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Ranges of Normal Values in Human Whole Blood (B), Plasma (P), or Serum (S)a Normal Value (Varies with Procedure Used) Determination Traditional Units SI Units Normal Value (Varies with Procedure Used) Determination Traditional Units SI Units Acetoacetate plus acetone (S) 0.3–2.0 mg/dL 3–20 mg/L Aldosterone (supine) (P) 3.0–10 ng/dL 83–227 pmol/L Alpha-amino nitrogen (P) 3.0–5.5 mg/dL 2.1–3.9 mmol/L Aminotransferases Alanine aminotransferase 3–48 units/L Aspartate aminotransferase 0–55 units/L Ammonia (B) 12–55 μmol/L 12–55 μmol/L Amylase (S) 53–123 units/L 884–2050 nmol s–1/L Ascorbic acid (B) 0.4–1.5 mg/dL (fasting) 23–85 μmol/L Bilirubin (S) Conjugated (direct): up to 0.4 mg/dL Up to 7 μmol/L Total (conjugated plus free): up to 1.0 mg/dL Up to 17 μmol/L Calcium (S) 8.5–10.5 mg/dL; 4.3–5.3 meq/L 2.1–2.6 mmol/L Carbon dioxide content (S) 24–30 meq/L 24–30 mmol/L Carotenoids (S) 0.8–4.0 μg/mL 1.5–7.4 μmol/L Ceruloplasmin (S) 23–43 mg/dL 240–430 mg/L Chloride (S) 100–108 meq/L 100–108 mmol/L Cholesterol (S) < 200 mg/dL < 5.17 mmol/L Cholesteryl esters (S) 60–70% of total cholesterol Copper (total) (S) 70–155 μg/dL 11.0–24.4 μmol/L Cortisol (P) (AM, fasting) 5–25 μg/dL 0.14–0.69 μmol/L Creatinine (P) 0.6–1.5 mg/dL 53–133 μmol/L Glucose, fasting (P) 70–110 mg/dL 3.9–6.1 mmol/L Iron (S) 50–150 μg/dL 9.0–26.9 μmol/L Lactic acid (B) 0.5–2.2 meq/L 0.5–2.2 mmol/L Lipase (S) 3–19 units/L Lipids, total (S) 450–1000 mg/dL 4.5–10 g/L Magnesium (S) 1.4–2.0 meq/L 0.7–1.0 mmol/L Osmolality (S) 280–296 mosm/kg H2O 280–296 mmol/kg H2O PCO2 (arterial) (B) 35–45 mm Hg 4.7–6.0 kPa Pepsinogen (P) 200–425 units/mL pH (B) 7.35–7.45 Phenylalanine (S) 0–2 mg/dL 0–120 μmol/L Phosphatase, acid (S) Males: 0–0.8 sigma unit/mL Females: 0.01–0.56 sigma unit/mL Phosphatase, alkaline (S) 13–39 units/L (adults) 0.22–0.65 μmol s–1/L Phospholipids (S) 9–16 mg/dL as lipid phosphorus 2.9–5.2 mmol/L Phosphorus, inorganic (S) 2.6–4.5 mg/dL (infants in first year: up to 6.0 mg/dL) 0.84–1.45 mmol/L PO2 (arterial) (B) 75–100 mm Hg 10.0–13.3 kPa Potassium (S) 3.5–5.0 meq/L 3.5–5.0 mmol/L Protein Total (S) 6.0–8.0 g/dL 60–80 g/L Albumin (S) 3.1–4.3 g/dL 31–43 g/L Globulin (S) 2.6–4.1 g/dL 26–41 g/L Pyruvic acid (P) 0–0.11 meq/L 0–110 μmol/L Sodium (S) 135–145 meq/L 135–145 mmol/L Urea nitrogen (S) 8–25 mg/dL 2.9–8.9 mmol/L Uric acid (S) Women 2.3–6.6 mg/dL 137–393 μmol/L Men 3.6–8.5 mg/dL 214–506 μmol/L aBased in part on Kratz A, et al. Laboratory reference values. N Engl J Med 2004;351:1548. Ranges vary somewhat from one laboratory to another depending on the details of the methods used, and specific values should be considered in the context of the range of values for the laboratory that made the determination. a LANGE medical book Ganong’s Review of Medical Physiology Twenty-Third Edition Kim E. Barrett, PhD Scott Boitano, PhD Professor Associate Professor, Physiology Department of Medicine Arizona Respiratory Center Dean of Graduate Studies Bio5 Collaborative Research Institute University of California, San Diego University of Arizona La Jolla, California Tucson, Arizona Susan M. Barman, PhD Heddwen L. Brooks, PhD Professor Associate Professor Department of Pharmacology/Toxicology Department of Physiology Michigan State University College of Medicine East Lansing, Michigan University of Arizona Tucson, Arizona New York Chicago San Francisco Lisbon London Madrid Mexico City Milan New Delhi San Juan Seoul Singapore Sydney Toronto Copyright © 2010 by The McGraw-Hill Companies, Inc. All rights reserved. Except as permitted under the United States Copyright Act of 1976, no part of this publication may be reproduced or distributed in any form or by any means, or stored in a database or retrieval system, without the prior written permission of the publisher. ISBN: 978-0-07-160568-7 MHID: 0-07-160568-1 The material in this eBook also appears in the print version of this title: ISBN: 978-0-07-160567-0, MHID: 0-07-160567-3. All trademarks are trademarks of their respective owners. Rather than put a trademark symbol after every occurrence of a trademarked name, we use names in an editorial fashion only, and to the benefit of the trademark owner, with no intention of infringement of the trademark. Where such designations appear in this book, they have been printed with initial caps. McGraw-Hill eBooks are available at special quantity discounts to use as premiums and sales promotions, or for use in corporate training programs. To contact a representative please e-mail us at [email protected]. Medicine is an ever-changing science. As new research and clinical experience broaden our knowledge, changes in treatment and drug therapy are required. The authors and the publisher of this work have checked with sources believed to be reliable in their efforts to provide information that is complete and generally in accord with the standards accepted at the time of publication. However, in view of the possibility of human error or changes in medical sciences, neither the authors nor the publisher nor any other party who has been involved in the preparation or publication of this work warrants that the information contained herein is in every respect accurate or complete, and they disclaim all responsibility for any errors or omissions or for the results obtained from use of the infor- mation contained in this work. Readers are encouraged to confirm the information contained herein with other sources. For example and in particular, readers are advised to check the product information sheet included in the package of each drug they plan to administer to be certain that the information contained in this work is accurate and that changes have not been made in the recommended dose or in the contraindications for administration. This recommendation is of particular importance in connection with new or infrequently used drugs. TERMS OF USE This is a copyrighted work and The McGraw-Hill Companies, Inc. (“McGraw-Hill”) and its licensors reserve all rights in and to the work. Use of this work is subject to these terms. Except as permitted under the Copyright Act of 1976 and the right to store and retrieve one copy of the work, you may not decompile, disassemble, reverse engineer, reproduce, modify, create derivative works based upon, transmit, distribute, disseminate, sell, publish or sublicense the work or any part of it without McGraw-Hill’s prior consent. You may use the work for your own noncommercial and personal use; any other use of the work is strictly prohibited. Your right to use the work may be terminated if you fail to comply with these terms. THE WORK IS PROVIDED “AS IS.” McGRAW-HILL AND ITS LICENSORS MAKE NO GUARANTEES OR WARRANTIES AS TO THE ACCURACY, ADEQUACY OR COMPLETENESS OF OR RESULTS TO BE OBTAINED FROM USING THE WORK, INCLUDING ANY INFORMATION THAT CAN BE ACCESSED THROUGH THE WORK VIA HYPERLINK OR OTHERWISE, AND EXPRESSLY DISCLAIM ANY WARRANTY, EXPRESS OR IMPLIED, INCLUDING BUT NOT LIMITED TO IMPLIED WARRANTIES OF MERCHANTABILITY OR FITNESS FOR A PARTICULAR PURPOSE. McGraw-Hill and its licensors do not warrant or guarantee that the functions contained in the work will meet your requirements or that its operation will be unin- terrupted or error free. Neither McGraw-Hill nor its licensors shall be liable to you or anyone else for any inaccuracy, error or omission, regardless of cause, in the work or for any damages resulting therefrom. McGraw-Hill has no responsibility for the content of any information accessed through the work. Under no circumstances shall McGraw-Hill and/or its licensors be liable for any indirect, incidental, special, punitive, consequential or similar damages that result from the use of or inability to use the work, even if any of them has been advised of the possibility of such damages. This limitation of liability shall apply to any claim or cause whatsoever whether such claim or cause arises in contract, tort or otherwise. Dedication to WILLIAM FRANCIS GANONG William Francis (“Fran”) Ganong was an outstanding scien- among physiologists. He was an excellent writer and far ahead tist, educator, and writer. He was completely dedicated to the of his time with his objective of distilling a complex subject into field of physiology and medical education in general. Chair- a concise presentation. Like his good friend, Dr. Jack Lange, man of the Department of Physiology at the University of Cal- founder of the Lange series of books, Fran took great pride in ifornia, San Francisco, for many years, he received numerous the many different translations of the Review of Medical Physi- teaching awards and loved working with medical students. ology and was always delighted to receive a copy of the new edi- Over the course of 40 years and some 22 editions, he was the tion in any language. sole author of the best selling Review of Medical Physiology, and He was a model author, organized, dedicated, and enthusias- a co-author of 5 editions of Pathophysiology of Disease: An tic. His book was his pride and joy and like other best-selling Introduction to Clinical Medicine. He was one of the “deans” of authors, he would work on the next edition seemingly every the Lange group of authors who produced concise medical text day, updating references, rewriting as needed, and always ready and review books that to this day remain extraordinarily popu- and on time when the next edition was due to the publisher. He lar in print and now in digital formats. Dr. Ganong made a did the same with his other book, Pathophysiology of Disease: gigantic impact on the education of countless medical students An Introduction to Clinical Medicine, a book that he worked on and clinicians. meticulously in the years following his formal retirement and A general physiologist par excellence and a neuroendocrine appointment as an emeritus professor at UCSF. physiologist by subspecialty, Fran developed and maintained a Fran Ganong will always have a seat at the head table of the rare understanding of the entire field of physiology. This greats of the art of medical science education and communi- allowed him to write each new edition (every 2 years!) of the cation. He died on December 23, 2007. All of us who knew Review of Medical Physiology as a sole author, a feat remarked him and worked with him miss him greatly. on and admired whenever the book came up for discussion iii Key Features of the 23rd Edition of Ganong’s Review of Medical Physiology •Thoroughly updatedto reflect the latest research and developments in important areas such as the cellular basis of neurophysiology •Incorporates examples from clinical medicinethroughout the chapters to illustrate important physiologic concepts •Delivers more detailed, clinically-relevant, high-yield information per pagethan any similar text or review •NEW full-color illustrations—the authors have worked with an outstanding team of medical illustrators, photographers, educators, and students to provide an unmatched collection of 600 illustrations and tables •NEW boxed clinical cases—featuring examples of diseases that illustrate important physiologic principles •NEW high-yield board reviewquestions at the end of each chapter •NEW larger 8½ X 11” trim-sizeenhances the rich visual content •NEW companion online learning center (LangeTextbooks.com) offers a wealth of innovative learning tools and illustrations NEW iPod-compatible review—Medical PodClass offers audio and text for study on the go Full-color illustrations enrich the text iv KEY FEATURES v Clinical Cases illustrate essential physiologic principles Summary tables and charts encapsulate important information Chapters conclude with Chapter Summaries and review questions About the Authors KIM E. BARRETT American Physiological Society (APS) and recently served on its council. She has also served as Chair of the Central Nervous Kim Barrett received her PhD in biological System Section of APS as well as Chair of both the Women in chemistry from University College London Physiology and Section Advisory Committees of APS. In her in 1982. Following postdoctoral training at spare time, she enjoys daily walks, aerobic exercising, and the National Institutes of Health, she joined mind-challenging activities like puzzles of various sorts. the faculty at the University of California, San Diego, School of Medicine in 1985, rising SCOTT BOITANO to her current rank of Professor of Medicine in 1996. Since 2006, she has also served the Scott Boitano received his PhD in University as Dean of Graduate Studies. Her genetics and cell biology from research interests focus on the physiology and pathophysiology Washington State University in of the intestinal epithelium, and how its function is altered by Pullman, Washington, where he commensal, probiotics, and pathogenic bacteria as well as in acquired an interest in cellular signaling. specific disease states, such as inflammatory bowel diseases. She He fostered this interest at University has published almost 200 articles, chapters, and reviews, and has of California, Los Angeles, where received several honors for her research accomplishments he focused his research on second including the Bowditch and Davenport Lectureships from the messengers and cellular physiology of the lung epithelium. He American Physiological Society and the degree of Doctor of continued to foster these research interests at the University of Medical Sciences, honoris causa, from Queens University, Belfast. Wyoming and at his current positions with the Department of She is also a dedicated and award-winning instructor of medical, Physiology and the Arizona Respiratory Center, both at the pharmacy, and graduate students, and has taught various topics University of Arizona. in medical and systems physiology to these groups for more than 20 years. Her teaching experiences led her to author a prior HEDDWEN L. BROOKS volume (Gastrointestinal Physiology, McGraw-Hill, 2005) and she is honored to have been invited to take over the helm of Heddwen Brooks received her PhD from Ganong. Imperial College, University of London and is an Associate Professor in the Department of Physiology at the University SUSAN M. BARMAN of Arizona (UA). Dr Brooks is a renal Susan Barman received her PhD in physiologist and is best known for her physiology from Loyola University School development of microarray technology of Medicine in Maywood, Illinois. Afterward to address in vivo signaling pathways she went to Michigan State University involved in the hormonal regulation of (MSU) where she is currently a Professor renal function. Dr Brooks’ many awards include the American in the Department of Pharmacology/ Physiological Society (APS) Lazaro J. Mandel Young Investigator Toxicology and the Neuroscience Program. Award, which is for an individual demonstrating outstanding Dr Barman has had a career-long interest in promise in epithelial or renal physiology. She will receive the neural control of cardiorespiratory function APS Renal Young Investigator Award at the 2009 annual with an emphasis on the characterization meeting of the Federation of American Societies for and origin of the naturally occurring discharges of sympathetic Experimental Biology. Dr Brooks is a member of the APS and phrenic nerves. She was a recipient of a prestigious National Renal Steering Section and the APS Committee of Institutes of Health MERIT (Method to Extend Research in Committees. She is on the Editorial Board of the American Time) Award. She is also a recipient of an Outstanding University Journal of Physiology-Renal Physiology (since 2001), and she Woman Faculty Award from the MSU Faculty Professional has also served on study sections of the National Institutes of Women's Association and an MSU College of Human Medicine Health and the American Heart Association. Distinguished Faculty Award. She has been very active in the vi Contents Preface ix I S E C T I O N 15. Electrical Activity of the Brain, Sleep–Wake CELLULAR & MOLECULAR BASIS FOR States, & Circadian Rhythms 229 MEDICAL PHYSIOLOGY 1 16. Control of Posture & Movement 241 17. The Autonomic Nervous System 261 1. General Principles & Energy Production in Medical Physiology 1 18. Hypothalamic Regulation of Hormonal Functions 273 2. Overview of Cellular Physiology in Medical Physiology 31 19. Learning, Memory, Language, & Speech 289 3. Immunity, Infection, & Inflammation 63 IV II S E C T I O N S E C T I O N ENDOCRINE & REPRODUCTIVE PHYSIOLOGY OF NERVE PHYSIOLOGY 301 & MUSCLE CELLS 79 20. The Thyroid Gland 301 4. Excitable Tissue: Nerve 79 21. Endocrine Functions of the Pancreas & Regulation of 5. Excitable Tissue: Muscle 93 Carbohydrate Metabolism 315 6. Synaptic & Junctional Transmission 115 22. The Adrenal Medulla & 7. Neurotransmitters & Neuromodulators 129 Adrenal Cortex 337 8. Properties of Sensory Receptors 149 23. Hormonal Control of Calcium and Phosphate Metabolism & 9. Reflexes 157 the Physiology of Bone 363 III 24. The Pituitary Gland 377 S E C T I O N CENTRAL & PERIPHERAL 25. The Gonads: Development & Function NEUROPHYSIOLOGY 167 of the Reproductive System 391 V 10. Pain & Temperature 167 S E C T I O N GASTROINTESTINAL 11. Somatosensory Pathways 173 PHYSIOLOGY 429 12. Vision 181 26. Overview of Gastrointestinal 13. Hearing & Equilibrium 203 Function & Regulation 429 14. Smell & Taste 219 vii viii CONTENTS VII 27. Digestion, Absorption, & S E C T I O N Nutritional Principles 451 RESPIRATORY PHYSIOLOGY 587 28. Gastrointestinal Motility 469 35. Pulmonary Function 587 29. Transport & Metabolic 36. Gas Transport & pH in the Lung 609 Functions of the Liver 479 37. Regulation of Respiration 625 VI S E C T I O N VIII CARDIOVASCULAR S E C T I O N PHYSIOLOGY 489 RENAL PHYSIOLOGY 639 30. Origin of the Heartbeat & the 38. Renal Function & Micturition 639 Electrical Activity of the Heart 489 39. Regulation of Extracellular Fluid 31. The Heart as a Pump 507 Composition & Volume 665 32. Blood as a Circulatory Fluid & the 40. Acidification of the Urine & Dynamics of Blood & Lymph Flow 521 Bicarbonate Excretion 679 33. Cardiovascular Regulatory Mechanisms 555 Answers to Multiple Choice Questions 687 34. Circulation Through Special Regions 569 Index 689

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