Ординатура / Офтальмология / Английские материалы / Development of the Ocular Lens_Lovicu, Lee Robinson_2004
.pdfx Contributors
Fisher, Marilyn, Department of Biology, University of Virginia, Gilmer Hall, P.O. Box 400328, Charlottesville, VA 22904-4328, USA, phone: (1-434)-982-5606, fax: (1-434)- 982-5626, e-mail: mf4b@virginia.edu
Goudreau, Guy, Department of Molecular Cell Biology, Max-Planck-Institute of Biophysical Chemistry, Am Fassberg 11, 37070 Gottingen,¨ Germany, phone: (49-551) 201 1361, fax: (49-551) 2011712, e-mail: guy.goudreau@sympatico.ca
Grainger, Robert M., Department of Biology, University of Virginia, Gilmer Hall, P.O. Box 400328, Charlottesville, VA 22904-4328, USA, phone: (1-434) 982-5495, fax: (1-434) 982-5495, e-mail: rmg9p@virginia.edu
Griep, Anne E., Department of Anatomy, University of Wisconsin Medical School, 1300 University Avenue, Madison, WI 53706, USA, phone: (1-608) 262-8988, fax: (1-608) 2627306, e-mail: aegriep@facstaff.wisc.edu
Gruss, Peter, Department of Molecular Cell Biology, Max-Planck-Institute for Biophysical Chemistry, Am Fassberg 11, 37070 Gottingen,¨ Germany, phone: (49-551) 201 1361, fax: (49-551) 201 1504, e-mail: peter.gruss@mpg-gv.mpg.de
Kantorow, Marc, Biomedical Sciences, Florida Atlantic University, Biomedical Sciences, 777 Glades Rd., P.O. Box 3091, Boca Raton, FL 33431-0991, USA, phone: (1-561) 2972910, e-mail: mkantoro@fau.edu
Kistler, Joerg, School of Biological Sciences, University of Auckland, Private Bag 92019, Auckland, New Zealand, phone: (64-9) 3737599 ext. 88250, fax: (64-9) 3737415, e-mail: j.kistler@auckland.ac.nz
Kuszak, Jer R., Departments of Pathology and Ophthalmology, Rush Presbyterian– St. Luke’s Medical Center, 1653 W. Congress Parkway, Chicago, IL 60612, USA, phone: (1-312) 942-5630, fax: (1-312) 942-2371, e-mail: jkuszak@rush.edu
Lang, Richard A., Divisions of Developmental Biology and Ophthalmology, Cincinnati Children’s Hospital Research Foundation, 3333 Burnet Avenue, Cincinnati, OH 45229, USA, phone: (1-513) 636-7030 (assistant), (1-513) 636-2700 (office), fax: (1-513) 6364317, e-mail: Richard.Lang@cchmc.org
Lovicu, Frank J., Save Sight Institute and Department of Anatomy and Histology (F13), Institute for Biomedical Research, University of Sydney, NSW, 2006, Australia, phone: (61-2) 9351 5170, fax: (61-2) 9351 2813, e-mail: lovicu@anatomy.usyd.edu.au
McAvoy, John W., Save Sight Institute and Department of Anatomy and Histology, University of Sydney, Sydney Eye Hospital, Macquarie Street, GPO Box 4337, Sydney 2001, Australia, phone: (61-2) 9382 7369, fax: (61-2) 9382 7318, e-mail: johnm@eye.usyd.edu.au
Menko, A. Sue, Department of Pathology, Anatomy and Cell Biology, Thomas Jefferson University, 571 Jefferson Alumni Hall, 1020 Locust St., Philadelphia, PA 19107-6799, USA, phone: (1-215) 503-2166, fax: (1-215) 923-3808, e-mail: Sue.Menko@jefferson.edu
Piatigorsky, Joram, Laboratory of Molecular and Developmental Biology, National Eye Institute, National Institutes of Health, Building 7, Room 100A, 7 Memorial Drive MSC 0704, Bethesda, MD 20892-2730, USA, phone: (1-301) 496-9467, fax: (1-301) 402-0781, e-mail: joramp@nei.nih.gov
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Prescott, Alan, School of Life Sciences, MSIWTB, University of Dundee, Millers Wynd, Dundee DD1 5EH, UK, phone: (441) 382-344884, fax: (441) 382-345893, e-mail: a.r.prescott@dundee.ac.uk
Quinlan, Roy A., Department of Biological Sciences, The Science Park, South Road, Durham DH1 3LE, UK, phone: (+44) 0191-334-1331, fax: (+44) 0191-334-1207, e-mail: r.a.quinlan@durham.ac.uk
Robinson, Michael L., Division of Molecular and Human Genetics, Children’s Research Institute, Wexner 492, 700 Children’s Drive, Columbus, OH 43205, USA, phone: (1-614) 722-2764, fax: (1-614) 722-2716, e-mail: RobinsoM@pediatrics.ohio-state.edu
Walker, Janice L., Department of Pharmacology, University of Pennsylvania, 167 Johnson Pavillion, 3620 Hamilton Way, Philadelphia, PA 19104, USA, e-mail: janicelwalker 2000@yahoo.com
Zhang, Pumin, Department of Molecular Physiology and Biophysics, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030, USA, phone: (1-713) 798-1866, fax: (1-713) 798-3475, e-mail: pzhang@bcm.tmc.edu
Preface
Like the beginning of the 20th century, the dawn of the 21st century is witnessing tremendous advances in developmental biology. Research on ocular lens development remains at the forefront of these advances, just as it was in 1900. The difference between then and now lies in the current synergism between developmental biology, genetics, and molecular biology that has led to the identification of the very molecules responsible for many of the inductive processes only descriptively defined earlier.
The period from the first descriptions of mice with transgenes targeted specifically to the lens until the present day, when lens-specific gene deletion has become almost routine, spans only about the past 20 years. It is in the context of this breathtaking influx of information that we thought it time to devote a text specifically and exclusively to lens development. Because of its simplicity and its predictable pattern of development and differentiation, the lens, arguably the sparkling jewel of anatomy, has attracted the attention of many developmental biologists. All stages of lens differentiation, from the proliferation of lens epithelial cells to the differentiation of mature, organelle-deficient fiber cells, are represented in the lens of any individual. Analogous to the rings of a tree, the components of the lens, from the newest cells born moments before tissue collection to the oldest cells originating during embryogenesis, offer a key to its life history. This fact, combined with the ability of researchers to observe and manipulate the lens in a living animal, ensures that this remarkable organ will remain a paradigm for fundamental biological investigations for decades to come.
It has been our good fortune to bring together for the first time an international panel of well-known developmental biologists currently working on research problems specifically related to aspects of lens induction, development, differentiation, growth, and maintenance. Each of these scientists has contributed to this up-to-date text covering virtually all aspects of lens development. The 12 chapters of Development of the Ocular Lens provide a current view of research in lens developmental biology, with an emphasis on recent technical and molecular breakthroughs. The introductory chapter covers historical and evolutionary aspects of lens development and structure. It is followed by chapters describing lens induction and formation as well as the transcription factors that provide the molecular control of these processes. Further chapters discuss the development of the lens structure, including its ultrastructure and fiber organization; crystallin synthesis and function; and lens cell membranes and cytoskeleton. The final chapters provide insights into the processes of lens development and growth at the molecular level, including the cell cycle, fiber cell differentiation, cell adhesion, and extracellular matrix and growth factor signaling. The last chapter describes the fascinating field of lens regeneration, bringing lens development full circle, from induction to regeneration. Although all the chapters are written from a developmental
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perspective, where appropriate, discussion of lens pathology related to developmental abnormalities complements the description of basic lens research.
Past texts have dealt with the lens in the context of cataract and molecular biology, and a few recent books have concentrated on eye development as a whole. Development of the Ocular Lens is unique in its unsurpassed depth of information specifically focused on lens development. Written by many of the leading researchers in the field, this text is an invaluable resource for anyone interested in lens biology, providing, among other things, a springboard into the primary research literature. We have been humbled and honored to have taken part in this endeavor, and it is our sincere hope that graduate students, residents, postdoctoral fellows, principal investigators, and clinicians alike will enjoy reading and using this book as much as we have enjoyed assembling it.
We owe a tremendous debt of gratitude to all of the contributing authors. We would sincerely like to thank them for their hard work as well as their patience during the long process of getting this work into its present form. Most importantly, we would like to thank our families, our wives (Maria Lovicu and Julia Robinson) and children (Christopher, Alexander, and Matthew Lovicu and Emily, Eric, Jessica, and Ian Robinson), for giving us the time to assemble and edit a book that we are very proud of.
Frank J. Lovicu
Michael L. Robinson
Acknowledgments
Acknowledgments for specific chapters are as follows: Chapter 1: FJL acknowledges support from the Sydney Foundation for Medical Research and the National Health and Medical Research Council (NHMRC) of Australia. MLR also acknowledges support from the National Eye Institute, USA (EY12995). The authors thank Barbara Van Brimmer of the Medical Heritage Center at the Ohio State University, USA, for her assistance in gathering information on the history of ophthalmology, as well as Professor John W. McAvoy (Save Sight Institute, Australia) for critically reviewing this chapter. Chapter 2: Work cited from the Grainger Lab was funded by NIH Grants EY06675, EY10283, and RR13221. We thank Dr. Nicolas Hirsch for critically reviewing the manuscript. Chapter 4: The authors gratefully acknowledge the contributions of K. J. Al-Ghoul, Ph.D.; H. Brown, M.D., Ph.D.; R. K. Zoltoski, Ph.D.; J. Chrisman; C. F. Freel; K. O. Gilliland; A. J. Kuszak; C. W. Lane; H. Mekeel; R. Nordgren; L. Novak; and K. L. Peterson. JRK has been supported by NIH NEI Grant EY06642, the Regenstein Foundation, the Louise C. Norton Trust, and the Alcon Research Institute. MJC has been supported by NIH NEI Grants EY08148 and EY05722. Human lenses were obtained from the NDRI and the North Carolina Eye Bank. Primate lenses were obtained from the Regional Primate Center in Seattle, Washington, and the Yerkes Primate Center in Atlanta, Georgia. Chapter 6: Research work in the authors’ laboratories was supported by the Health Research Council of New Zealand, the Marsden Fund, the Lotteries Grant Board, the University of Auckland Research Committee, and the Auckland Medical Research Foundation. Chapter 11: We thank Tina Coburn for assistance in the preparation of this chapter. This work was supported by Grants EY03177 (JMcA.) and EY11234, EY10559, EY12370, and EY12370 (RAL) from the National Institutes of Health and by a grant from the National Health and Medical Research Council (NHMRC) of Australia. JMcA acknowledges support from the Sydney Foundation for Medical Research and an Infrastructure Grant from the NSW Department of Health. The Lang Laboratory is also supported by funds from the Abrahamson Pediatric Eye Institute Endowment at Children’s Hospital Medical Center of Cincinnati.
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Part One
Early Lens Development
1
The Lens: Historical and Comparative
Perspectives
Michael L. Robinson and Frank J. Lovicu
1.1. Lens Anatomy and Development (Pre-1900)
The past decade has witnessed a tremendous increase in the basic understanding of the molecules and signal transduction pathways required to initiate embryonic lens development. Other advances in this time period have elucidated structural and physiological properties of lens cells, often in an evolutionary context, making it possible to frame many pathological conditions of the lens as errors of specific developmental events. All of these recent advances rest on the fundamental observations of talented investigators in previous decades and centuries. While several texts describe the history of ophthalmology as a clinical discipline, the conceptual history of basic eye research as a science, and in particular the history of lens development research, is a much less traversed subject. Though it is inevitable that we cannot include all of the many important experiments and personalities that have played fundamental roles in shaping the field of lens development, we hope to stimulate appreciation for those pioneers, both past and present, to whom we owe a debt of gratitude for their contributions to the field.
Throughout human history, the sense of sight has been both treasured and revered. Without doubt, visual loss resulting from lens dysfunction has always plagued the human family. In the early years of lens development research, investigations of the eye were intertwined with the genesis of the field of ophthalmology. Two valuable texts, extensively cited in this chapter, provide much more detail on the origins of this medical discipline than we are able to offer here. For those particularly interested in the history of ophthalmology, we recommend The History of Ophthalmology, edited by Daniel Albert and Diane Edwards (1996), as well as Julius Hirschberg’s eleven-volume series The History of Ophthalmology, translated by Frederick C. Blodi. We also highly recommend Howard B. Adelmann’s Marcello Malpighi and the Evolution of Embryology (1966). Adelmann’s text presents a good history of ocular embryology in volume 3 under Excursus XII, “The Eyes.”
For many, the history of research in eye lens development largely dates back to the famous experiments of Hans Spemann and his work on lens induction at the turn of the twentieth century. However, descriptive knowledge of all the basic ocular structures was well established by the time Spemann began his experiments. Spemann’s fundamental experiments on lens induction, along with those of Mencl and Lewis, are reviewed in subsequent chapters (see, e.g., chap. 2). One of the aims of the present chapter is to review the major recorded advances in the understanding of the anatomy, pathology, and development of the ocular lens from antiquity up to 1900.
The ancient Egyptians may have been the first to document cases of cataract, as this was likely the disease state referred to under the descriptions ‘darkening of the pupil’ and ‘white
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