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Guyton & Hall Physiology ReviewSecond EditionJohn E. Hall, PhDArthur C. Guyton,Professor and Chair , Associate Vice Chancellor for Research, Department of Physiology and Biophysics, University of Mississippi Medical Center, Jackson,MississippiSaunders

Copyright1600 John F. Kennedy Blvd., Ste. 1800Philadelphia, PA 19103–2899GUYTON & HALL PHYSIOLOGY REVIEW, SECOND EDITION ISBN: 978-1-4160-5452-8Copyright 2011, 2006 by Saunders, an imprint of Elsevier Inc. All rights reserved.No part of this publication may be reproduced or transmitted in any form or by any means, electronicor mechanical, including photocopying, recording, or any information storage and retrieval system,without permission in writing from the publisher. Details on how to seek permission, further informationabout the Publisher’s permissions policies and our arrangements with organizations such as the CopyrightClearance Center and the Copyright Licensing Agency, can be found at our website:www.elsevier.com/permissions.This book and the individual contributions contained in it are protected under copyright by thePublisher (other than as may be noted herein).NoticesKnowledge and best practice in this field are constantly changing. As new research and experiencebroaden our understanding, changes in research methods, professional practices, or medical treatmentmay become necessary.Practitioners and researchers must always rely on their own experience and knowledge in evaluatingand using any information, methods, compounds, or experiments described herein. In using suchinformation or methods they should be mindful of their own safety and the safety of others, includingparties for whom they have a professional responsibility.With respect to any drug or pharmaceutical products identified, readers are advised to check the mostcurrent information provided (i) on procedures featured or (ii) by the manufacturer of each product to beadministered, to verify the recommended dose or formula, the method and duration of administration, andcontraindications. It is the responsibility of practitioners, relying on their own experience and knowledgeof their patients, to make diagnoses, to determine dosages and the best treatment for each individualpatient, and to take all appropriate safety precautions.To the fullest extent of the law, neither the Publisher nor the authors, contributors, or editors, assumeany liability for any injury and/or damage to persons or property as a matter of products liability,negligence or otherwise, or from any use or operation of any methods, products, instructions, or ideascontained in the material herein.ISBN: 978-1-4160-5452-8Acquisitions Editor: William R. SchmittDevelopmental Editors: Christine AbshirePublishing Services Manager: Patricia TannianSenior Project Manager: Sarah WunderlyDesign Direction: Louis ForgionePrinted in ChinaLast digit is the print number: 9 8 7 6 5 4 3 2 1

ContributorsThomas H. Adair, PhD, Professor of Physiology and Biophysics, University of Mississippi Medical Center, Jackson,Mississippi, Unit II, Unit XII, and Unit XIIIDavid J. Dzielak, PhD, Professor of Surgery, Professor of Health Sciences, Associate Professor of Physiology and Biophysics,University of Mississippi Medical Center, Jackson, Mississippi, Unit IX, Unit X, andUnit XIJoey P. Granger, PhD, Billy Guyton Professor of Physiology and Biophysics and Medicine, Dean of the School of GraduateStudies, University of Mississippi Medical Center, Jackson, Mississippi, Unit IVJohn E. Hall, PhD, Arthur C. Guyton Professor and Chair, Associate Vice Chancellor for Research, Department of Physiologyand Biophysics, University of Mississippi Medical Center, Jackson, Mississippi, Unit I, Unit V, andUnit XIIIRobert L. Hester, PhD, Professor of Physiology and Biophysics, University of Mississippi Medical Center, Jackson,Mississippi, Unit VI, Unit VII, andUnit VIIIThomas E. Lohmeier, PhD, Professor of Physiology and Biophysics, University of Mississippi Medical Center, Jackson,Mississippi, Unit XIVR. Davis Manning, PhD, Professor of Physiology and Biophysics, University of Mississippi Medical Center, Jackson,Mississippi, Unit III, Unit IV, andUnit XVDavid B. Young, PhD, Professor Emeritus of Physiology and Biophysics, University of Mississippi Medical Center, Jackson,Mississippi, Unit XIV

PrefaceSelf-assessment is an important component of effective learning, especially when studying a subjectas complex as medical physiology. The Guyton & Hall Physiology Review is designed to provide acomprehensive review of medical physiology through multiple-choice questions and explanations of theanswers. Medical students preparing for the United States Medical Licensure Examinations (USMLE)will also find this book useful, since test questions have been constructed according to the USMLEformat.The questions and answers in this review are based on Guyton and Hall’s Textbook of MedicalPhysiology, twelfth edition (TMP 12). More than 1000 questions and answers are provided, and eachanswer is referenced to the Textbook of Medical Physiology to facilitate a more complete understandingof the topic and self-assessment of your knowledge. Illustrations are used to reinforce basic concepts.Some of the questions incorporate information from multiple chapters in the Textbook of MedicalPhysiology to test your ability to apply and integrate the principles necessary for the mastery of medicalphysiology.An effective way to use the review is to allow an average of 1 minute for each question in a givenunit, approximating the time limit for a question in the USMLE examination. As you proceed, indicateyour answer next to each question. After finishing the questions and answers, spend as much time asnecessary to verify your answers and to carefully read the explanations provided. Read the additionalmaterial referred to in the Textbook of Medical Physiology , especially for questions where incorrectanswers were chosen.Guyton & Hall Physiology Review should not be used as a substitute for the comprehensiveinformation contained in the Textbook of Medical Physiology . It is intended mainly as a means ofassessing your knowledge of physiology and of strengthening your ability to apply and integrate thisknowledge.We have attempted to make this review as accurate as possible, and we hope that it will be avaluable tool for your study of physiology. We invite you to send us your critiques, suggestions forimprovement, and notifications of any errors.I am grateful to each of the contributors for their careful work on this book. I also wish to express mythanks to William Schmitt, Rebecca Gruliow, Christine Abshire, and the rest of the Elsevier staff for theireditorial and production excellence. I am especially indebted to the late Dr. Arthur C. Guyton, who wrotethe first eight editions of the Textbook of Medical Physiology , beginning nearly 50 years ago. I had theprivilege of working with him on the ninth and tenth editions and have attempted in the last two editions tocontinue his practice of accurately presenting the complex principles of physiology in language that iseasy for students to read and understand.John E. Hall, PhD, Jackson, Mississippi

Table of ContentsCopyrightContributorsPrefaceUnit 1: The cell and general physiologyUnit 2: Membrane physiology, nerve, and muscleUnit 3: The heartUnit 4: The circulationUnit 5: The body fluids and kidneysUnit 6: Blood cells, immunity, and blood coagulationUnit 7: RespirationUnit 8: Aviation, space, and deep-sea diving physiologyUnit 9: The nervous system: a. general principles and sensory physiologyUnit 10: The nervous system: b. the special sensesUnit 11: The nervous system: c. motor and integrative neurophysiologyUnit 12: Gastrointestinal physiologyUnit 13: Metabolism and temperature regulationUnit 14: Endocrinology and reproductionUnit 15: Sports physiology

UNIT IThe cell and general physiology1. The term “glycocalyx” refers toA) the negatively charged carbohydrate chains that protrude into the cytosol from glycolipids andintegral glycoproteinsB) the negatively charged carbohydrate layer on the outer cell surfaceC) the layer of anions aligned on the cytosolic surface of the plasma membraneD) the large glycogen stores found in “fast” musclesE) a mechanism of cell–cell attachment2. Messenger RNA (mRNA)A) carries the genetic code to the cytoplasmB) carries activated amino acids to the ribosomesC) is single-stranded RNA molecules of 21 to 23 nucleotides that can regulate gene transcriptionD) forms ribosomes3. Which of the following statements is true for both pinocytosis and phagocytosis?A) Involves the recruitment of actin filamentsB) Occurs spontaneously and non-selectivelyC) Endocytotic vesicles fuse with ribosomes that release hydrolases into the vesiclesD) Is only observed in macrophages and neutrophilsE) Does not require ATP4. In comparing two types of cells from the same person, the variation in the proteins expressed by eachcell type reflectsA) differences in the DNA contained in the nucleus of each cellB) differences in the numbers of specific genes in their genomesC) cell-specific expression and repression of specific genesD) differences in the number of chromosomes in each cellE) the age of the cells5. Micro RNAs (miRNAs)A) are formed in the cytoplasm and repress translation or promote degradation of mRNA before itcan be translatedB) are formed in the nucleus and then processed in the cytoplasma by the dicer enzymeC) are short (21 to 23 nucleotides) double-stranded RNA fragments that regulate gene expressionD) repress gene transcriptionQuestions 6–8A) Nucleolus

B) NucleusC) Agranular endoplasmic reticulumD) Granular endoplasmic reticulumE) Golgi apparatusF) EndosomesG) PeroxisomesH) LysosomesI) CytosolJ) CytoskeletonK) GlycocalyxL) MicrotubulesFor each of the scenarios described below, identify the most likely subcellular site listed above for thedeficient or mutant protein.6. Studies completed on a 5-year-old boy show an accumulation of cholesteryl esters and triglyceridesin his liver, spleen, and intestines and calcification of both adrenal glands. Additional studies indicate thecause to be a deficiency in acid lipase A activity.7. The abnormal cleavage of mannose residues during the post-translational processing of glycoproteinsresults in the development of a lupus-like autoimmune disease in mice. The abnormal cleavage is due to amutation of the enzyme α-mannosidase II.8. The observation that abnormal cleavage of mannose residues from glycoproteins causes anautoimmune disease in mice supports the role of this structure in the normal immune response.Questions 9–11A) NucleolusB) NucleusC) Agranular endoplasmic reticulumD) Granular endoplasmic reticulumE) Golgi apparatusF) Endosomes

G) PeroxisomesH) LysosomesI) CytosolJ) CytoskeletonK) GlycocalyxL) MicrotubulesMatch the cellular location for each of the steps involved in the synthesis and packaging of a secretedprotein listed below with a term listed above.9. Initiation of translation10. Protein condensation and packaging11. Gene transcription12. “Redundancy” or “degeneration” of the genetic code occurs during which of the following steps ofprotein synthesis?A) DNA replicationB) TranscriptionC) Post-transcriptional modificationD) TranslationE) Protein glycosylation13. Which of the following does not play a direct role in the process of transcription?A) HelicaseB) RNA polymeraseC) Chain-terminating sequenceD) “Activated” RNA moleculesE) Promoter sequence14. Which of the following proteins is most likely to be the product of a proto-oncogene?A) Growth factor receptorB) Cytoskeletal proteinC) Na channelD) Ca -ATPaseE) Myosin light chain15. Which of the following events does not occur during the process of mitosis?A) Condensation of the chromosomesB) Replication of the genomeC) Fragmentation of the nuclear envelopeD) Alignment of the chromatids along the equatorial plate

E) Separation of the chromatids into two sets of 46 “daughter” chromosomes16. Which of the following characteristics of a biological membrane is most influenced by itscholesterol content?A) ThicknessB) Ion permeabilityC) FluidityD) GlycosylationE) Hydrophobicity17. The appearance of which of the following distinguishes eukaryotic cells from lower units of life?A) DNAB) RNAC) MembranesD) ProteinE) Nucleus18. Assume that excess blood is transfused into a patient whose arterial baroreceptors are nonfunctionaland blood pressure increases from 100 to 150 mm Hg. Then, assume that the same volume is blood isinfused into the same patient under conditions where his arterial baroreceptors are functioning normallyand blood pressure increases from 100 to 125 mm Hg. What is the approximate feedback “gain” of thearterial baroreceptors in this patient when they are functioning normally?A) 1.0B) 2.0C) 0.0D) 1.0E) 2.0Answers1.B) The cell “glycocalyx” is the loose negatively charged carbohydrate coat on the outside of thesurface of the cell membrane. The membrane carbohydrates usually occur in combination with proteins orlipids in the form of glycoproteins or glycolipids, and the “glyco” portion of these molecules almostinvariably protrudes to the outside of the cell.TMP12 142.A) mRNA molecules are long, single RNA strands that are suspended in the cytoplasm, and arecomposed of several hundred to several thousand RNA nucleotides in unpaired strands. The mRNAcarries the genetic code to the cytoplasm for controlling the type of protein formed. The transfer RNA(tRNA) transports activated amino acids to the ribosomes. Ribosomal RNA, along with about 75 differentproteins, forms ribosomes. MicroRNAs are single-stranded RNA molecules of 21 to 23 nucleotides thatregulate gene transcription and translation.TMP12 313.A) Both pinocytosis and phagocytosis involve movement of the plasma membrane. Pinocytosisinvolves invagination of the cell membrane whereas phagocytosis involves evagination. Both eventsrequire the recruitment of actin and other cytoskeleton elements. Phagocytosis is not spontaneous and is

selective, being triggered by specific receptor-ligand interactions.TMP12 194.C) The variation in proteins expressed by each cell reflects cell-specific expression and repression ofspecific genes. Each cell contains the same DNA in the nucleus and the same number of genes. Sodifferentiation results not from differences in the genes but from selective repression and /or activation ofdifferent gene promoters.TMP12 39–405.A) MicroRNAs (miRNA) are formed in the cytoplasm from pre-miRNAs and processed by theenzyme dicer that ultimately assembles RNA-induced silencing complex (RISC), which then generatesmiRNAs. The miRNAs regulate gene expression by binding to the complementary region of the RNA andrepressing translation or promoting degradation of mRNA before it can be translated by the ribosome.TMP 12 32–336.H) Acid lipases, along with other acid hydrolases, are localized to lysosomes. Fusion of endocytoticand autolytic vesicles with lysosomes initiates the intracellular process that allows cells to digest cellulardebris and particles ingested from the extracellular milieu, including bacteria. In the normal acidicenvironment of the lysosome, acid lipases use hydrogen to convert lipids into fatty acids and glycerol.Other acid lipases include a variety of nucleases, proteases, and polysaccharide-hydrolyzing enzymes.TMP12 157.E) Membrane proteins are glycosylated during their synthesis in the lumen of the rough endoplasmicreticulum. Most post-translational modification of the oligosaccharide chains, however, occurs during thetransport of the protein through the layers of the Golgi apparatus matrix, where enzymes such as αmannosidase II are localized.TMP12 158.K) The oligosaccharide chains that are added to glycoproteins on the luminal side of the roughendoplasmic reticulum, and subsequently modified during their transport through the Golgi apparatus, areattached to the extracellular surface of the cell. This negatively charged layer of carbohydrate moieties iscollectively called the glycocalyx. It participates in cell–cell interactions, cell–ligand interactions, andthe immune response.TMP12 14; see also Chapter 349.I) Initiation of translation, whether of a cytosolic protein, a membrane-bound protein, or a secretedprotein, occurs in the cytosol and involves a common pool of ribosomes. Only after the appearance of theN-terminus of the polypeptide is it identified as a protein destined for secretion. At this point, theribosome attaches to the cytosolic surface of the rough endoplasmic reticulum. Translation continues, andthe new polypeptide is extruded into the matrix of the endoplasmic reticulum.TMP12 32–3310.E) Secreted proteins are condensed, sorted, and packaged into secretory vesicles in the terminalportions of the Golgi apparatus, also known as the trans-Golgi network. It is here that proteins destinedfor secretion are separated from those destined for intracellular compartments or cellular membranes.TMP12 15

11.B) All transcription events occur in the nucleus, regardless of the final destination of the proteinproduct. The resulting messenger RNA molecule is transported through the nuclear pores in the nuclearmembrane and translated into either the cytosol or the lumen of the rough endoplasmic reticulum.TMP12 3112.D) During both replication and transcription, the new nucleic acid molecule is an exact complementof the parent DNA molecule. This is a result of predictable, specific, one-to-one base pairing. During theprocess of translation, however, each amino acid in the new polypeptide is encoded by a codon, a seriesof three consecutive nucleotides. Whereas each codon encodes a specific amino acid, most amino acidscan be encoded for by multiple codons. Redundancy results because 60 codons encode a mere 20 aminoacids.TMP12 3113.A) Helicase is one of the many proteins involved in the process of DNA replication. It does not playa role in transcription. RNA polymerase binds to the promoter sequence and facilitates the addition of“activated” RNA molecules to the growing RNA molecule until the polymerase reaches the chainterminating sequence on the template DNA molecule.TMP12 30–3114.A) An oncogene is a gene that is either abnormally activated or mutated in such a way that itsproduct causes uncontrolled cell growth. A proto-oncogene is simply the “normal” version of anoncogene. By definition, proto-oncogenes are divided into several families of proteins, all of whichparticipate in the control of cell growth. These families include, but are not limited to, growth factors andtheir receptors, protein kinases, transcription factors, and proteins that regulate cell proliferation.TMP12 40–4115.B) DNA replication occurs during the S phase of the cell cycle and precedes mitosis. Condensationof the chromosomes occurs during the prophase of mitosis. Fragmentation of the nuclear envelope occursduring the prometaphase of mitosis. The chromatids align at the equatorial plate during metaphase andseparate into two complete sets of daughter chromosomes during anaphase.TMP12 3716.C) The cholesterol content of a membrane determines the packing density of phospholipids. Thehigher the cholesterol content, the more fluid the membrane and the greater the lateral mobility ofmembrane components, including proteins and phospholipid molecules themselves. To a lesser extent,cholesterol content also affects the “leakiness” of a membrane to water-soluble molecules.TMP12 1317.E) Nucleic acids and proteins, together, constitute the fundamental replicable unit of life,exemplified by viruses. Membranes and even organelles appear in prokaryotic cells, but only eukaryoticcells possess a nucleus.TMP12 17–1818.A) The feedback gain of the control system is calculated as the amount of correction divided by theremaining error of the system. In this example, blood pressure increased from 100 to 150 mm Hg when thebaroreceptors were not functioning. When the baroreceptors

Guyton & Hall Physiology Review should not be used as a substitute for the comprehensive information contained in the Textbook of Medical Physiology. It is intended mainly as a means of assessing your knowledge of physiology and of strengthening your ability to apply and integrate this knowledge.

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