RobertB.Rucker/HandbookofVitamins,FourthEdition 4022_C000 FinalProof page i 5.5.2007 2:33pm CompositorName:BMani F O U RT H E D I T I O N Handbook of V ITA MINS RobertB.Rucker/HandbookofVitamins,FourthEdition 4022_C000 FinalProof page ii 5.5.2007 2:33pm CompositorName:BMani RobertB.Rucker/HandbookofVitamins,FourthEdition 4022_C000 FinalProof page iii 5.5.2007 2:33pm CompositorName:BMani F O U RT H E D I T I O N Handbook of V ITA MINS Edited by Janos Zempleni Robert B. Rucker Donald B. McCormick John W. Suttie Boca Raton London New York CRC Press is an imprint of the Taylor & Francis Group, an informa business RobertB.Rucker/HandbookofVitamins,FourthEdition 4022_C000 FinalProof page iv 5.5.2007 2:33pm CompositorName:BMani CRC Press Taylor & Francis Group 6000 Broken Sound Parkway NW, Suite 300 Boca Raton, FL 33487-2742 © 2007 by Taylor & Francis Group, LLC CRC Press is an imprint of Taylor & Francis Group, an Informa business No claim to original U.S. Government works Printed in the United States of America on acid-free paper 10 9 8 7 6 5 4 3 2 1 International Standard Book Number-10: 0-8493-4022-5 (Hardcover) International Standard Book Number-13: 978-0-8493-4022-2 (Hardcover) This book contains information obtained from authentic and highly regarded sources. Reprinted material is quoted with permission, and sources are indicated. A wide variety of references are listed. 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Trademark Notice: Product or corporate names may be trademarks or registered trademarks, and are used only for identification and explanation without intent to infringe. Library of Congress Cataloging-in-Publication Data Handbook of vitamins / editors, Robert B. Rucker ... [et al.]. -- 4th ed. p. ; cm. Includes bibliographical references and index. ISBN-13: 978-0-8493-4022-2 (hardcover : alk. paper) ISBN-10: 0-8493-4022-5 (hardcover : alk. paper) 1. Vitamins. 2. Vitamins in human nutrition. I. Rucker, Robert B. [DNLM: 1. Nutrition Physiology--Handbooks. 2. Vitamins--Handbooks. QU 39 H361 2007] QP771. H35 2007 612.3’99--dc22 2006100138 Visit the Taylor & Francis Web site at http://www.taylorandfrancis.com and the CRC Press Web site at http://www.crcpress.com RobertB.Rucker/HandbookofVitamins,FourthEdition 4022_C000 FinalProof page v 5.5.2007 2:33pm CompositorName:BMani Table of Contents Preface. .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... vii Editors . .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... ix Contributors .. .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... xi Chapter 1 VitaminA: NutritionalAspectsof Retinoids and Carotenoids. .. ... .. ... . 1 A. Catharine Ross andEarlH. Harrison Chapter 2 VitaminD . ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... 41 Anthony W. Norman and Helen L. Henry Chapter 3 VitaminK . ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. .. 111 John W.Suttie Chapter 4 VitaminE.. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. .. 153 Maret G.Traber Chapter 5 BioorganicMechanisms Importantto CoenzymeFunctions... .. ... .. .. 175 Donald B. McCormick Chapter 6 Niacin .. .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. .. 191 James B. Kirkland Chapter 7 Riboflavin (Vitamin B ) . ... .. ... .. ... .. ... .. ... .. ... .. ... .. .. 233 2 Richard S. Rivlin Chapter 8 Thiamine .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. .. 253 Chris J.Bates Chapter 9 Pantothenic Acid . ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. .. 289 RobertB. Rucker and KathrynBauerly Chapter 10 VitaminB ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. .. 315 6 Shyamala Dakshinamurti and Krishnamurti Dakshinamurti v RobertB.Rucker/HandbookofVitamins,FourthEdition 4022_C000 FinalProof page vi 5.5.2007 2:33pm CompositorName:BMani vi TableofContents Chapter 11 Biotin. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. 361 DonaldM.Mock Chapter 12 FolicAcid. .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. 385 LynnB. Bailey Chapter 13 Vitamin B . ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. 413 12 Ralph GreenandJoshua W. Miller Chapter 14 Choline... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. 459 TimothyA. Garrow Chapter 15 Ascorbic Acid... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. 489 CarolS. Johnston,Francene M. Steinberg, and Robert B.Rucker Chapter 16 Vitamin-Dependent Modifications of Chromatin:Epigenetic Events and Genomic Stability.. .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. 521 James B.Kirkland, Janos Zempleni, Linda K. Buckles, and Judith K. Christman Chapter 17 Accelerator Mass Spectrometry in the Study ofVitamins and Mineral Metabolism in Humans . ... .. ... .. ... .. ... .. ... .. ... .. 545 FabianaFonsecade Moura, Betty Jane Burri, and Andrew J.Clifford Chapter 18 DietaryReference Intakes for Vitamins. ... .. ... .. ... .. ... .. ... .. 559 SuzanneP. Murphy and Susan I.Barr Index . ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. ... .. 571 RobertB.Rucker/HandbookofVitamins,FourthEdition 4022_C000 FinalProof page vii 5.5.2007 2:33pm CompositorName:BMani Preface In keeping with the tradition of previous editions, the fourth edition of the Handbook of Vitamins was assembled to update and provide contemporary perspectives on dietary acces- soryfactorscommonlyclassifiedasvitamins.Oneofthechallengesinassemblingthisvolume was to maintain the clinical focus of previous editions, while addressing important concepts thathaveevolved inrecentyears owingtotheadvancesinmolecular andcellularbiology as wellasthoseinanalyticalchemistryandnanotechnology.Thereaderwillfindcomprehensive summariesthatfocusonchemical,physiological,andnutritionalrelationshipsandhighlights of newly described and identified functions for all the recognized vitamins. Our goal was to assemblethebestcurrentlyavailablereferencetextonvitaminsforanaudiencerangingfrom basicscientiststoclinicianstoadvancedstudentsandeducatorswithacommitmenttobetter understanding vitamin function. Asexamples,apparentvitamin-dependentmodificationsthatareimportanttoepigenetic events and genomic stability are described, as well as new information on the role and importance of maintaining optimal vitamin status for antioxidant and anti-inflammatory defense.Importantanalyticaladvancesinvitaminanalysisandassessmentarediscussedina chapterdealingwithacceleratedmassspectrometry(AMS)applications.RecentAMSappli- cationshaveprovidedthebasisforstudiesofvitaminmetabolismandturnoverinhumansat levels corresponding to physiological concentrations and fluxes. It is also important to underscore that much of the interest in vitamins stems from an appreciation that there remainsregrettablysizablepopulationsatriskforvitamindeficiencies.Inthisregard,classic examples are included along with examples of vitamin-related polymorphisms and genetic factorsthat influence therelativeneedsfor given vitamins. Thisvolumeiswrittenbyagroupofauthorswhohavemademajorcontributionstoour understanding of vitamins. Over half of the authors are new to this series; each chapter is written by individuals who have made clearly important contributions in their respective areas of research as judged by the scientific impact of their work. In addition, Dr. Janos Zempleni joins the group of editors who assembled the third edition. Dr. Zempleni adds a molecular biology perspectiveto complementthe biochemicaland physiological expertise of the other editors. We also wish to note that we miss the input of Dr. Lawrence Machlin, a renownedresearcheronvitaminE,whowassoleeditorofthefirsttwoeditionsinthisseries andwhodiedshortlyafterthereleaseofthethirdedition.Weknowthathewouldbepleased with theprogress and advancesin vitamin research summarized in thefourth edition. This volume comes at an important time and represents a new treatment of this topic. With the possible exception of the earlier days of vitamin discovery, this period of vitamin research is particularly exciting because of the newly identified roles of vitamins in cellular and organismal regulation and their obvious and continuing importance in health and disease. JanosZempleni RobertB. Rucker Donald B.McCormick John W.Suttie vii RobertB.Rucker/HandbookofVitamins,FourthEdition 4022_C000 FinalProof page viii 5.5.2007 2:33pm CompositorName:BMani RobertB.Rucker/HandbookofVitamins,FourthEdition 4022_C000 FinalProof page ix 5.5.2007 2:33pm CompositorName:BMani Editors JanosZemplenireceivedhisundergraduateandgraduatetraininginnutritionattheUniver- sityofGiesseninGermany.Hereceivedpostdoctoraltraininginnutrition,biochemistry,and molecularbiologyattheUniversityofInnsbruck(Austria),EmoryUniversity,andArkansas Children’sHospitalResearchInstitute.JanosZempleniiscurrentlyanassistantprofessorof molecular nutrition at the University of Nebraska at Lincoln. He has published more than 100 manuscripts and books and is the recipient of the 2006 Mead Johnson Award by the AmericanSocietyforNutrition.Zempleni’sresearchfocusesonrolesofthevitaminbiotinin chromatinstructure. Robert B. Rucker received his PhD in biochemistry from Purdue University in 1968 and workedfortwoyearsasapostdoctoralfellowattheUniversityofMissouri,beforejoiningthe faculty of nutrition at the University of California (UC), Davis, in 1970. He currently holds the title of distinguished professor. He serves as vice chair of the department of nutrition in the College of Agriculture and Environmental Sciences and holds an appointment in endo- crinology, nutrition, and vascular medicine, department of internal medicine, UC Davis, Schoolof Medicine. Dr. Rucker’s research focuses on cofactor function. His current research addresses problems associated with extracellular matrix assembly, the role of copper in early growth and development, and the physiological roles of quinone cofactors derived from tyrosine, such as pyrroloquinolinequinone. Honorsandactivitiesincludeservingasapastpresident,AmericanSocietyforNutrition; appointmentasafellowintheAmericanAssociationfortheAdvancementofScienceandthe American Society for Nutrition; service as chair or cochairperson for FASEB Summer and Gordon Conferences; service on Program and Executive Committees for American Society for Nutrition and FASEB, as well as service on committees of the Society for Experimental BiologyandMedicine.Healsocurrentlyservesasseniorassociateeditor,AmericanJournalof Clinical Nutrition and is a past editorial board member of the Journal of Nutrition, Experi- mentalBiologyandMedicine,NutritionResearch,andtheAnnualReviewofNutrition.Heisa pastrecipient ofUC Davis and theAmerican Society for NutritionResearch awards. DonaldB.McCormickearnedhisbachelor’sdegree(chemistryandmath,1953)anddoctorate (biochemistry, 1958) at Vanderbilt University in Nashville, Tennessee. His dissertation was onpentoseandpentitolmetabolism.HewasthenanNIHpostdoctoralfellow(1958–1960)at the University of California-Berkeley, where his research was on enzymes that convert vitamin B to the coenzyme pyridoxal phosphate. He has had sabbatics in the chemistry 6 departments in Basel University (Switzerland) and in the University of Arizona, and in the biochemistry department inWageningen (Netherlands). Dr.McCormick’sacademicappointmentshavebeenatCornellUniversity(1960–1978)in Ithaca, New York, where he became the Liberty Hyde Bailey Professor of nutritional biochemistry and at Emory University (1979–present) in Atlanta, Georgia, where he served astheFullerE.Callawayprofessorandchairmanofthedepartmentofbiochemistryandthe executiveassociatedeanforbasicsciencesintheschoolofmedicine.Hisresearchhasbeenon ix
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