Human Anatomy & Physiology Ninth Edition Elaine N. Marieb, R.N., Ph.D. Holyoke Community College Katja Hoehn, M.D., Ph.D. Mount Royal University Boston Columbus Indianapolis New York San Francisco Upper Saddle River Amsterdam Cape Town Dubai London Madrid Milan Munich Paris Montreal Toronto Delhi Mexico City São Paulo Sydney Hong Kong Seoul Singapore Taipei Tokyo # 105016 Cust: Benjamin Cummings/CA Au: Marieb Pg. No. i C/M/Y/K cDEaSIGrN SlERiVIsCESl OeF Title: Anatomy & Physiology Server: S4C Short / Normal Publishing Services Editor-in-Chief: Serina Beauparlant Cover Designer: Riezebos Holzbaur Design Group Acquisitions Editor: Gretchen Puttkamer Art Houses: Imagineering STA Media Services Inc. and Electronic Associate Project Editor: Shannon Cutt Publishing Services Inc., NYC Director of Development: Barbara Yien Art Coordinator: Jean Lake Development Editor: Alice Fugate Photo Image Lead: Donna Kalal Art Development Manager: Laura Southworth Photo Researcher: Kristin Piljay Senior Managing Editor: Debbie Cogan Copyeditor: Anita Wagner Production and Design Manager: Michele Mangelli Proofreader: Martha Ghent Production Supervisor: David Novak Compositor: S4Carlisle Publishing Services Media Producer: Aimee Pavy Senior Manufacturing Buyer: Stacey Weinberger Editorial Assistant: Lisa Damerel Marketing Manager: Derek Perrigo Text Designer: tani hasegawa Cover photo of Olympic Gold Medalist Hope Solo © Annie Leibovitz/Contact Press Images. 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Where those designations appear in this book, and the publisher was aware of a trademark claim, the designations have been printed in initial caps or all caps. Library of Congress Cataloging-in-Publication Data Marieb, Elaine Nicpon Human anatomy & physiology / Elaine N. Marieb, Katja Hoehn.—9th ed. p. ; cm. ISBN-13: 978-0-321-74326-8 (student ed.) ISBN-10: 0-321-74326-1 (student ed.) I. Hoehn, Katja. II. Title. [DNLM: 1. Anatomy. 2. Physiological Phenomena. QS 4] LC classification not assigned 612—dc23 2011038702 ISBN 10: 0-321-74326-1; ISBN 13: 978-0-321-74326-8 (Student edition) ISBN 10: 0-321-80217-9; ISBN 13: 978-0-321-80217-0 (Instructor’s Review Copy) 1 2 3 4 5 6 7 8 9 10—RRD—15 14 13 12 11 www.pearsonhighered.com # 105016 Cust: Benjamin Cummings/CA Au: Marieb Pg. No. ii C/M/Y/K cDEaSIGrN SlERiVIsCESl OeF Title: Anatomy & Physiology Server: S4C Short / Normal Publishing Services About the Authors We dedicate this work to our students both present and past, who always inspire us to “push the envelope.” Elaine N. Marieb For Elaine N. Marieb, taking the student’s perspective into ac- are returning to college after a hiatus or attending college for count has always been an integral part of her teaching style. Dr. the first time and who would be unable to continue their studies Marieb began her teaching career at Springfield College, where without financial support. She funds the E. N. Marieb Science she taught anatomy and physiology to physical education ma- Research Awards at Mount Holyoke College, which promotes jors. She then joined the faculty of the Biological Science Divi- research by undergraduate science majors, and has underwrit- sion of Holyoke Community College in 1969 after receiving ten renovation and updating of one of the biology labs in Clapp her Ph.D. in zoology from the University of Massachusetts Laboratory at that college. Dr. Marieb also contributes to the at Amherst. While teaching at Holyoke Community College, University of Massachusetts at Amherst where she generously where many of her students were pursuing nursing degrees, provided funding for reconstruction and instrumentation of she developed a desire to better understand the relationship a cutting-edge cytology research laboratory. Recognizing the between the scientific study of the human body and the clinical severe national shortage of nursing faculty, she underwrites the aspects of the nursing practice. To that end, while continuing Nursing Scholars of the Future Grant Program at the university. to teach full time, Dr. Marieb pursued her nursing education, In 1994, Dr. Marieb received the Benefactor Award from which culminated in a Master of Science degree with a clinical the National Council for Resource Development, American specialization in gerontology from the University of Massachu- Association of Community Colleges, which recognizes her setts. It is this experience that has informed the development of ongoing sponsorship of student scholarships, faculty teaching the unique perspective and accessibility for which her publica- awards, and other academic contributions to Holyoke Com- tions are known. munity College. In May 2000, the science building at Holyoke Dr. Marieb has partnered with Benjamin Cummings for Community College was named in her honor. over 30 years. Her first work was Human Anatomy & Physiol- Dr. Marieb is an active member of the Human Anatomy ogy Laboratory Manual (Cat Version), which came out in 1981. and Physiology Society (HAPS) and the American Association In the years since, several other lab manual versions and study for the Advancement of Science (AAAS). Additionally, while guides, as well as the softcover Essentials of Human Anatomy actively engaged as an author, Dr. Marieb serves as a consultant & Physiology textbook, have hit the campus bookstores. This for the Benjamin Cummings Interactive Physiology® CD-ROM textbook, now in its 9th edition, made its appearance in 1989 series. and is the latest expression of her commitment to the needs of When not involved in academic pursuits, Dr. Marieb is students studying human anatomy and physiology. a world traveler and has vowed to visit every country on this Dr. Marieb has given generously to provide opportunities planet. Shorter term, she serves on the scholarship committee for students to further their education. She contributes to the of the Women’s Resources Center and on the board of directors New Directions, New Careers Program at Holyoke Commu- of several charitable institutions in Sarasota County. She is an nity College by funding a staffed drop-in center and by provid- enthusiastic supporter of the local arts and enjoys a competitive ing several full-tuition scholarships each year for women who match of doubles tennis. iii # 105016 Cust: Benjamin Cummings/CA Au: Marieb Pg. No. iii C/M/Y/K cDEaSIGrN SlERiVIsCESl OeF Title: Anatomy & Physiology Server: S4C Short / Normal Publishing Services iv About the Authors Katja Hoehn Dr. Katja Hoehn is an associate professor in the Department second-year medical students at Dalhousie University and at of Chemical and Biological Sciences at Mount Royal Univer- the University of Calgary. sity in Calgary, Canada. Dr. Hoehn’s first love is teaching. Her Dr. Hoehn has been a contributor to several books and has teaching excellence has been recognized by several awards dur- written numerous research papers in Neuroscience and Phar- ing her 17 years at Mount Royal University. These include a macology. She oversaw a recent revision of the Benjamin Cum- PanCanadian Educational Technology Faculty Award (1999), mings Interactive Physiology® CD-ROM series modules, and a Teaching Excellence Award from the Students’ Association coauthored the newest module, The Immune System. of Mount Royal (2001), and the Mount Royal Distinguished Following Dr. Marieb’s example, Dr. Hoehn provides fi- Faculty Teaching Award (2004). nancial support for students in the form of a scholarship that Dr. Hoehn received her M.D. (with Distinction) from she established in 2006 for nursing students at Mount Royal the University of Saskatchewan, and her Ph.D. in Pharma- University. cology from Dalhousie University. In 1991, the Dalhousie Dr. Hoehn is also actively involved in the Human Anat- Medical Research Foundation presented her with the Max omy and Physiology Society (HAPS) and is a member of the Forman (Jr.) Prize for excellence in medical research. Dur- American Association of Anatomists. When not teaching, she ing her Ph.D. and postdoctoral studies, she also pursued her likes to spend time outdoors with her husband and two sons, passion for teaching by presenting guest lectures to first- and compete in triathlons, and play Irish flute. # 105016 Cust: Benjamin Cummings/CA Au: Marieb Pg. No. iv C/M/Y/K cDEaSIGrN SlERiVIsCESl OeF Title: Anatomy & Physiology Server: S4C Short / Normal Publishing Services Introduce yourself to the chapter Improved readability and navigability makes the text more accessible and easier to study. Chapter 14 The Autonomic Nervous System 527 ■ Dilates the bronchioles in the lungs, increasing ventilation Parasympathetic Sympathetic (and thus increasing oxygen delivery to body cells) Eye Eye ■ Causes the liver to release more glucose into the blood to ac- Brain stem commodate th1e increased energy4 needs of body cells Salivary Skin* At the same time, the sympathetic division temporarily glands Cranial damps nonessential activities, such as gastrointestinal tract mo- Sympathetic Sglaalnivdasry twilaitity!. IItf iys ofuar amreo rrue nimnipnogr tfaronmt t oa gmivueg ygoeur,r dmiguessctilnesg elvuenrcyhth cinang Heart Cervical ganglia they need to get you out of danger. In such active situations, the sympathetic division generates a head of steam that enables Lungs Lungs the body to cope with situations that threaten homeostasis. It T1 Heart provides the optimal conditions for an appropriate response to some threat, whether that response is to run, see distant objects Stomach better, or think more clearly. Thoracic Stomach Pancreas We have just looked at two extreme situations in which one or the other branch of the ANS dominates. Think of the para- Liver sympathetic division as the D division [digestion, defecation, Pancreas and gall- Chapter Outlines and diuresis (urination)], and the sympathetic division as the L1 bladder E division (exercise, excitement, emergency, embarrassment). Liver and Adrenal Chapter outlines provide Table 14.4 (p. 536) presents a more detailed summary of how gall- Lumbar gland each division affects various organs. bladder a preview of the chapter Remember, however, that the two ANS divisions rarely work in an all-or-none fashion as described above. A dynamic antag- and help you locate onism exists between the divisions, and both make continuous Bladder Bladder fine adjustments to maiTntainh homeeosta sis.Autonomic Nervous System information easily. Genitals Sacral Genitals Check Your Understanding 1. Name the three types of effectors of the autonomic nervous Figure 14.3 The subdivisions of the ANS. The parasympathetic 14 system. and sympathetic divisions differ anatomically in the (1) sites where Learning Objectives 2. Wquhicickhly ,r ethlaey ss oinmstartuicc tnioenrvso furos msy sthteem C oNrS t htoe mANusSc?le Esx mploairne why. tphhoeerseitrg bnayen rsgvCyleinhosa anopiprcsit egfie isbnrieta e1rtsse4,), . a(2nT)hd re e( 3lAa) utliotvocena ltoeionmngistch oNsf e otrfhv etohiure sgir a Spnyrgsetlgieaam n(ingdlioicnaitce ad5n 2d7 ■ D3il.a tWesh itchhe b brarnocnhc ohOfi otvhleeesr AviniNe Swt hw eo( pulupldn. 5pg2rse,4 d–ion5mc2r7ien)aastein igf yvoeun wtielartei on TP*aArltahsoyumghhpe asyt hmheputaictmheaticn in nbeorvadtiyon itso tehex sqkinu iiss mitaeppleyd tsoe thnSes ycimetrivpviacaethl ettioc changes in its internal Learning objectives ■ (Caanudslw yetiahsn vugteh sso e?in n l Wictvhrheeeri ac btsheoi nab rcgreah lCon eexocanhNymsj egoewp eyrmaonivnrou ogidlusrd eotes hl n piSgevr ylo eeussdfrutc oyentoh mm tsaeoeisnn Sbida(no optttemphdo ei.ya f t 5 thsyci2oocee5u lu abl–n snw5ld)od2 eoo 6rAdfe) u tothtonoe anaco -mic rfiebgeiorsnE. yheeefroenr, varleli rnsooeruvnermsc teoes t nuhetn ,pd eaernirp dhee vereyn rcg-aacrrghye apsnoB isgnrtgai naianng lgs ictlfieoeomnlnoicdn siygtmi soptnartush.Ege Atygiecll eth toou bgahla anllc be ocdomy spyesttienmg sd ceomnatrnidbs- are integrated into the comsmurofbdoaaterd t haen din ac rsehAaasNrkeS da D peipnveiesarirogeynd s n we(ipethpdi.sn 5 oa2f 6 fbe–ow5d2 f7ye) ecte lolfs you? utSea, ltivhaery stability of our internal environment dependSsk ilna*rgely on the autonomic nervous cohf awptheart acnodn tgeivnet iyso tuo ac opmreev i ew dtwAialaimtiAtyN!.pt IsItt fS nh iysoe o n fuAasear s amsmrneeeno tarritueaimt ln iamonecAi,tp nmNitovPSghSyiar temtf rDyiaAra eiopnsssvnym,ayti a smstmtiu htoopaocepn ahgmta miti tc(havyhp use(e peT g tg(t.yhigp iFac5ocoeop s(2 urrrt.C ,a7 drra 5roc– dnia2mvoi5sinn7wigl2suui–te9oeie5mssor)sscs3tant,bil3 inecs na)resgtraa ee) ell ml)A Dvut preinpavprcocieyshtnrti oha dmcrniixnaio lHngy -. ■Fmsa yinggasdllLlAutkaoiHo eenegcret cmd cealaeaaasltr vtode tn1(iesdjoAdu4re ny s.siNt3 n t omo(m SF fittol )ihehltg,umnhee tsu C tehsetvsierrpn eriaaers i tvgtnscs, ie yce1asansarisnl4 ge alttcgce.nhlo1 lemaeeirs)aslffds.s .eo ae. s Mrcaft yrtnmooe dtsraootSg m oo atypeotnrm nahfggrrrpseal oainuaarnsmt reChsykeu.emr e vatorSyiincpoysp idcaanmteltiismhffrp ateeahaltrt iloehac sntre u ggtciiapnaeSgcnnsplna a,ggslo nei wvlaidrriannatsvhr t gyfaiaoolctri hretaeh s b eso CmdNyo oSac,t hatin vadint iadeus ct. oaInnrd orimeascipc mo nnuesrsevc teloes and what you are expected tthhee yFtsh oynerm e tsephitdaee t thpooeaf t rgCiaceNs tyd Smyi voopirusiaig otoihnnuVe ,tg it lsi(eooccpnec fpa aerd.nart 5ialado t2nR ens9egsys–fle m a5ore .3pfh x 3Iaeegn)sata hdns(e upgotc.lif ich5 as 3 d,ta e3iacva)ntimsidvio etgnh essani,tt e duereanatsailc obrnilbesse, acarhdeajLW usnuusngetmg sibn sbmeglg oaicornoid zno epdudrir te iidnsose nuTtsaar,ieb l tealhdene de1 xA 4bp.No1lod.Sry a stthieoumnnp tosef r btahtloueo rAed,N atSno d w“ niintehce rLdteuhyane”sg easasnr oae-ra sd, escpreeaesdess ostro smloawchs sheecarrett iroantes,. to learn. the fibboedry p taot hcwopayes w.ithA NsiStu Pahtiyosniso ltohgaty th(prpea. t5e3n3 –h5o3m9)eostasis. It tomMicoalslty o sfi mthpilse fir npaer-Tat1usynminpga tohcectuicr ds iwviistihoonu.t our awarHeneaersts or attention. Can you tell when provides the optimal conNdeiutrioontrsa nfosrm aitnt earps parnodp rRieactee prteosprso n se to your arteries are constricting or your pupils are dilating? Probably not—but if you’ve Bulleted Narrative sbAd■oe inmfftWtSaeeeitrrteo , te imhohnsrr aie ocvtahaeft l i,ljn oyuw,krs hi ttgm heliotneoh o.r eskeyPr e mcatdlhre paaTIaanastthr yt tlr(etheympew .reEspaoptpf.cif acot5eeti n3xcohat3stnnesre– tsed o5i imoc s3fp f5tDe fiaot) rbrhs uraieetugsru yAnssa m ,ut (aisptporeo.eane n5 t sodhc3 rimi5ensat)tini accwi n ohdts iioacvcbhisrj eiaoocln—ntess PThemfuSvatineeonr cmrd ptabai aocoesrhnefay nssiytm, smsb toopupwtcaahktn htT t,hie hhnytooeio rcasaut e cid’c ivccwhiev ee i(bsc’Crikeeoeo rnanua w itnvs a leiriarnoelys eso o,a fa aw cancaandlrrdlee a yd toloh )futo hDvrse ief si uccwvrelleiar PsS’nabratelilio naoo amncdscnraoaedtcactiehv.sr riIa tnlwy d. aThese cdeo, nAastNr tahSc etci otnengrt maros alisuf atiotll n htohamdes iaec or tthhee yo tohreigr ibnraatne cinh tohf et hberDa AiinvNi s(iSco rndason mi(upimpn.a) 5 tae3ns5.d– Th5 s3a7icn)rka l osfp tinhea l pcaorrad-. d(aivuitsoio 5n bseeclfa; unsoe mit s5 pr gegoavnegrnlio) nimic pfilbieesr,s tshpirsi nmgo ftroorm sL uiovbeprdpiovsiisteio n of the peripheral nervous Tbuhlele nteadrr wathiveer ehvaesr b peoesns i ble saETe■ynaa mcdbdhStsThRli hpievy edod oaimlien1saitur ivh4 atporshii.eceysanv4atiistme ciosihc(( s npaepe ll oedn t(x.aa iunneidt5cffvhrggr 3 eifiicl etsu6ncpibhtis)mtaoirese sctenp r ,i bgv oos rdH(eaaa aefnpaixrnr ssovr.) ictC e gre]iomh5itseno,l thu 3iietge(ntoaoehp9esismintrnn oo o )por aDdin erosfi.rc han tlsg5m b tac aatdoa3hoeo,tsnon8 fiefri lv ed rc–sus Attiem .5.sh hsmI u sy3mieThedheotm9b oreosnbo)agtppner paare tio[a—ilpnn pdtlampehoacaintodglyriseh ccsae,i ct ettesissFoeegcyutysur m ia mm odcondonobnr.cigpmfvitna, glai itriodasdithornihrnieanoetyae fisni eott scAoei cmn ca fNfiai — sntebdhS in oteiott hvthhrnw)iseee.,-. fvcessivrpnLgbPnayaioialtstiasvdvaenmelcnteldosd.-eacer d l m Thrloraetuei hanfnracn edle soth tt h rmreaCaeedrrs Nom t yCh(atS FoieN n ni crngaSeae exLs—rulra uyovtgrrmasntoelahtiysbuL nnee a1sm1asg ryb al4llnsir,m . ayata4wh ispo)tnehth.sue eiaswThnmc ttwh etal ,mye ii oiet wnh f tpc aod hrflpn oiueitcodcgshnash eaettct n hgettsrogasieteo n lr tfliosnuhgoarecalne cictwlo riu tlcanosiir nlt iec hacidrsatxi e sntenotgi hp eonisabtAgoueeunhsnllaydnnra dbe dro eno d eicdgonxdtnfneeoaaf tsanmnr rlen ltlg-elnchsooeredce--ts.i toTh oufs e it csAo enNfftrSeo cilts,o orasrl.s oth cea glleend etrhael to make the text easier to Rperemgeamngblieorn, hico wfibeverDesre , atvrheeal tos htphomer tet wnanotad Al tNAhsSep dpeiocvstitssgi ooannf sgt lhriaoern eAilcyN fiwSb oerrks oOrgavnse. Vrevry isheowrt postganglionic axons issue from the terminal read and navigate. ionn aisanmr ea llelox-noisgrt-.sn boentew feaesnh (itpoh.n e5 a3ds9i vd)iessiocrnisb,e adn adb boovteh. Am daykne acmonict iannutoaugs- gBanlagdldiear and synapse with effector cells in their immediBaltaed daerera. fine adjustments to maintain homeostasis. Define autonomic nervous system and explain its relationship to the peripheral Gennietarlvsous system. Sacral Genitals Check Your Check Your Understanding Compare the somatic and autonomic nervous systems relative to effectors, efferent 1. Name the three types of effectors of the autonomic nervous Figupraet h1w4.a3y sT, haen sdu nbdeuivriositornasn osmf tihttee ArsN rSe.l eThaese pda.rasympathetic 14 UConncdeeptr cshteackn ding #Ti 1tl0e5:23 0A..1 nsWqW6ay ut shhoit ciimeCcckmhhluyy s,.rb& tet:r hlPaBaenhey nycsss hojiian ommosloftai r5ngttuiyh 2cCc e t4n u ioA me nrNm vsSSo ife unrwrosgv omses/ruyC: sltSdAht4e e pCm rCeA oNduro:S tMm htoaein r miAaetbNue sS icf?Pl e ygEso.x umNp lowoa.ir nee5r 2ew7 h y. ShatphhonoeCerdrst/ei tr CMg s/ bny oaN/ymenmY rosgp/vyprKleainmosata hnroapeeilrcst i egiafic in sbndidetaievrt ssecis,)o,i . oa(nc2nntD)sdr EP ar aSdue(Ib3sGillrfatiN)fs the tlSiiolvrnhE gRceae iVSa nIel stCerfaviEuonitcSlengo nOsstmecFh toisicfo a otnlfhl yset hioinre f git rtha hpnere ge( 1lgpia)a a sn(riitngaedlsisoyi cwnmaihtcpe eadarne tdh etic and sympathetic divisions. *Although sympathetic innervation to the skin is mapped to the cervical questions are tied to lying on the beach enjoying the sun and the sound of the region here, all nerves to the periphery carry postganglionic sympathetic waves? Which branch would predominate if you were on a fibers. the sections' Learning surfboard and a shark appeared within a few feet of you? For answers, see Appendix H. ■ Location of their ganglia. Most parasympathetic ganglia are Objectives and ask you located in the visceral effector organs. Sympathetic ganglia to stop, think, and check ANS Anatomy lie close to the spinal cord. Figure 14.3 illustrates these and other key differences, which your understanding For the parasympathetic and sympathetic divisions, describe are summarized in Table 14.1. the site of CNS origin, locations of ganglia, and general We begin our detailed exploration of the ANS with the ana- before moving on. fiber pathways. tomically simpler parasympathetic division. Anatomically, the sympathetic and parasympathetic divisions differ in Parasympathetic (Craniosacral) Division ■ Sites of origin. Parasympathetic fibers are craniosacral— The parasympathetic division is also called the craniosacral MasteringAthe&y origPinat®e in the brain (cranium) and sacral spinal cord. division because its preganglionic fibers spring from opposite Sympathetic fibers are thoracolumbar—they originate in the ends of the CNS—the brain stem and the sacral region of the thoracic and lumbar regions of the spinal cord. spinal cord (Figure 14.4). The preganglionic axons extend ■ Relative lengths of their fibers. The parasympathetic divi- from the CNS nearly all the way to the structures they inner- sion has long preganglionic and short postganglionic fibers. vate. There the axons synapse with postganglionic neurons lo- Reading Questions The sympathetic division has the opposite condition—the cated in terminal ganglia that lie close to or within the target keep you on track. parree gloanngg.lionic fibers are short and the postganglionic fibers ogargnagnlisa. Vanedry s syhnoarpts pe owstitgha negffleioctnoirc caexlolsn isn i stshueeir f rimomm tehdei ateter marienaa.l # 105016 Cust: Benjamin Cummings/CA Au: Marieb Pg. No. 527 C/M/Y/K cDEaSIGrN SlERiVIsCESl OeF Title: Anatomy & Physiology Server: S4C Short / Normal Publishing Services # 105016 Cust: Benjamin Cummings/CA Au: Marieb Pg. No. v C/M/Y/K cDEaSIGrN SlERiVIsCESl OeF Title: Anatomy & Physiology Server: S4C Short / Normal Publishing Services Follow complex processes step by step Focus Figures help you grasp tough topics in A&P by walking you through carefully developed step-by-step illustrations that use a big-picture layout and dramatic art to provide a context for understanding the process. MasteringA&P® Focus Figure Tutorials FOCUS Bulk Flow Across Capillary Walls All Focus Figures have related tutorials in MasteringA&P Overview Figure 19.17 Bulk fluid flow across capillary walls causes that your instructor can continuous mixing of fluid between the plasma and the Each Overview quicky assign and that will guide summarizes the key interstitial fluid compartments, and maintains the How do the pressures drive fluid flow across a capillary? you through the figures interstitial environment. idea of the figure. step by step. Net filtration occurs at the arteriolar end of a capillary. Big Picture The big picture Capillary Boundary Interstitial fluid (capillary wall) Fluid filters from capillaries at their arteriolar Orientation end and flows through the interstitial space. Most is reabsorbed at the venous end. Hydrostatic pressure in capillary HP = 35 mm Hg The big picture provides “pushes” fluid out of capillary. c you with a concrete Arteriole starting point for the Osmotic pressure in capillary process. “pulls” fluid into capillary. OPc = 26 mm Hg Fluid moves through the interstitial space. To determine the pressure driving the For all capillary beds, fluid out of the capillary at any given 20 L of fluid is filtered Hydrostatic pressure in point, we calculate the net filtration out per day—almost 7 HPif = 0 mm Hg interstitial fluid pressure (NFP)––the outward pressures Blue Text tviomluems teh!e total plasma “cappuislhlaersy”. fluid into (pHrePscs aunreds O (HPPif) manindu sO tPhe) . inSwo,ard if c This text acts as the OP = 1 mm Hg Osmotic pressure in if interstitial fluid “pulls” NFP = (HP + OP) – (HP + OP) instructor’s voice fluid out of capillary. = (35 +c 1) – (if0 + 26)if c and explains difficult = 10 mm Hg (net outward pressure) concepts. In some As a result, fluid moves from the capillary NFP= 10 mm Hg into the interstitial space. figures the text is broken into numbered Net filtration pressure (NFP) determines the direction of fluid movement. Two kinds of pressure drive fluid flow: steps to help you Net reabsorption occurs at the venous end of a capillary. Hydrostatic pressure (HP) Osmotic pressure (OP) more easily understand difficult processes. • Due to fluid pressing against a • Due to nondiffusible solutes that Capillary Boundary Interstitial fluid boundary cannot cross the boundary (capillary wall) • HP “pushes” fluid across the • OP “pulls” fluid across the Hydrostatic pressure in capillary boundary boundary • In blood vessels, is due to blood • In blood vessels, is due to “ppreussshuerse” h flaus iddr oouptp eodf cbaepcialluasrey. oTfh e HPc = 17 mm Hg pressure plasma proteins resistance encountered along the capillaries. Piston 17 L of fluid per day is reabsorbed Osmotic pressure in capillary OP = 26 mm Hg into the capillaries “pulls” fluid into capillary. c Solute at the venous end. molecules About 3 L per day (proteins) of fluid (and any Boundary Boundary leaked proteins) are removed by the Again, we calculate the NFP: lymphatic system HP = 0 mm Hg Hydrostatic pressure in (see Chapter 20). if interstitial fluid “pushes” “Pushes” “Pulls” fluid into capillary. NFP = (HP + OP) – (HP + OP) c if if c Lymphatic = (17 + 1) – (0 + 26) Venule capillary OPif = 1 mm Hg Oinstmerosttiitci aplr eflsusiudr e“ pinu lls” fluid = –8 mm Hg (net inward pressure) out of capillary. Notice that the NFP at the venous end is a negative number. This means that reabsorption, not filtration, is occurring and so fluid moves from the interstitial NFP= –8 mm Hg space into the capillary. # 105016 Cust: Benjamin Cummings/CA Au: Marieb Pg. No. vi C/M/Y/K cDEaSIGrN SlERiVIsCESl OeF Title: Anatomy & Physiology Server: S4C Short / Normal Publishing Services Follow complex processes step by step Focus Figures help you grasp tough topics in A&P by walking you through carefully developed step-by-step illustrations that use a big-picture layout and dramatic art to provide a context for understanding the process. MasteringA&P® Focus Figure Tutorials FOCUS Bulk Flow Across Capillary Walls All Focus Figures have related tutorials in MasteringA&P Overview Figure 19.17 Bulk fluid flow across capillary walls causes that your instructor can continuous mixing of fluid between the plasma and the Each Overview quicky assign and that will guide summarizes the key interstitial fluid compartments, and maintains the How do the pressures drive fluid flow across a capillary? you through the figures interstitial environment. idea of the figure. step by step. Net filtration occurs at the arteriolar end of a capillary. Big Picture The big picture Capillary Boundary Interstitial fluid (capillary wall) Fluid filters from capillaries at their arteriolar Orientation end and flows through the interstitial space. Most is reabsorbed at the venous end. Hydrostatic pressure in capillary HP = 35 mm Hg The big picture provides “pushes” fluid out of capillary. c you with a concrete Arteriole starting point for the Osmotic pressure in capillary process. “pulls” fluid into capillary. OPc = 26 mm Hg Fluid moves through the interstitial space. To determine the pressure driving the For all capillary beds, fluid out of the capillary at any given 20 L of fluid is filtered Hydrostatic pressure in point, we calculate the net filtration out per day—almost 7 HPif = 0 mm Hg interstitial fluid pressure (NFP)––the outward pressures Blue Text tviomluems teh!e total plasma “cappuislhlaersy”. fluid into (pHrePscs aunreds O (HPPif) manindu sO tPhe) . inSwo,ard if c This text acts as the OP = 1 mm Hg Osmotic pressure in if interstitial fluid “pulls” NFP = (HP + OP) – (HP + OP) instructor’s voice fluid out of capillary. = (35 +c 1) – (if0 + 26)if c and explains dif- = 10 mm Hg (net outward pressure) ficult concepts. In As a result, fluid moves from the capillary NFP= 10 mm Hg into the interstitial space. some figures the text is broken into Net filtration pressure (NFP) determines the direction of fluid movement. Two kinds of pressure drive fluid flow: numbered steps to Net reabsorption occurs at the venous end of a capillary. Hydrostatic pressure (HP) Osmotic pressure (OP) help you more easily understand difficult • Due to fluid pressing against a • Due to nondiffusible solutes that Capillary Boundary Interstitial fluid boundary cannot cross the boundary (capillary wall) processes. • HP “pushes” fluid across the • OP “pulls” fluid across the Hydrostatic pressure in capillary boundary boundary • In blood vessels, is due to blood • In blood vessels, is due to “ppreussshuerse” h flaus iddr oouptp eodf cbaepcialluasrey. oTfh e HPc = 17 mm Hg pressure plasma proteins resistance encountered along the capillaries. Piston 17 L of fluid per day is reabsorbed Osmotic pressure in capillary OP = 26 mm Hg into the capillaries “pulls” fluid into capillary. c Solute at the venous end. molecules About 3 L per day (proteins) of fluid (and any Boundary Boundary leaked proteins) are removed by the Again, we calculate the NFP: lymphatic system HP = 0 mm Hg Hydrostatic pressure in (see Chapter 20). if interstitial fluid “pushes” “Pushes” “Pulls” fluid into capillary. NFP = (HP + OP) – (HP + OP) c if if c Lymphatic = (17 + 1) – (0 + 26) Venule capillary OPif = 1 mm Hg Oinstmerosttiitci aplr eflsusiudr e“ pinu lls” fluid = –8 mm Hg (net inward pressure) out of capillary. Notice that the NFP at the venous end is a negative number. This means that reabsorption, not filtration, is occurring and so fluid moves from the interstitial NFP= –8 mm Hg space into the capillary. # 105016 Cust: Benjamin Cummings/CA Au: Marieb Pg. No. vii C/M/Y/K cDEaSIGrN SlERiVIsCESl OeF Title: Anatomy & Physiology Server: S4C Short / Normal Publishing Services 188 Unit 2 Covering, Support, and Movement of the Body IMBALANCE Calcium homeostasis of blood: 9–11 mg/100 ml BALANCE BALANCE Stimulus Falling blood IMBALANCE Ca2+ levels Thyroid gland Osteoclasts degrade bone Parathyroid matrix and release glands Parathyroid Ca2+ into blood. glands release parathyroid hormone (PTH). 284 UNIT 2 Covering, Support, and Movement of the Body 6 PTH Thin filament (actin) Myosin heads Thick filament (myosin) mitochondria and glycogen granules, both involved in produc- Figure 6.12 Parathyroid hormone (PTH) control of blood calcium levels. ing the energy used during contraction. stimulated to contract. As you will see, calcium provides the When blood levels of ionic calcium decline, PTH is released through an additional pathway mediated by the hypothalamus, (Figure 6.12). The increased PTH level stimulates osteoclasts which activates sympathetic nerves serving bones. However, the to resorb bone, releasing calcium into blood. Osteoclasts are no full scope of leptin’s bone-modifying activity in humans is still T Tubules At each A band–I band junction, the sarcolemma respecters of matrix age: When activated, they break down both being worked out. of the muscle cell protrudes deep into the cell interior, form- ing an elongated tube called the T tubule (T for “transverse”). old and new matrix. As blood concentrations of calcium rise, It is also evident that the brain, intestine, and skeleton have the stimulus for PTH release ends. The decline of PTH reverses ongoing conversations that help regulate the balance between its effects and causes blood Ca21 levels to fall. bone formation and destruction, with serotonin serving as a fusing tubelike caveolae (inpocketings of the sarcolemma), In humans, calcitonin appears to be a hormone in search of a hormonal go-between. Serotonin is better known as a neu- the lumen (cavity) of the T tubule is continuous with the function because its effects on calcium homeostasis are negligi- rotransmitter that regulates mood and sleep, but most of the Figure 9.4 Myosin heads forming cross bridges that gener- extracellular space. ble. When administered at pharmacological (abnormally high) body’s serotonin is made in the gut (intestine) and the blood- ate muscular contractile force. Part of a sarcomere is seen in a Along its length, each T tubule runs between the paired ter- doses, it does lower blood calcium levels temporarily. brain barrier (see Chapter 12) bars it from entering the brain. transmission electron micrograph (277,000 ). Study figures as you read thmineal citsteernsx oft the SR, forming triads, successive groupings These hormonal conPtrorls eacpt toa prreeser vfe oblorod ycaolciuum r fThue troule rofe gu t cseraotornein eis srtill poorly understood. What is of the three membranous structures (terminal cistern, T tubule, homeostasis, not the skeleton’s strength or well-being. In fact, known is that when we eat, serotonin is secreted and circulated Sarcoplasmic Reticulum Shown in blue in Figure 9.5, the sar- if blood calcium levels are low for an extended time, the bones via the blood to the bones where it interferes with osteoblast ac- 9 Secloepclats mpiiec creetisc uoluf ma r(StR p) riso avni deleab omraoter sem ovoisthu eanl dcooplnastmeinc t andne oxtf, ttheen T h tuabvuele s also encircle each sarcomere. become so demineralizedC tlhinati tchaely cdoevveelorpa lgaerg ea,n pdun cchaesde- osuttu-dies thivaitvye. R bedeuecnti oenx opf baonndee tudr ntohvreor uaftgehro eautitn.g may lock calcium Muscle contraction is ultimately controlled by nerve- streeptic-ublyu-ms t(seepe ptpe. x00t– 0t0h)a. Itt sh inetleprcso nynoeuct inbge ttutbeurle su snurdroeurnsdt and structure, looking holes. Thus, the bones serve as a storehouse from which in bone when new calcium is flooding into the bloodstream. initiated electrical impulses that travel along the sarcolemma. functions, and processes. ionic calcium is drawn as needed. This is a troubling finding for those taking Prozac and other Because T tubules are continuations of the sarcolemma, they surrounds your arm. antidepressant drugs that inhibit serotonin uptake, making it conduct impulses to the deepest regions of the muscle cell and ▶ ▶co3m-Dmu naincaatitnog mwiyth aearcth other at the H zone. Others called - Homeostatic Imbalance 6.1 mHoorem aveaiolabsltea ttoi bco n e cells. Such patients have lower bone den- sIimty abnadl saunffcere m ore fractures than people not taking these drugs. teSrtmuninninagl c3i-sDte rannsa t(o“emnyd a srat ciss” r)e fnodremre lda rigne ar, dprearmpaetnicdailclyu lar cross Minute changes from the homeostatic range for blood calcium chmaonrnee dlsy anta tmhiec ,A r ebaalinstdic– Is tbyalen dth jautn ucsteios nvsib arnadnt t, hseaytu arlawteady s occur can lead to severe neuromuscular problems ranging from hyper- RHeosmpoenosseta ttico ImMbeaclahnacnei cseacl tiSotnrse s s The second set of controls inc oploairrss t. oC hloeslpe lyyo aus svoiscuiaaltiezed kweiyt ha ntahteo mSRic aal rset rluacrtguer ensu.mbers of excitability (when blood Ca21 levels are too low) to nonrespon- raergeu ilnatteignrga tebdo nweit hrienm thoed teelxint gan, db one’s response to mechanical siveness and inability to function (with high blood Ca21 levels). satlreersts y (omu utosc tlhee p cuolnl)s eaqnude ngcreasv oitfy , keeps the bones strong where In addition, sustained high blood levels of Ca21, a condition sbtroedsyso sryss taerme asc ntiontg f.unctioning Part of a skeletal I band A band I band known as hypercalcemia (hi0per-kal-se9me-ah), can lead to un- opWtimolaffll’ys. lTawhe hseo lpdast hthoalot gai cbaol n e grows or remodels in response muscle fiber (cell) 318 Unit d2eCosveirrinag, bSulpepo rtd, aendp Moovesmietnst ofo thfe Bcodaylcium salts in the blood vessels, kidneys, tcoo tnhdeit idoenms aarned isn tpelgarcaetde do nw iitth. The first thing to understand is Z disc H zone Z disc 25. Define EPOC.and other soft organs, wh i2c. hWh emn a sauiycid eh vicatimm waps foeurnd ,t thhe ceoriorne rf wuas nuncablte itoo n. ✚ tthhaet tae xbto tnoe ’sc laarniafyt oamndy irlleuflmeicntast eth e common stresses it encoun- 26. Smooth muscle has some unique properties, such as low energy remove the drug vial clutched in his hand. Explain the reasons for M usage, and the ability to maintain contraction over long periods. Tie this. If the victim had been discovered three days later, would the tneorsr.m Faolr f uenxcatmiopnlien,g a. bone is loaded (stressed) whenever weight line these properties to the function of smooth muscle in the body. coroner have had the same difficulty? Explain. Other hormones are al s3.o M uisnclev-reolaxlinvg edrdugs airne ad mminiosterdedi tof ya piantiengt d ubrinog mnajeor density bears down on it or muscles pull on it. This loading is usually surgery. Which of the two chemicals described next would be a Myofibril Sarcolemma 1. JjoimiMn eFdit caCaQh lnr oudidctaeeai ccslCi atdhliliaaeeso ndaTnlin dhttcshhtai anidclt lkp Auheib inps oba gppnh ldsrioyc bseaineqtig iuotaeennn il ets o ftt s “mgupuuumcrehApn t ,oir oopbne” vld& taehesriyerreed s.tP,i m saoFe sh o®er or lee x i4.na ■■gMm mo uiCCorcsdhhhce aleespeemmkg lce iiiellccseule aalta allen l,xABl sm caw flibltueoiatenrstoiciidndoplsengsn t r tatoaeh gn lsieaaed nnxmr daicbenuo ,sbtsn ol ocaotlafrnec aqnk dcc us tew eihlAeolshsntC’yi o.och? dIyn nrtsroe e rdcpemeelsnapapsltoimonsnorg sswi ewont itftio hytmh e “C. uWs a skB2cher1llaee.et etc aellls lest.a ksnedo wbny obffon cee notne ro anned s itdene dasn tdo sbuebnjedc tths eit b toon tee.n Bsieonnd i(nstgr ectocmhipnrge)s soens tthhee weekly. After thfreoe mro nitthss o fe trffaineingc, dtusri nog wnhic hw hee liftiegd ht andp reotneine chragngeys s hbapae wlhaenn Cca21e b i(ndss eto eit? ”p hep wr.it e9s 40–941), in other (Figure 6.13). increasingly heavier weights, he noticed that his arm and chest “tropomyosin.” What should he have responded and what is the muscles were suabsntanitmially alarlge r.s Etxpulaidn tihee sstru cltuerapl antdi n appearesrulst o f tthoat cailcniumh ioinb biintdi ngo?steoblasts. It does so functional basis of these changes. Triad: NEW! Homeostatic Imbalance • T tubule ATC lTinhicea lC QlIuNeIsCtions can be • Terminal 9 Relaatsesdi gCnliendic atlo T eyromus by your Sarcolemma cisterns Fibromiynossitits (rfiubroc 5t fioberr; itois 5n i nMflamamsatitoen) rAlisno kgnoAwn& P. on chromosome 19. Because the number of repeats tends to of the SR (2) as fibromyalgia; a group of conditions involving chronic increase from generation to generation, subsequent generations in3flTa1m8hmeatioyn oUhf an emitul s2pcle C, isotsv tceorrinnenge,c nStiuvgpe pttioshsrut,e ea cnondv eM riynogvose amnuedn rt of the Body develop more severe symptoms. No effective treatment. taenn 2udd5 io.n nnDvso,ed lafivnene dev acErra#PypsiOsn utg1Cl ead.0se ong5fr end0eesa 1oribfn 6yte jgno di n etro sn. eSCfsysm uahspstsoooctmi:as tw eaBdre we nitotnhnh ssjppaeeeccmi ififi cc in CRuImCEtr Ame a2ctr.m oiWnenynhmtge fn ofso ra r/a sr CupeiusctlAi,ld eiced e v m, iccu otismmAclpe wru, eoasrs:s i efooMxnuc,en asdans,id vrt heeilelyee cvsobtarrteoi otncneh .r eTh Pdw ateges ns u.tdan onNandbsal ero dto . 188 C/M/Y/K cDEaSIGrN SlERiVIsCESl OeF tri 2g6g.e rS pmooinottsh, ams uwsecllle a hsa fsa tsiogmuee aunndi qfrueeq pureonpt earwtiaeks,e snuinchg afrso lmow s leeneepr.gy or ligarmemenotvse. the drug vial clutched in his hand. Explain the reasons for Herniab Prooutsrdaugseiyo, an n wTodf tiahotne l oarebrgik:lai tnysA tt hontr mooauagi nthsto aititnsm a bcooyndyty r iac&ncatvi io tbnPy wohavaellryla. lMosnniagyoc p eleroio d gs.y T ie Sp as m AS sutehdirds.ev Infe, tihnrev: ov liSuctn4imtaCr hya tdw bitecehn i nd issmcoovoetrhe do rth srkeeel edtaayl sm lautsecrl,e w ould the Short / Normal Publishing Services Tubules of be contgheenseit parl o(poweritniegs ttoo ftahielu fruen octfi omnu osfc lsem fuoositohn m duusrcilneg i n the body. rangincgo rfroonmer mhaevree lhy aidrr tithaet isnagm toe dveiffiryc pualtiyn?f uElx; pmlaaiyn .be due to Mthyeo SfibRrils Myaldomgbeiuaoaves es(cinlmtoulyepi dda-mtani sl ed9oonwj erstd-u)CaQaf,eb nrhhbrus .i;udet beaaqti cslCumgatateiliaolin o nl sTth5nat im hcos aapniftunalesp ikccnAnli )enppie s Mwg pce.eau liauscnkcsaeleetnds ipi obn anygwin . h erheasvuyel tliinnftgin figrto om rf aanyll s ccmaha p alelr3semos.d alio ■■Msggct unoiaicnurgCClos ggseid,hhmec dp treelsh esbymmsk-ey.pae r cWiilaaleccahesfflaantaomhaellxcl iclABo;eci tnmghsu e fl.bgi ds ucoIo iu dnnasfaoa clr drt dlslufehlsepaysag e act r ts oomsteth owcam leaaacrr onxesysum da rcmoahnus hdfbe t asaemtll cypaohtml n iecenbenik diecnceigs aseydwi htllnAe eslthtlsv rCihd’oyode ecehr?dl y vos sa rctterpftoeor dafcipe a.base rlcSm eap peitdsatrax.mo leteAn ritmr esce wc cnhxuoiristtisfta n e hwdcmm.gl uoeC puarsuain, sn il2cdsd1gl e .bm cee aaljl os.r Myofas c1i.a lJ ipma iFni tscyhn ddercoimdeed P tahiant chaius spehdy bsiyq ua et ilgehftt emnuedch b taon db eo df esired, so he Strain C4o. mMmicohnaleyl cisa lalends wa e“rpinugll ead s meruiessc olef,” q au estsrtaioinn si sd eexacliensgsi vwei th skeletal muscljeo fiinbeedrs a, wlohciaclh h tewalitthch c lwuhb eann dth bee sgkainn toov e“pr uthmepm i riso nto”u tchhreeed .t imes stretchminugs calne dc eplol essxicbiltea ttieoanr ianngd o cf oan mtruacsctiloe nd.u Ien troe smpuosncslee to “What Mitochondria MyopMaothstyl ywi(n maecserisk-eooalycps.ii 9anAatgheftld-yet rwhh teeiht;a hrpve aioeetv hrme w5rouen sidgtehidhsset osao,rs f h este,tr r sanauiinoffntieenicrdgei n,dp dg otu)hs rAtauitnnr hgay li w dsm iahsuriemcsahcs l eaeh nsoe.df l icftheeds t oinvflearums“peetrrd ooo t(rpem aoinbmy uocysshoeiat.sin isThng),.e” eas W nisndhhj auaaprdtee js adwhc omehuneultnds j c oChlieean 2bth1sea cabvorieenm rdueessss upt poaoa lnilitynd? ”eifmu dhl mleay nw odbr iwitleihzsea dt .is the musclme.uscles were substantially larger. Explain the structural and Tetanus (1r)e Asu sltta otef tohfa stu csatalciinuemd cioonn tbriancdtiionng ?of a muscle that Myotonic dfuynstcrtoiopnhayl Aba fsoisr mof othf emseu scchualnagr edsy.strophy that is less is a normal aspect of skeletal muscle functioning. (2) An common than DMD; in the U.S. it affects about 14 of 100,000 acute infectious disease caused by the anaerobic bacterium people. Symptoms include a gradual reduction in muscle mass Clostridium tetani and resulting in persistent painful spasms of and cAontTrol ofT thhe skeelet aCl mulscIleNs, abInCormal heart rhythm, some skeletal muscles. Progresses to fixed rigidity of the jaws and diabetes mellitus. May appear at any time; not sex-linked. (lockjaw) and spasms of trunk and limb muscles. Usually fatal Figure 9.5 Relationship of the net of communicating channels at the level between the terminal cisterns. (See detailed Underlying genetic defect is multiple repeats of a particular gene due to respiratory failure. 9 Related Clinical Terms sarcoplasmic reticulum and T tubules to of the H zone and saclike elements called view in Figure 9.11, pp.290-291) Sites of close Fibromyositis (fibro 5 fibCer; aitiss 5e i nSflatmumdatiyon) AlMso kunsocwun lar System on chromosome 19. Because the number of repeats tends to MasteringA&P® myofibrils of skeletal muscle. The tubules terminal cisterns abutting the A-I junctions. contact of these three elements (terminal as fibromyalgia; a group of conditions involving chronic increase from generation to generation, subsequent generations inflammation of a muscle, its connective tissue coverings and develop more severe symptoms. No effective treatment. o“fh othleey S”R s l(ebeluvee.) Tehnecisrec lteu ebauclehs m fuysoefi tbor ilf olirkme aa Tohf eth Te tsuabrcuoleles m(gmraay )t haaret rinuwn adrede ipn vinatgoin tahteio cnesl l cciasltleerdn t,r Tia dtusb.ule, and terminal cistern) are Herntatbierneaing d cPdg oiorennornv tgspore,o ulanvisneniit dotavsn lac, r (aaooypsfi ws wnau LDtdmignenhlee ele duigltosts S’ea assr ottgtcs gioefcil rumiaf peonnaflnaeahtngee isitatag t l uhulooriunonbrrfrbeoeo uoeyt s.’aek ueseojnnio grnfmoddiv hngmua ee ft irtrrsadutni e.sott siq eSac bcntsuyallhosesemel eda nopofsp yutfnrfs t osa ocoohMiwbacomtevileanrraeissks tt m s.d eyea mkdnru swee ,irawl n iaendngiltotelg ah.fn lrM ssoppmaeeycc s iifilfiecce p. 12.. 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Wfarnohdma et t lceraocccmantmha rspialeepclrsemaclosidtlcal iio asogcttnitainicnimglo s goei,nmu dfpts lh a sbtsiheyptna icelaoa ahm sffnsnomec uclocio;esa tfcgsute .lihids eccIue naansarl rt elsrrfelparyuiagvac c oehmsttocm?uta cr rlWsysuee grmo hhasf e nm ayatly pdhtau enbrefs eieunceg lyndpiehnec asttlvt ihsoioodsoerrli n v vodsaee rpeftr d earfReea.ss rOcSmdu eitM?larxt. le eA tm rc cchuirsisanecm.gle pasn, isd prepare you for your future career. MasteringA&P® Her phMysyicoiafans coiradle pras idna silyyn pdarsosmivee Praaning ec-aoufs-emd obtyio an t i(gRhOteMn)e dex bearncidse o f 4. ExpSlatirna itnh eC roemasmononinlgy cbaellheidn da “thpue ldleide tmaruys rcelec,o” ma smtreanind aist ieoxncse.ssive and electricmalu sstcimle ufilbateirosn, wfohri chhe trw riigtchht wlehge nan tdh ea s dkiient ohviegrh t hine m is touched. stretching and possible tearing of a m(Aunsswclee rds uine Atop pmenudscixl eH ) protein, carMboohsytldyr aastseosc, iaantedd vwitiatmh oinv eCr.used or strained postural muscles. overuse or abuse. The injured muscle becomes painfully M09_MAR#I3 120685_00196_S E _CCHu0s9t:.i nBdedn j2a8m4in Cummings/CA Au: Marieb Pg. No. 284 C/M/Y/K CDEASIGRN SLERIVISCESL OEF 4/20/11 9:16 AM Myopmauthscyl e(.mi-op9ah-the; path 5 disease, suffering) Any disease of Tetaninufls a(m1)e Ad (smtayteo osift issu),s taaninde add cjaocnetnrta cjotiionnts o afr ea umsuusaclllye timhamt obilized. 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Ufousfl u tshapleal ysja mfwasts ao lf 4/20/11 9:16 AM End-of-chapter sections now contain assess your knowledge of terms an At the Clinic feaCtuarsee, Swtuhdicyh hMeulspc uylaor uS y stem and structures as well as the order apply what you’ve L elte’s caonrtinnuee odur. t aBle yof Mlers.a rning re 1.l aDetsceribde the step-by-step process of wound healing that of steps and elements involved in DeStephano’s medical problems, will occur in her fleshy (muscle) wounds, and note the clinical terms and trheis taimde lionokgin gs aht thoe rnott esC maadse e Studcioensseq u ences of the specific restorative process that occurs. physiological processes. and answering quedmesutastciluiinolagt unorbess.e,rv aytioonsu o f wher islkle lebtael gin t 2o. Wv a hscaut lacor m(bploliocdat vioenssse iln) dhaemaliangge ciann t hbee raignhtitc ilpeagt?ed owing to prep■■■ aSDTtrhreaaevemn elsora eewgfc leteoai orct oenlrir m tao htbfyie o)t ,hnbo jesulo sousotcf di arat bhvt ioeecfsv msneuee utlrshtsv ceseuel er(ritsrvgh ionheefgt lt a kthrnchgeee eaer ringirgehhertvt l eelee gsge ra ra.nvnidnd gk k nmneeoeest of 3. Wfarnohdma et t lceraocntmrsiepcaclitlc iasottniimo onufs la tihtnieo m nsc uoisafct lihece nsrt errruigvceht?ut rlWeeg ha nmy dau rfseuc lnpecast siosorinvde er erResOduM?lt Her physician orders daily passive range-of-motion (ROM) exercise 4. Explain the reasoning behind the dietary recommendations. and electrical stimulation for her right leg and a diet high in (Answers in Appendix H) protein, carbohydrates, and vitamin C. M09_MAR#TI3i 1t2l06e85: _0A019n6_aS t Eo _mCCHuy0 s&9t:. i nPBdhedny s j3ia1om8loingy C u m m Sienrgvse/rC: SA4 C Au: Marieb Pg. No. 318 ShoCrt/ M/ N/Yo/rKmal cDEPaSuIbGlriNsh SilnEgR iVSIesCrvEicSle OseF 4/20/11 9:16 AM # 105016 Cust: Benjamin Cummings/CA Au: Marieb Pg. No. viii C/M/Y/K cDEaSIGrN SlERiVIsCESl OeF Title: Anatomy & Physiology Server: S4C Short / Normal Publishing Services 188 Unit 2 Covering, Support, and Movement of the Body IMBALANCE Calcium homeostasis of blood: 9–11 mg/100 ml BALANCE BALANCE Stimulus Falling blood IMBALANCE Ca2+ levels Thyroid gland Osteoclasts degrade bone Parathyroid matrix and release glands Parathyroid Ca2+ into blood. glands release parathyroid hormone (PTH). 284 UNIT 2 Covering, Support, and Movement of the Body 6 PTH Thin filament (actin) Myosin heads Thick filament (myosin) mitochondria and glycogen granules, both involved in produc- Figure 6.12 Parathyroid hormone (PTH) control of blood calcium levels. ing the energy used during contraction. stimulated to contract. As you will see, calcium provides the When blood levels of ionic calcium decline, PTH is released through an additional pathway mediated by the hypothalamus, (Figure 6.12). The increased PTH level stimulates osteoclasts which activates sympathetic nerves serving bones. However, the to resorb bone, releasing calcium into blood. Osteoclasts are no full scope of leptin’s bone-modifying activity in humans is still T Tubules At each A band–I band junction, the sarcolemma respecters of matrix age: When activated, they break down both being worked out. of the muscle cell protrudes deep into the cell interior, form- ing an elongated tube called the T tubule (T for “transverse”). old and new matrix. As blood concentrations of calcium rise, It is also evident that the brain, intestine, and skeleton have the stimulus for PTH release ends. The decline of PTH reverses ongoing conversations that help regulate the balance between its effects and causes blood Ca21 levels to fall. bone formation and destruction, with serotonin serving as a fusing tubelike caveolae (inpocketings of the sarcolemma), In humans, calcitonin appears to be a hormone in search of a hormonal go-between. Serotonin is better known as a neu- the lumen (cavity) of the T tubule is continuous with the function because its effects on calcium homeostasis are negligi- rotransmitter that regulates mood and sleep, but most of the Figure 9.4 Myosin heads forming cross bridges that gener- extracellular space. ble. When administered at pharmacological (abnormally high) body’s serotonin is made in the gut (intestine) and the blood- ate muscular contractile force. Part of a sarcomere is seen in a Along its length, each T tubule runs between the paired ter- doses, it does lower blood calcium levels temporarily. brain barrier (see Chapter 12) bars it from entering the brain. transmission electron micrograph (277,000 ). Study figures as you read thmineal citsteernsx oft the SR, forming triads, successive groupings These hormonal conPtrorls eacpt toa prreeser vfe oblorod ycaolciuum r fThue troule rofe gu t cseraotornein eis srtill poorly understood. What is of the three membranous structures (terminal cistern, T tubule, homeostasis, not the skeleton’s strength or well-being. In fact, known is that when we eat, serotonin is secreted and circulated Sarcoplasmic Reticulum Shown in blue in Figure 9.5, the sar- if blood calcium levels are low for an extended time, the bones via the blood to the bones where it interferes with osteoblast ac- 9 Secloepclats mpiiec creetisc uoluf ma r(StR p) riso avni deleab omraoter sem ovoisthu eanl dcooplnastmeinc t andne oxtf, ttheen T h tuabvuele s also encircle each sarcomere. become so demineralizedC tlhinati tchaely cdoevveelorpa lgaerg ea,n pdun cchaesde- osuttu-dies thivaitvye. R bedeuecnti oenx opf baonndee tudr ntohvreor uaftgehro eautitn.g may lock calcium Muscle contraction is ultimately controlled by nerve- streeptic-ublyu-ms t(seepe ptpe. x00t– 0t0h)a. Itt sh inetleprcso nynoeuct inbge ttutbeurle su snurdroeurnsdt and structure, looking holes. Thus, the bones serve as a storehouse from which in bone when new calcium is flooding into the bloodstream. initiated electrical impulses that travel along the sarcolemma. functions, and processes. ionic calcium is drawn as needed. This is a troubling finding for those taking Prozac and other Because T tubules are continuations of the sarcolemma, they surrounds your arm. antidepressant drugs that inhibit serotonin uptake, making it conduct impulses to the deepest regions of the muscle cell and ▶ ▶co3m-Dmu naincaatitnog mwiyth aearcth other at the H zone. Others called - Homeostatic Imbalance 6.1 mHoorem aveaiolabsltea ttoi bco n e cells. Such patients have lower bone den- sIimty abnadl saunffcere m ore fractures than people not taking these drugs. teSrtmuninninagl c3i-sDte rannsa t(o“emnyd a srat ciss” r)e fnodremre lda rigne ar, dprearmpaetnicdailclyu lar cross Minute changes from the homeostatic range for blood calcium chmaonrnee dlsy anta tmhiec ,A r ebaalinstdic– Is tbyalen dth jautn ucsteios nvsib arnadnt t, hseaytu arlawteady s occur can lead to severe neuromuscular problems ranging from hyper- RHeosmpoenosseta ttico ImMbeaclahnacnei cseacl tiSotnrse s s The second set of controls inc oploairrss t. oC hloeslpe lyyo aus svoiscuiaaltiezed kweiyt ha ntahteo mSRic aal rset rluacrtguer ensu.mbers of excitability (when blood Ca21 levels are too low) to nonrespon- raergeu ilnatteignrga tebdo nweit hrienm thoed teelxint gan, db one’s response to mechanical siveness and inability to function (with high blood Ca21 levels). satlreersts y (omu utosc tlhee p cuolnl)s eaqnude ngcreasv oitfy , keeps the bones strong where In addition, sustained high blood levels of Ca21, a condition sbtroedsyso sryss taerme asc ntiontg f.unctioning Part of a skeletal I band A band I band known as hypercalcemia (hi0per-kal-se9me-ah), can lead to un- opWtimolaffll’ys. lTawhe hseo lpdast hthoalot gai cbaol n e grows or remodels in response muscle fiber (cell) 318 Unit d2eCosveirrinag, bSulpepo rtd, aendp Moovesmietnst ofo thfe Bcodaylcium salts in the blood vessels, kidneys, tcoo tnhdeit idoenms aarned isn tpelgarcaetde do nw iitth. The first thing to understand is Z disc H zone Z disc 25. Define EPOC.and other soft organs, wh i2c. hWh emn a sauiycid eh vicatimm waps foeurnd ,t thhe ceoriorne rf wuas nuncablte itoo n. ✚ tthhaet tae xbto tnoe ’sc laarniafyt oamndy irlleuflmeicntast eth e common stresses it encoun- 26. Smooth muscle has some unique properties, such as low energy remove the drug vial clutched in his hand. Explain the reasons for M usage, and the ability to maintain contraction over long periods. Tie this. If the victim had been discovered three days later, would the tneorsr.m Faolr f uenxcatmiopnlien,g a. bone is loaded (stressed) whenever weight line these properties to the function of smooth muscle in the body. coroner have had the same difficulty? Explain. Other hormones are al s3.o M uisnclev-reolaxlinvg edrdugs airne ad mminiosterdedi tof ya piantiengt d ubrinog mnajeor density bears down on it or muscles pull on it. This loading is usually surgery. Which of the two chemicals described next would be a Myofibril Sarcolemma 1. JjoimiMn eFdit caCaQh lnr oudidctaeeai ccslCi atdhliliaaeeso ndaTnlin dhttcshhtai anidclt lkp Auheib inps oba gppnh ldsrioyc bseaineqtig iuotaeennn il ets o ftt s “mgupuuumcrehApn t ,oir oopbne” vld& taehesriyerree ds.tP,i m saoFe sh o®er or lee x i4.na ■■gMm mo uiCCorcsdhhhce aleespeemmkg lce iiiellccseule aalta allen l,xABl sm caw flibltueoiatenrstoiciidndoplsengsn t r tatoaeh gn lsieaaed nnxmr daicbenuo ,sbtsn ol ocaotlafrnec aqnk dcc us tew eihlAeolshsntC’yi o.och? dIyn nrtsroe e rdcpemeelsnapapsltoimonsnorg sswi ewont itftio hytmh e “C. uWs a skB2cher1llaee.et etc aellls lest.a ksnedo wbny obffon cee notne ro anned s itdene dasn tdo sbuebnjedc tths eit b toon tee.n Bsieonnd i(nstgr ectocmhipnrge)s soens tthhee weekly. After thfreoe mro nitthss o fe trffaineingc, dtusri nog wnhic hw hee liftiegd ht andp reotneine chragngeys s hbapae wlhaenn Cca21e b i(ndss eto eit? ”p hep wr.it e9s 40–941), in other (Figure 6.13). increasingly heavier weights, he noticed that his arm and chest “tropomyosin.” What should he have responded and what is the muscles were suabsntanitmially alarlge r.s Etxpulaidn tihee sstru cltuerapl antdi n appearesrulst o f tthoat cailcniumh ioinb biintdi ngo?steoblasts. It does so functional basis of these changes. Triad: NEW! Homeostatic Imbalance • T tubule ATC lTinhicea lC QlIuNeIsCtions can be • Terminal 9 Relaatsesdi gCnliendic atlo T eyromus by your Sarcolemma cisterns Fibromiynossitits (rfiubroc 5t fioberr; itois 5n i nMflamamsatitoen) rAlisno kgnoAwn& P. on chromosome 19. Because the number of repeats tends to of the SR (2) as fibromyalgia; a group of conditions involving chronic increase from generation to generation, subsequent generations in3flTa1m8hmeatioyn oUhf an emitul s2pcle C, isotsv tceorrinnenge,c nStiuvgpe pttioshsrut,e ea cnondv eM riynogvose amnuedn rt of the Body develop more severe symptoms. No effective treatment. taenn 2udd5 io.n nnDvso,ed lafivnene dev acErra#PypsiOsn utg1Cl ead.0se ong5fr end0eesa 1oribfn 6yte jgno di n etro sn. eSCfsysm uahspstsoooctmi:as tw eaBdre we nitotnhnh ssjppaeeeccmi ififi cc in CRuImCEtr Ame a2ctr.m oiWnenynhmtge fn ofso ra r/a sr CupeiusctlAi,ld eiced e v m, iccu otismmAclpe wru, eoasrs:s i efooMxnuc,en asdans,id vrt heeilelyee cvsobtarrteoi otncneh .r eTh Pdw ateges ns u.tdan onNandbsal ero dto . 188 C/M/Y/K cDEaSIGrN SlERiVIsCESl OeF tri 2g6g.e rS pmooinottsh, ams uwsecllle a hsa fsa tsiogmuee aunndi qfrueeq pureonpt earwtiaeks,e snuinchg afrso lmow s leeneepr.gy or ligarmemenotvse. the drug vial clutched in his hand. Explain the reasons for Herniab Prooutsrdaugseiyo, an n wTodf tiahotne l oarebrgik:lai tnysA tt hontr mooauagi nthsto aititnsm a bcooyndyty r iac&ncatvi io tbnPy wohavaellryla. lMosnniagyoc p eleroio d gs.y T ie Sp as m AS sutehdirds.ev Infe, tihnrev: ov liSuctn4imtaCr hya tdw bitecehn i nd issmcoovoetrhe do rth srkeeel edtaayl sm lautsecrl,e w ould the Short / Normal Publishing Services Tubules of be contgheenseit parl o(poweritniegs ttoo ftahielu fruen octfi omnu osfc lsem fuoositohn m duusrcilneg i n the body. rangincgo rfroonmer mhaevree lhy aidrr tithaet isnagm toe dveiffiryc pualtiyn?f uElx; pmlaaiyn .be due to Mthyeo SfibRrils Myaldomgbeiuaoaves es(cinlmtoulyepi dda-mtani sl ed9oonwj erstd-u)CaQaf,eb nrhhbrus .i;udet beaaqti cslCumgatateiliaolin o nl sTth5nat im hcos aapniftunalesp ikccnAnli )enppie s Mwg pce.eau liauscnkcsaeleetnds ipi obn anygwin . h erheasvuyel tliinnftgin figrto om rf aanyll s ccmaha p alelr3semos.d alio ■■Msggct unoiaicnurgCClos ggseid,hhmec dp treelsh esbymmsk-ey.pae r cWiilaaleccahesfflaantaomhaellxcl iclABo;eci tnmghsu e fl.bgi ds ucoIo iu dnnasfaoa clr drt dlslufehlsepaysag e act r ts oomsteth owcam leaaacrr onxesysum da rcmoahnus hdfbe t asaemtll cypaohtml n iecenbenik diecnceigs aseydwi htllnAe eslthtlsv rCihd’oyode ecehr?dl y vos sa rctterpftoeor dafcipe a.base rlcSm eap peitdsatrax.mo leteAn ritmr esce wc cnhxuoiristtisfta n e hwdcmm.gl uoeC puarsuain, sn il2cdsd1gl e .bm cee aaljl os.r Myofas c1i.a lJ ipma iFni tscyhn ddercoimdeed P tahiant chaius spehdy bsiyq ua et ilgehftt emnuedch b taon db eo df esired, so he Strain C4o. mMmicohnaleyl cisa lalends wa e“rpinugll ead s meruiessc olef,” q au estsrtaioinn si sd eexacliensgsi vwei th skeletal muscljeo fiinbeedrs a, wlohciaclh h tewalitthch c lwuhb eann dth bee sgkainn toov e“pr uthmepm i riso nto”u tchhreeed .t imes stretchminugs calne dc eplol essxicbiltea ttieoanr ianngd o cf oan mtruacsctiloe nd.u Ien troe smpuosncslee to “What Mitochondria MyopMaothstyl ywi(n maecserisk-eooalycps.ii 9anAatgheftld-yet rwhh teeiht;a hrpve aioeetv hrme w5rouen sidgtehidhsset osao,rs f h este,tr r sanauiinoffntieenicrdgei n,dp dg otu)hs rAtauitnnr hgay li w dsm iahsuriemcsahcs l eaeh nsoe.df l icftheeds t oinvflearums“peetrrd ooo t(rpem aoinbmy uocysshoeiat.sin isThng),.e” eas W nisndhhj auaaprdtee js adwhc omehuneultnds j c oChlieean 2bth1sea cabvorieenm rdueessss upt poaoa lnilitynd? ”eifmu dhl mleay nw odbr iwitleihzsea dt .is the musclme.uscles were substantially larger. Explain the structural and Tetanus (1r)e Asu sltta otef tohfa stu csatalciinuemd cioonn tbriancdtiionng ?of a muscle that Myotonic dfuynstcrtoiopnhayl Aba fsoisr mof othf emseu scchualnagr edsy.strophy that is less is a normal aspect of skeletal muscle functioning. (2) An common than DMD; in the U.S. it affects about 14 of 100,000 acute infectious disease caused by the anaerobic bacterium people. Symptoms include a gradual reduction in muscle mass Clostridium tetani and resulting in persistent painful spasms of and cAontTrol ofT thhe skeelet aCl mulscIleNs, abInCormal heart rhythm, some skeletal muscles. Progresses to fixed rigidity of the jaws and diabetes mellitus. May appear at any time; not sex-linked. (lockjaw) and spasms of trunk and limb muscles. Usually fatal Figure 9.5 Relationship of the net of communicating channels at the level between the terminal cisterns. (See detailed Underlying genetic defect is multiple repeats of a particular gene due to respiratory failure. 9 Related Clinical Terms sarcoplasmic reticulum and T tubules to of the H zone and saclike elements called view in Figure 9.11, pp.290-291) Sites of close Fibromyositis (fibro 5 fibCer; aitiss 5e i nSflatmumdatiyon) AlMso kunsocwun lar System on chromosome 19. Because the number of repeats tends to MasteringA&P® myofibrils of skeletal muscle. The tubules terminal cisterns abutting the A-I junctions. contact of these three elements (terminal as fibromyalgia; a group of conditions involving chronic increase from generation to generation, subsequent generations inflammation of a muscle, its connective tissue coverings and develop more severe symptoms. No effective treatment. o“fh othleey S”R s l(ebeluvee.) Tehnecisrec lteu ebauclehs m fuysoefi tbor ilf olirkme aa Tohf eth Te tsuabrcuoleles m(gmraay )t haaret rinuwn adrede ipn vinatgoin tahteio cnesl l cciasltleerdn t,r Tia dtusb.ule, and terminal cistern) are Herntatbierneaing d cPdg oiorennornv tgspore,o ulanvisneniit dotavsn lac, r (aaooypsfi ws wnau LDtdmignenhlee ele duigltosts S’ea assr ottgtcs gioefcil rumiaf peonnaflnaeahtngee isitatag t l uhulooriunonbrrfrbeoeo uoeyt s.’aek ueseojnnio grnfmoddiv hngmua ee ft irtrrsadutni e.sott siq eSac bcntsuyallhosesemel eda nopofsp yutfnrfs t osa ocoohMiwbacomtevileanrraeissks tt m s.d eyea mkdnru swee ,irawl n iaendngiltotelg ah.fn lrM ssoppmaeeycc s iifilfiecce p. 12.. DwcWvoaeihslnslcRS acsoueprtIci lqCabaccsuoreuE mtore m(rr atbrA nehn ipAlnlaccoeigtleg r i oimhscossanudate mngeoedt ryn vip ffedomfl etrn-te noesbfhtn osmssfsyeeo ,rh.- i l irnsysa)mn t p r dev(hpeeemopauesrcltm ealuiu,plfil lesyiniarccndco tgi elga rrceme,err e)c ciys isutowant astosmn twoctori hlnpaubifeteg trn,ecwi e vohd trsaeooir ssgin i,unevpo htaxn eirntsccnord m,iley pdce ash eoag pnsnesoit?advseoati ehndltetilehn lf yoeou ag tvrswlht t;a st remotihkeni coeatgacclntye hu t.t eobraThsdle . mtdee uunsetsda ctonloend sa rd NincErWea!s eC yaoseu rS pturodby leCmo-ascohlivning gA scktiillvsi taiensd ■■■ SDTthreaaevmn Melsoraeeywgc aletealdomi gorctbe ioenulaveir m s estao(cilhmttoblfiyee op)ti ,dha-mnb ajniesulsloed 9ososnjo tercsft dd-ui )aata ,ebb hvthbrisoee.c;ue sv aqtmsne leumge utleirshaosvn c esset5elt em r(ro itsvpghft uioanheesifgtc nn llt ak )ethi rns hMwge eec eeeaur riausngikgcsehelehretndvt pl i eebnlea gygsigne .ha r aernvneaidnsdv ugyk kl ntlmniienfteoegiens ftgr oo omfr any 3. Wfarnohdma et t lceraocccmantmha rspialeepclrsemaclosidtlcal iio asogcttnitainicnimglo s goei,nmu dfpts lh a sbtsiheyptna icelaoa ahm sffnsnomec uclocio;esa tfcgsute .lihids eccIue naansarl rt elsrrfelparyuiagvac c oehmsttocm?uta cr rlWsysuee grmo hhasf e nm ayatly pdhtau enbrefs eieunceg lyndpiehnec asttlvt ihsoioodsoerrli n v vodsaee rpeftr d earfReea.ss rOcSmdu eitM?larxt. le eA tm rc cchuirsisanecm.gle pasn, isd prepare you for your future career. MasteringA&P® Her phMysyicoiafans coiradle pras idna silyyn pdarsosmivee Praaning ec-aoufs-emd obtyio an t i(gRhOteMn)e dex bearncidse o f 4. ExpSlatirna itnh eC roemasmononinlgy cbaellheidn da “thpue ldleide tmaruys rcelec,o” ma smtreanind aist ieoxncse.ssive and electricmalu sstcimle ufilbateirosn, wfohri chhe trw riigtchht wlehge nan tdh ea s dkiient ohviegrh t hine m is touched. stretching and possible tearing of a m(Aunsswclee rds uine Atop pmenudscixl eH ) protein, carMboohsytldyr aastseosc, iaantedd vwitiatmh oinv eCr.used or strained postural muscles. overuse or abuse. The injured muscle becomes painfully M09_MAR#I3 120685_00196_S E _CCHu0s9t:.i nBdedn j2a8m4in Cummings/CA Au: Marieb Pg. No. 284 C/M/Y/K CDEASIGRN SLERIVISCESL OEF 4/20/11 9:16 AM Myopmauthscyl e(.mi-op9ah-the; path 5 disease, suffering) Any disease of Tetaninufls a(m1)e Ad (smtayteo osift issu),s taaninde add cjaocnetnrta cjotiionnts o afr ea umsuusaclllye timhamt obilized. Title: Anatomy & NPhEyWsio!l oAgyr t L a bSeelrivnegr :a Sn4dC Ranking/ Short / Normal Publishing Services Myotcoonmicm doyns ttrhoapnh Dy MA Dfo;r imn tohf em Uu.Ssc. uitl aarff decytsst raobpohuyt t1h4a to fis 1 l0e0ss,0 00 iasc au tneo irnmfeaclt iaosupse cdti soefa sskee cleatuasle md ubsyc tlhe efu anncateiroonbinicg b. 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Ufousfl u tshapleal ysja mfwasts ao lf 4/20/11 9:16 AM End-of-chapter sections now contain assess your knowledge of terms an At the Clinic feaCtuarsee, Swtuhdicyh hMeulspc uylaor uS y stem and structures as well as the order apply what you’ve L elte’s caonrtinnuee odur. t aBle yof Mlers.a rning re 1.l aDetsceribde the step-by-step process of wound healing that of steps and elements involved in DeStephano’s medical problems, will occur in her fleshy (muscle) wounds, and note the clinical terms and trheis taimde lionokgin gs aht thoe rnott esC maadse e Studcioensseq u ences of the specific restorative process that occurs. physiological processes. and answering quedmesutastciluiinolagt unorbess.e,rv aytioonsu o f wher islkle lebtael gin t 2o. 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No. 318 ShoCrt/ M/ N/Yo/rKmal cDEPaSuIbGlriNsh SilnEgR iVSIesCrvEicSle OseF 4/20/11 9:16 AM # 105016 Cust: Benjamin Cummings/CA Au: Marieb Pg. No. ix C/M/Y/K cDEaSIGrN SlERiVIsCESl OeF Title: Anatomy & Physiology Server: S4C Short / Normal Publishing Services