that cross-react with
• Introductory materials cover key Distal tubuleMacula denSsamooth muscle Mesangial cell Cenydtoopthlaeslimal of atonrtsib. oThdeiesse taon tthibeo mdiuessc dlee sttyrpoey osof mniec ootfi nthice crehcoelpintoerrsg iacn rde bceinpd- apclaesttyilcc; haonldin 1e0 r–e1c5e%pt ohrasv. eIn a mthoysmt opmatiae.nts, the thymus is hyper-
cell others to neighboring receptors, triggering their removal by
principles of endocrine regulation endocytosis. Normally, the number of quanta released from THERAPEUTIC HIGHLIGHTS
in physiology CPodoEcnydteothBealisuaml lamina Basal lamina Endothelium DFenestrations Fofo poFoti ltdprBsorapooctaicwyoectnmees sssaelnits’s tsacoItncTasplehinhhtylantphnreaoii n eennmevepsdotsr ahsei et epnscbmu.sampp e eaaelyaelnaItseciorl nn aci. etadfgo actojefoaI noo tenrnsl pmreecaocrrai edmnlmtd aewtevei sn7 tuc ,ffet ayeoa 0 eow trrtlrmffacrsef–urehre e stvme t9svce eiithaoepncve0tch m,k ufaeti a l%aet tets tnxrryasbteh nhdas the .onrre nerdile Tmasea fadmr,o e ,ah n eo mq sgicrdbreuan ehcmeurriinrlo ssuaeaasslhaec oatelvapllan aalolmlraaisoedi sstrmrrtegrsswa , eua ee ema .eelna tcsn,i , Po nntn lcrreluadiiwmeearle ntdunseeytdltyroecepriudhs na ueofls a,lise eal cmann cmaisci zisywtenteb lln eaeu suemda.tcinj ud. tr os orTl woehyfumre cS hrase aar ui td akttimpssfbuslslihnioseupatgnr edoah drllircougrememim lece lthnetad yoernsyaastas dae sfsh ilrwwev nas osasiame rleh i tiisftytevtdt gh omhoo hrtucedear h hwe ec saaiiff teenssaracopcv heesuu plniteritedeaipcsh soomh ta ietteyarwsgnmriebansasltater cidsi e tsfnagaathptteoa.auijvs ehhmvnooieIikeeliddneeesstrr--,.. i(sittniapmfMatnmnoehhyunfiygtgadureyelpoper c ,ttm usngip trhrhacdrch ioprrebaoloeaeaoeeevrp o m lvsnss eergpie ls intwo di4sarera simae.aerg5 ner mtimEvnsiem%smaiuovits d tskassdeoeii oi. sobnucl nrn f ssndlTfoo aeseeit aeipophesfsdin .mc s, enceasyyr e ttlec t m uaithdrpinCistieyezlnneoutareehlnna o dchsetc ootg3itihed n tioshrli5tb ootui.n wnhniro%ailmoc e.ti efti nttlopi thsmeIinyheaeomrtrea rone iny is sanmsdursdaao ena aeuetds dmisud n,tncivtm e mhnhn hdeedo ceyie ol pianrohcnpmua sriniapaniaoscsura tovom tythvtep riemnhtceeiagdperb luesntbar aonrii aosenttoe,etss o svsts senypsc ohiprmtnssomw soifyeo ig m ivrmismpicfttmmih etihnptpaaeey oo pieenlrrcadnmlwmesnyr)t sr oevao dtsnyuschie vcam auf ageneto etisrnnoeaysstr----
Capillary lumen Clinical cases add CLINICAL BOX 6–3
Basal lamina
Lambert–Eaton Syndrome prostate, bladder, kidney, or gallbladder. Clinical signs usually
atgFonlIo dtGmh peUeo rcduRaolpaEcirly lp3atero7ysly –etaon2n dgi ooloSntmht.reuelricuutmluarr.a Dcla )d pEeinltllaaarirliges seo.m fC ge) nlDotem otafe itrlh uoelfu rtesh.ce At aw)n Sagyel cept oiinod nCo t cthyort osehus ogfowhr mvtha sfiec lpturolaadtri oopcnoy lstelei,t spsh roooncw etihsnsege bsc.aa Tsphailell al afrumyz lzionyoa m,p asan.t Bder) ti Rahlee ol arnet iltoahtneio oirnf s moufre ftashacene lsga iimas li nceal ls rteoa tlh-we oterlxdt relevance Itactnihhuo eatnaonn. iniTmrceeh mllsiassy tu nidivnndeee crl tyrohae metarta saernec ese(k Lr tcvEhaoeMegn adSenin)inot,d isrommti nnuaog lsncs cC aelalea lte2 o +wdf t i hentLaeflhak uemnn xee vbtsuohesrl aortitatsm– gccEueaaas-uugtcsosuaeenltsade rad m bc juyeyC tnaaaycs2nl+--- pcaalrsenoTc oeiHmcdceEpu aRtrih raA eCf tPade2Eri+a tUcghhneTao IunsCsnise e H olo ffIfu GcanamHcntciiLnoeInorG. .gAHly scTyonSsdidroem aen tsiibmioilatirc sto, wLhEMichS
More than agbivoeunt t1i2m me 2i.s Th30e– 4vo0 lmumL.e of blood in the renal capillaries at any gplroomloenrgu alanru firlitar ainti oancu rtaet ek (idGnFeRy) i annjudr iys (cAlaKimI)e.d to enhance and cadhbisooeluaints ee1 trchealaestae as epp.pe Tre ha1er0s 0i nt,o0c0 idh0ea nvpece eoa p oslfie mL; EiiltMa irsS o uicnsc uutahrlreley nU acnnei tiaendd mu Sltte-anot enassn eidst Sctheiner,c teihn tedh iefivrirdes tui sat rale haailtgsmoh e chnoatm ss tocrarabtneicdgeiytry ia swn tidoth, d isfem tseaorl,ml ctioenl le tl ruwenahgte ctthhaneart-
women. Proximal muscles of the lower extremities are primar- appropriately. In patients without cancer, immunother-
600 full-color LYMPHATICS INNERVATION OF itlhye a affremcste. dR,e pperotidtiuvcei nstgi mau wlaatidodnl inofg thgea itm aontodr dnieffirvcue ltfayc rilaitisaitnegs apphye reiss isin, iatinadte idn. trParveednnoisuosn iem madumnoingislotrbatuiolinn, aprela ssommae-
The kidneys have an abundant lymphatic supply that drains THE RENAL VESSELS accumulation of Ca2+ in the nerve terminal and increases ace- examples of effective therapies for LEMS. Also, the use of
via the thoracic duct into the venous circulation in the thorax. tylcholine release, leading to an increase in muscle strength. aminopyridines facilitates the release of acetylcholine
illustrations The renal nerves travel along the renal blood vessels as they This is in contrast to myasthenia gravis in which symptoms are in the neuromuscular junction and can improve muscle
CAPSULE enter the kidney. They contain many postganglionic sym- exacerbated by repetitive stimulation. About 40% of patients strength in LEMS patients. This class of drugs causes
pathetic efferent fibers and a few afferent fibers. There also with LEMS also have cancer, especially small cell cancer of the blockade of presynaptic K+ channels and promote activa-
Thepdreeesm sruaertneoa u(l rsc,e antphasleu ilnceta episrs sutthlietii nal ilbm puritte sst ostuuhrgee h)s. wrIifes eltlsih.n eThg ,k iiasdn nddee cytr hebeae scetoissm stuheees abopunptic ei atisrn sn ftueorn vbcaetti ioao ncn h icoso liumnneecrseg rpictra iiimnnn.a erThrilvyea tfirsooynmm vp itaaht ehth eleoti wvc aepgru rteshg noaernargcvlieic-, latmuotnta LalgiEcg.kM On taSnhn.e eLt t EtchhMaeynoSmc rheyoar im ssc taeah;ll slaasot n mabdne actyeaib nnao clasdesoirse ooasct ftt ihtaahtaceetkd hbC awraev2iaet+hs bct ,lhe yseamtnnopn mpheralooscsd,h alu,er cccaeooddlmion tnago;, tisniyohmnib poittfo ovmrosl stc aoagfn eL -EbgMea tSue.sde dC ab2+u ct hoafntenne lds.o A cneotty lacmhoelliinoerasttee rtahsee
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About the Authors
KIM E. BARRETT and an MSU College of Human Medicine Distinguished Faculty
Award. She has been very active in the American Physiologi-
Kim Barrett received her PhD in biological
cal Society (APS) and served as its 85th President. She has also
chemistry from University College London
served as a Councillor as well as Chair of the Central Nervous
in 1982. Following postdoctoral training at
System Section of APS, Women in Physiology Committee and
the National Institutes of Health, she joined
Section Advisory Committee of APS. She is also active in the
the faculty at the University of California, San
Michigan Physiological Society, a chapter of the APS.
Diego, School of Medicine in 1985, rising to
the rank of Professor of Medicine in 1996, SCOTT BOITANO
and was named Distinguished Professor of
Scott Boitano received his PhD in
Medicine in 2015. Since 2006, she has also
genetics and cell biology from Washing-
served the University as Dean of the Graduate Division. Her
ton State University in Pullman, Wash-
research interests focus on the physiology and pathophysiology
ington, where he acquired an interest
of the intestinal epithelium, and how its function is altered by
in cellular signaling. He fostered this
commensal, probiotic, and pathogenic bacteria as well as in
interest at University of California, Los
specific disease states, such as inflammatory bowel diseases. She
Angeles, where he focused his research
has published more than 200 articles, chapters, and reviews, and
on second messengers and cellular
has received several honors for her research accomplishments
physiology of the lung epithelium. How
including the Bowditch and Davenport Lectureships from the
the airway epithelium contributes to lung health has remained a
American Physiological Society and the degree of Doctor of
central focus of his research at the University of Wyoming and
Medical Sciences, honoris causa, from Queens University, Belfast.
in his current positions with the Departments of Physiology and
She has been very active in scholarly editing, serving currently
Cellular and Molecular Medicine, the Arizona Respiratory Center
as the Deputy Editor-in-Chief of the Journal of Physiology. She
and the Bio5 Collaborative Research Institute at the University of
is also a dedicated and award-winning instructor of medical,
Arizona. Dr. Boitano remains an active member of the American
pharmacy, and graduate students, and has taught various topics
Physiological Society and served as the Arizona Chapter’s Presi-
in medical and systems physiology to these groups for more than
dent from 2010–2012.
20 years. Her efforts as a teacher and mentor were recognized
with the Bodil M. Schmidt-Nielson Distinguished Mentor and HEDDWEN L. BROOKS
Scientist Award from the American Physiological Society (APS)
in 2012, and she also served as the 86th APS President from Heddwen Brooks received her PhD from
2013–14. Her teaching experiences led her to author a prior Imperial College, University of London and
volume (Gastrointestinal Physiology, McGraw-Hill, 2005; second is a Professor in the Departments of Physi-
edition published in 2014) and she was honored to have been ology and Pharmacology at the University
invited to take over the helm of Ganong in 2007 for the 23rd and of Arizona (UA). Dr Brooks is a renal phys-
subsequent editions, including this one. iologist and is best known for her develop-
ment of microarray technology to address
SUSAN M. BARMAN
in vivo signaling pathways involved in the
hormonal regulation of renal function. Dr
Susan Barman received her PhD in physiol-
Brooks’ many awards include the American Physiological Society
ogy from Loyola University School of Med-
(APS) Lazaro J. Mandel Young Investigator Award, which is for
icine in Maywood, Illinois. Afterward she
an individual demonstrating outstanding promise in epithelial
went to Michigan State University (MSU)
or renal physiology. In 2009, Dr Brooks received the APS Renal
where she is currently a Professor in the
Young Investigator Award at the annual meeting of the Federation
Department of Pharmacology/Toxicology
of American Societies for Experimental Biology. Dr Brooks served
and the Neuroscience Program. Dr Barman
as Chair of the APS Renal Section (2011–2014) and currently
has had a career-long interest in neural con-
serves as Associate Editor for the American Journal of Physiology-
trol of cardiorespiratory function with an
Regulatory, Integrative and Comparative Physiology and on the
emphasis on the characterization and origin
Editorial Board for the American Journal of Physiology-Renal
of the naturally occurring discharges of sympathetic and phrenic
Physiology (since 2001). Dr Brooks has served on study sections
nerves. She was a recipient of a prestigious National Institutes of
of the National Institutes of Health, the American Heart Associa-
Health MERIT (Method to Extend Research in Time) Award. She
tion and recently was a member of the Nephrology Merit Review
is also a recipient of an Outstanding University Woman Faculty
Board for the Department of Veterans’ Affairs.
Award from the MSU Faculty Professional Women’s Association
vii
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Contents
Preface xi
13 Autonomic Nervous System 255
S E C T I O N Cellular & 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 33 S E C T I O N
Endocrine & Reproductive
III
3 Immunity, Infection, & Inflammation 67 Physiology 297
4 Excitable Tissue: Nerve 85
16 Basic Concepts of Endocrine
5 Excitable Tissue: Muscle 99
Regulation 299
6 Synaptic & Junctional Transmission 121
17 Hypothalamic Regulation of Hormonal
Functions 307
7 Neurotransmitters & Neuromodulators 137
18 The Pituitary Gland 321
19 The Thyroid Gland 337
S E C T I O N
Central & Peripheral
II
Neurophysiology 157 20 The Adrenal Medulla & Adrenal Cortex 351
21 Hormonal Control of Calcium, & Phosphate
Metabolism & the Physiology of Bone 375
8 Somatosensory Neurotransmission:
Touch, Pain, & Temperature 159
22 Reproductive Development & Function of the
Female Reproductive System 389
9 Vision 177
23 Function of the Male Reproductive
10 Hearing & Equilibrium 199
System 417
11 Smell & Taste 217
24 Endocrine Functions of the Pancreas
12 Reflex & Voluntary Control of & Regulation of Carbohydrate
Posture & Movement 227 Metabolism 429
ix
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