Basic and Applied Bone Biology PDF
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BASIC AND APPLIED BONE BIOLOGY http://booksite.elsevier.com/9780124160156 Basic and Applied Bone Biology David B. Burr and Matthew R. Allen, Editors Resources for Professors: All figures from this book available as both Power Point slides and JPEG files BASIC AND APPLIED BONE BIOLOGY Edited by D B. B AVID URR DepartmentofAnatomyandCellBiology,IndianaUniversitySchoolofMedicine,and DepartmentofBiomedicalEngineering, IndianaUniversity-PurdueUniversityIndianapolis(IUPUI), Indianapolis,Indiana,USA M R. A ATTHEW LLEN DepartmentofAnatomyandCellBiology,IndianaUniversitySchoolofMedicine, Indianapolis,Indiana,USA AMSTERDAM(cid:129)BOSTON(cid:129)HEIDELBERG(cid:129)LONDON(cid:129)NEWYORK(cid:129)OXFORD PARIS(cid:129)SANDIEGO(cid:129)SANFRANCISCO(cid:129)SINGAPORE(cid:129)SYDNEY(cid:129)TOKYO AcademicPressisanimprintofElsevier AcademicPressisanimprintofElsevier 32JamestownRoad,LondonNW17BY,UK 225WymanStreet,Waltham,MA02451,USA 525BStreet,Suite1800,SanDiego,CA92101-4495,USA Copyrightr2014ElsevierInc.Allrightsreserved Covercreditlines: Mainimage:Photographofhumanhumerus.Dr.MatthewR.Allen Upperleft:Photomicrographoffracturehealinginaratbone.Dr.DavidB.BurrandDr.KeithW.Condon Leftpaneltwo:Photomicrographoffluorochromelabelsinamouseulna.Dr.AlexanderG.Robling Leftpanelthree:Scanningelectronmicrographofosteocyte-lacunarnetwork.ImagebyDrs.LilianPlotkin,TeresitaBellido, andLyndaBonewaldandpseudocolorbyDr.AlexanderG.Robling Leftlowerpanel:Atomicforcemicroscopyimageofcollagenfibernetworkinbone.Dr.JosephWallace. 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Becauseofrapidadvancesinthemedicalsciences,inparticular,independentverificationofdiagnosesanddrugdosages shouldbemade BritishLibraryCataloguing-in-PublicationData AcataloguerecordforthisbookisavailablefromtheBritishLibrary LibraryofCongressCataloging-in-PublicationData AcatalogrecordforthisbookisavailablefromtheLibraryofCongress ISBN:978-0-12-416015-6 ForinformationonallAcademicPresspublications visitourwebsiteatelsevierdirect.com TypesetbyMPSLimited,Chennai,India www.adi-mps.com PrintedandboundinChina 14 15 16 17 10 9 8 7 6 5 4 3 2 1 Dedications This book isdedicated to the wonderful teachers Ihave had over many years—Denny, Bruce,Harold, Eric,Mitch, andCharles, among many others—who have sopatiently taught me about the beauties and intricacies of our skeleton. And to my wife, Lisa, and son, Erik, who havetolerated and supported myobsession with bone. —DavidB. Burr This bookis dedicated to my late father,who was,and always willbe, themostinfluential teacher in mylife. And to Kristine, Sophie, and Gus who provide me with a daily reminder that there is somuch moreto life than bone biology. —Matthew R. Allen Preface More than 10 years ago when we began to teach Basic and Applied Bone Biology covers those topics that our graduate-level Basic Bone Biology course at we feel are relevant to a modern course in skeletal biol- Indiana University, there were several excellent refer- ogy. The book is organized, like bone, in a somewhat ence works available, primarily targeted to research- hierarchical manner. The first section begins with the ers working in a wide range of areas in skeletal basic construction of bone, including its cellular struc- biology. These included (and still include) Principles tureanddynamicsandthebasicphysiologicalprocesses of Bone Biology, edited by John Bilezikian, Lawrence that bone uses to grow and adapt itself over a lifetime. Raisz, and John Martin, which has since been This is succeeded by several chapters related to the expanded to two volumes; “Big Red” (Osteoporosis), technical aspects used to assess bone in health and the excellent and very complete reference edited by disease—various imaging modalities; biomechanical Bob Marcus, David Feldman, and Jennifer Kelsey; measurements useful for assessing bone properties; his- and the more succinct Primer of Metabolic Bone tomorphometric techniques to evaluate the dynamics of Diseases, updated and republished every few years by bonemodelingandremodeling;andgeneticapproaches the American Society of Bone and Mineral Research. usedtoteaseouttherolesofspecificgenesandproteins, These are still available, and still excellent, but they and epigenetic influences in the basic metabolic do not serve well as textbooks for a bone biology functions of bone. These early chapters provide the course either because they are too extensive, too foundation for the next several chapters on skeletal expensive, or do not cover relevant topics in sufficient adaptation, highlighting mechanically-induced adaption depth. Therefore, we have chosen over the years to ofbone,fracturehealing,andadaptationoftheoralcav- use primary reference materials—mostly, recent ity associated with orthodontics and implants. papers published in the peer reviewed literature—for Following this, the text transitions (gradually we hope) our course. From a didactic standpoint, this is an into areas that are more clinically related, the applied acceptable approach, and even a desirable one, espe- aspects referred to in the title. These chapters address cially for a graduate course in which the goal is to growth and development, metabolic and hormonal pro- teach the student how to read and evaluate the litera- cesses, and how these are related to health and disease. ture. However, it became clear over time that this was The text ends with a chapter on pharmaceutical treat- not a sufficient surrogate for a true textbook. ments for osteoporosis, which we hope incorporates As the skeletal biology group at Indiana University boththeclinicalelementsoftreatmentandthebiological grew over the years, we incorporated topic experts to reasonsfor,andeffectsof,thesetreatments. deliver lectures intheir area ofexpertise.Wesoon real- Skeletal biology is, by its nature, interdisciplinary. ized that the course and content experts provided the The course that we teach at Indiana University typi- foundation for building a textbook on basic and cally includes students in the basic medical sciences, applied skeletal biology. As we discussed this idea general biology, the dental sciences, several engineer- with our colleagues here at Indiana University and up ing subspecialties, foods and nutrition, kinesiology, the road at Purdue University, there was universal and rehabilitation sciences. We have written this text- support and enthusiasm. Discussion with colleagues book to cover a range of topics that we feel would be outside of our group made it clear that, beyond our relevant to these groups, and have attempted to write own requirements, there was a need and a desire by various chapters in a way that will be understandable the academic community for such a text. Writing this to those students whose particular expertise and inter- textbook began as something of a selfish idea—we est may not be in the area covered by a given chapter. needed it for our course—but we truly hope that it We have also attempted to write the chapters so that will be welcomed and used by others who find it they will be suitable for students at various levels of appropriate for their owncourses,or as a more modest study, including undergraduate, graduate, and even reference than existing books on a wide range of topics postgraduate. We realize that the danger of this is that in skeletal biology. some chapters may be too superficial for students who xi xii PREFACE are more expert in the area covered by that chapter. be a single “correct” answer. We hope that they will However, the textbook is meant to be supplemented permit further exploration of the chapter topic, at the by additional readings that delve into specific topics in level appropriate for thestudent. greater depth for those who wish to specialize in that Finally, we have not only had a lot of fun putting area. To this end, we have included a list of 10(cid:1)15 this text together but have also learned a lot in areas suggested readings at the end of each chapter that can that are not within our own expertise. We sincerely serve as a starting point for supplementary reading hope that it serves the same purpose for you. and discussion. Further, we have incorporated study questions at the end of each chapter. We have resisted David B. Burr,PhD the temptation to include answers to these questions. Matthew R. Allen, PhD They are intended to be used for discussion (although they could also be used for testing), and there may not 16 February 2013 Acknowledgements We would like to thank the students/post-doctoral We are grateful to the following individuals for con- fellows who reviewed and provided critical critiques tributingto original figures: of each chapter: Dr. Keith W. Condon for several histological images in chapters 1,2, and 7 Rachel Dirks Mr. Drew Brown for numerous illustrations in Paul Childress Chapter 4 as well as figures 1.4, 9.5, 9.6, 17.6, 17.9, and Chris Newman 17.11 Lisa Cole Dr. Joseph Wallace for figures 1.1 and 1.3 Dr. JunSun Dr. Mitch Schaffler for figures 1.9 and 1.11 Joshua Gargac Dr. NicolettaBivi for figure 2.2 Dr. HeatherCoan Dr. Alex Robling for figure 4.6 Amy Sato Dr. StuartWardenfor figure 5.15 and9.17 Dr. Emily Farrow Rafael Pacheco-Costa and Dr. Lynn Neff for Perla Reyes-Fernandez figure 2.3 DennisJoseph Drs. Lilian I. Plotkin, Teresita Bellido, and Lynda Rebecca McCreedy Bonewald for the original SEM in 9.11 MaximeGallant Dr. Hau Zhou for the clinical photomicrographs in We would also like to thank Dr. Jason Organ for figures 7.16,7.17, 7.18 and 7.21 editing thequestionsat the end ofeachchapter. Dr. Steven Doty for the EM images in 2.9 We would also like to thank Dr. James Fleet for con- tributingto the early drafts ofChapter 13. xiii List of Contributors Ozan Akkus, PhD Department of Mechanical and Kathleen M. Hill Gallant, PhD, RD Department of AerospaceEngineering,BiomedicalEngineering,and AnatomyandCellBiology,IndianaUniversitySchoolof OrthopaedicSurgery,CaseWesternReserveUniversity, Medicine,Indianapolis,Indiana,USA Cleveland,Ohio,USA Erik A. Imel, MD Departments of Medicine and Matthew R. Allen, PhD Department of Anatomy and pediatrics,IndianaUniversitySchoolofMedicine, CellBiology,IndianaUniversitySchoolofMedicine, Indianapolis,Indiana,USA Indianapolis,Indiana,USA Daniel L. Koller, PhD Department of Medical and William J Babler, PhD Department of Oral Biology, MolecularGenetics,IndianaUniversitySchoolof IndianaUniversitySchoolofDentistry,Indianapolis, Medicine,Indianapolis,Indiana,USA Indiana,USA Kelly Krohn, MD Senior Medical Advisor, Eli Lilly Teresita Bellido, PhD Department of Anatomy and Cell Laboratories,Indianapolis,Indiana,USA Biology,IndianaUniversitySchoolofMedicine, Jiliang Li,PhD, MD DepartmentofBiologyandCenterfor Indianapolis,Indiana,USA DevelopmentalandRegenerativeBiology,Indiana Nicoletta Bivi, PhD Department of Anatomy and Cell University-PurdueUniversityIndianapolis,Indianapolis, Biology,IndianaUniversitySchoolofMedicine, Indiana,USA Indianapolis,Indiana,USA BruceH.Mitlak,MD DistinguishedMedicalFellow,Lilly Angela Bruzzaniti, PhD Department of Oral Biology, ResearchLaboratories,Indianapolis,Indiana,USA IndianaUniversitySchoolofDentistryandDepartmentof Lilian I. Plotkin, PhD Department of Anatomy and Cell Anatomy&CellBiology,IndianaUniversitySchoolof Biology,IndianaUniversitySchoolofMedicine, Medicine,Indianapolis,Indiana,USA Indianapolis,Indiana,USA David B. Burr, PhD Department of Anatomy and Cell AlexanderG.Robling,PhD DepartmentofAnatomyand Biology,IndianaUniversitySchoolofMedicine, CellBiology,IndianaUniversitySchoolofMedicine, Indianapolis,Indiana,USA Indianapolis,Indiana,USA Tim Corbin, MS Department of Therapeutic Discovery, Sean Shih-Yao Liu, DDS, MS, PhD Department of TransgenicDivision,AmgenInc.,ThousandOaks, Orthodontics,IndianaUniversitySchoolofDentistry, California,USA Indianapolis,Indiana,USA Linda A. DiMeglio, MD, MPH Department of Pediatrics, David L. Stocum, PhD Department of Biology and IndianaUniversitySchoolofMedicine,Indianapolis, CenterforDevelopmentalandRegenerativeBiology, Indiana,USA IndianaUniversity-PurdueUniversityIndianapolis, Robyn K. Fuchs, PhD Department of Health and Indianapolis,Indiana,USA RehabilitationSciences,IndianaUniversitySchoolof Joseph M. Wallace, PhD Department of Biomedical Medicine,DepartmentofPhysicalTherapy,Schoolof Engineering,IndianaUniversity-PurdueUniversity HealthandRehabilitationSciences,IndianaUniversity, Indianapolis,Indianapolis,Indiana,USA Indianapolis,USA Connie M. Weaver, PhD Nutrition Science, Purdue Tien-Min Gabriel Chu, DDS, PhD Department of University,WestLafayette,Indianapolis,USA RestorativeDentistry,IndianaUniversitySchoolof Dentistry,Indianapolis,Indiana,USA Kenneth E. White, PhD Department of Medical and MolecularGenetics,IndianaUniversitySchoolof Medicine,Indianapolis,Indiana,USA xv C H A P T E R 1 Bone Morphology and Organization 1 2 David B. Burr and Ozan Akkus 1Department ofAnatomy and Cell Biology, Indiana University School of Medicine, Indianapolis, Indiana, USA 2Department ofMechanical and Aerospace Engineering, Biomedical Engineering, andOrthopaedic Surgery,Case Western Reserve University,Cleveland, Ohio, USA THE FUNCTIONS OF BONE to absorb impact energy. Developmentally, bones that serve a protective function (e.g. calvarium and ribs) Bone is multifunctional, playing roles in mechanical are formed, at least in part, through intramembranous support and protection, mineral homeostasis, and ossification rather than through endochondral ossifica- hematopoiesis. In recent years, it has become clear that tion, (see Chapter 4)and are part of the axial skeleton. bone also serves an important endocrinefunction. It is not widely realized that bone is a blood- The mechanical functions of bone are by far the forming (hematopoietic) organ, but regions composed most widely recognized and studied. Both trabecular largely of spongy bone such as the iliac crest, vertebrae, and cortical bone serve this function, although the and proximal femur are good sources of red blood nature of this function is partly specific to each. The cells throughout life. The marrow cavity within the dense cortical bone comprises most of the bone mass bone is an important site of red marrow, indicative of and takes on most of the role for load bearing. hematopoiesis, during growth and development, but is Although the more porous cancellous bone also sup- largelycomposed of yellowfat in adults. White fat and ports load, one of its important functions is to redirect brown fat are also found in the human body, and stresses to the stronger cortical shell. The mechanical while these are acted upon by osteocalcin, which is function of bone extends beyond simple load bearing, produced by osteoblasts, these types of fat are not which requires a certain degree of strength and actually found within the bone marrow itself. Yellow stiffness. Because of its organization as a multiscale marrow fat originates from the same precursor cells material, it is also highly adapted to avoid fractures that differentiate to become bone-forming osteoblasts. caused by repetitive loading at physiologic levels, It provides an energy store and may contribute to lipid i.e. failure in fatigue. metabolism by regulating triglycerides. Because of the Bone also serves a protective function, especially in large surface area, regions with a lot of cancellous those vital areas such as the torso and head where bone are also responsible for rapid turnover of bone injury can be fatal. In these locations, the bone micro- tissue and play an important role in the long-term con- structure is not different from that of bone in other trol of calciumbalance. locations, but it is organized in a manner that can Bone turnover can be sensitive to changes in energy absorb maximum energy with minimum trauma to the metabolism that occur as a function of aging, hormone bone itself. For instance, the cranial vault is con- deficiency, or the production of skeletal hormones, and structed of two thin plates of dense bone that sand- this provides the means for long-term exchange of cal- wich porous cancellous bone (the porous appearance cium and phosphate (as well as other minerals such as of this bone is why it is sometimes called spongy bone). iron and magnesium). Although calcium provides Ribs are also constructed in this way, but with less bone with its stiffness and much of its strength, cal- dense cancellous bone. In the case of the ribs, the cium ions in the mineral phase are also important for inherent curvature of the bone also increases its ability enzyme reactions, blood clotting, muscle contraction, BasicandAppliedBoneBiology. DOI:http://dx.doi.org/10.1016/B978-0-12-416015-6.00001-0 3 ©2014ElsevierInc.Allrightsreserved.
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