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HANDBOOK OF NUTRITION, DIET, AND THE EYE Edited by V r. P ictor reedy King’s College London, London, UK AMSTERDAM • BOSTON • HEIDELBERG • LONDON NEW YORK • OXFORD • PARIS • SAN DIEGO SAN FRANCISCO • SINGAPORE • SYDNEY • TOKYO Academic Press is an imprint of Elsevier Academic Press is an imprint of Elsevier The Boulevard, Langford Lane, Kidlington, Oxford, OX5 1GB, UK 225 Wyman Street, Waltham, MA 02451, USA First published 2014 Copyright © 2014 Elsevier Inc. All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any means, electronic or 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 information about the Publisher’s permissions policies and our arrangement with organizations such as the Copyright Clearance 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 the Publisher (other than as may be noted herein). Notices Knowledge and best practice in this field are constantly changing. As new research and experience broaden our understanding, changes in research methods, professional practices, or medical treatment may become necessary. Practitioners and researchers must always rely on their own experience and knowledge in evaluating and using any information, methods, compounds, or experiments described herein. In using such information or methods they should be mindful of their own safety and the safety of others, including parties for whom they have a professional responsibility. To the fullest extent of the law, neither the Publisher nor the authors, contributors, or editors, assume any 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 ideas contained in the material herein. British Library Cataloguing in Publication Data A catalogue record for this book is available from the British Library Library of Congress Cataloguing in Publication Data A catalogue record for this book is available from the Library of Congress ISBN: 978-0-12-401717-7 For information on all Academic Press publications visit our website at store.elsevier.com Printed and bound in the United States 14 15 16 17 10 9 8 7 6 5 4 3 2 1 Contributors Winsome Abbott-Johnson Princess Alexandra Hospital, Laurie T. Butler School of Psychology and Clinical Language Woolloongabba, Qld, Australia Sciences, University of Reading, Reading, UK Niyazi Acar Eye and Nutrition Research Group, University Aldo Caporossi Policlinico Universitario A. Gemelli, Rome, of Burgundy, Centre des Sciences du Goût et de Italy l’Alimentation, Dijon, France Stefano Caragiuli Azienda Ospedaliera Universitaria Senese, Vaishali Agte Agharkar Research Institute, Pune, India Siena, Italy Daniel Agudelo Department of Chemistry–Biology, Univer- Chloé Cartier Département de psychologie, Université du sity of Québec at Trois-Rivières, Trois-Rivières, Québec, Canada Québec à Montréal, Montréal, Québec, Canada Maria Antonietta Altea Department of Clinical and Experi- Cristina Cartiglia University of Genoa, Genoa, Italy mental Medicine, University of Pisa, Pisa, Italy Chi-Ming Chan School of Medicine, Fu Jen Catholic Univer- R.A. Armstrong Vision Sciences, Aston University, Birming- sity, New Taipei City, Taiwan; Department of Ophthalmol- ham, UK ogy, Cardinal Tien Hospital, New Taipei City, Taiwan Tin Aung Singapore National Eye Centre, Singapore Min-Lee Chang Jean Mayer USDA Human Nutrition Bahri Aydın Istanbul Medeniyet University Medical School, Research Center on Aging at Tufts University, Boston, MA Istanbul, Turkey Bashira A. Charles Center for Research on Genomic and Fereshteh Bahmani Kashan University of Medical Sciences, Global Health, National Human Genome Research Institute, Bethesda, Maryland, USA Kashan, Iran D. Balmer IRO, Institute for Research in Ophthalmology, Emily Y. Chew National Eye Institute, National Institutes of Sion, Switzerland Health, Bethesda, Maryland, USA S. Zahra Bathaie Tarbiat Modares University, Tehran, Iran Ching-Yu Cheng Singapore Eye Research Institute, Singapore; Yong Loo Lin School of Medicine, National University of Lynne Bell School of Psychology and Clinical Language Singapore Sciences, University of Reading, Reading, UK Chung-Jung Chiu Jean Mayer USDA Human Nutrition Tos T.J.M. Berendschot University Eye Clinic Maastricht, Research Center on Aging at Tufts University, Boston, MA Maastricht, The Netherlands Deepika Chopra Government Medical College, Amritsar, Paul S. Bernstein Moran Eye Center, University of Utah Punjab, India School of Medicine, Salt Lake City, UT Patricia Coelho de Velasco Instituto de Biologia, Universi- Brian M. Besch Moran Eye Center, University of Utah School dade Federal Fluminense, Niterói, Brazil of Medicine, Salt Lake City, UT David Coman Department of Metabolic Medicine, The Royal Philippe Bourassa Department of Chemistry–Biology, Children’s Hospital, Brisbane, Queensland, Australia University of Québec at Trois-Rivières, Trois-Rivières, Québec, Canada Nicole Combe ITERG – Equipe Nutrition Métabolisme & Santé, Bordeaux, France R.B. Bozard Department of Cellular Biology and Anatomy, Georgia Health Sciences University, Augusta, GA, USA; Dolores Corella Genetic and Molecular Epidemiology Unit, Department of Ophthalmology, Georgia Health Sciences Department of Preventive Medicine and Public Health, University, Augusta, GA, USA School of Medicine, University of Valencia, Valencia, Spain; CIBER Fisiopatología de la Obesidad y Nutrición, ISCIII, Lionel Bretillon Eye and Nutrition Research Group, Univer- Valencia, Spain sity of Burgundy, Centre des Sciences du Goût et de l’Alimentation, Dijon, France M. Cossenza Program of Neurosciences, Fluminense Federal University, Niterói, Brazil; Departament of Physiology and Alain M. Bron Eye and Nutrition Research Group, Univer- Pharmacology, Biomedical Institute, Fluminense Federal sity of Burgundy, Centre des Sciences du Goût et de University, Niterói, Brazil l’Alimentation, Dijon, France Simonetta Costa Division of Neonatology, Catholic Univer- Benjamin Buaud ITERG – Equipe Nutrition Métabolisme & sity of Rome, Rome, Italy Santé, Bordeaux, France Catherine P. Creuzot-Garcher Eye and Nutrition Research Gabriëlle H.S. Buitendijk Erasmus Medical Center, Rotter- Group, University of Burgundy, Centre des Sciences du dam, The Netherlands Goût et de l’Alimentation, Dijon, France xv xvi CONTRIBUTORS Maria Cristina de Oliveira Izar Cardiology Division, Depart- Jose J. Garcia-Medina Department of Ophthalmology, Uni- ment of Medicine, Federal University of São Paulo, São versity General Hospital Reina Sofía, Murcia, Spain; Depart- Paulo, Brazil ment of Ophthalmology, School of Medicine, University of R.P. Cubbidge Vision Sciences, Aston University, Birming- Murcia, Murcia, Spain ham, UK Pitchairaj Geraldine Department of Animal Science, School Alyssa Cwanger FM Kirby Center for Molecular Ophthal- of Life Sciences, Bharathidasan University, Tiruchirappalli, Tamilnadu, India mology, Scheie Eye Institute, University of Pennsylvania, Philadelphia, Pennsylvania, USA Carmen Giannantonio Division of Neonatology, Catholic University of Rome, Rome, Italy Cécile Delcourt Universite de Bordeaux, Bordeaux, France; Inserm, ISPED, Centre INSERM U897-E pidemiologie- C.R. Gibson Wyle Science, Technology and Engineering, Biostatistique, Bordeaux, France Houston, Texas, USA, and Coastal Eye Associates, Webster, Texas, USA Marie-Noëlle Delyfer Universite de Bordeaux, Bordeaux, France; Inserm, ISPED, Centre INSERM U897-Epidemiologie- Snehal Gite Agharkar Research Institute, Pune, India Biostatistique, Bordeaux, France; Service d’Ophtalmologie, N. Goldenberg-Cohen Pediatric Ophthalmology Unit, CHU de Bordeaux, Bordeaux, France Schneider Children’s Medical Center of Israel, Petach Tikva, I.C.L. Domith Program of Neurosciences, Fluminense Israel; The Krieger Eye Research Laboratory, Felsenstein Federal University, Niterói, Brazil Medical Research Center, Sackler School of Medicine, Tel Aviv University, Tel Aviv, Israel David Dunaief Medical Compass, MD Private Practice, New York, USA Glen Gole Paediatrics and Child Health, University of Queensland, Queensland, Australia Joshua L. Dunaief FM Kirby Center for Molecular Ophthal- mology, Scheie Eye Institute, University of Pennsylvania, Ian R. Gorovoy University of California, San Francisco, CA Philadelphia, Pennsylvania, USA Julia A. Haller Wills Eye Hospital, Philadelphia, PA Rajan Elanchezhian Department of Animal Science, School Lisa Hark Wills Eye Hospital, Philadelphia, PA of Life Sciences, Bharathidasan University, Tiruchirappalli, Rijo Hayashi Department of Ophthalmology, Koshigaya Tamilnadu, India Hospital, Dokkyo University, School of Medicine, Kosh- Andrew W. Eller Retina Service, UPMC Eye Center, Univer- igaya, Saitama, Japan sity of Pittsburgh School of Medicine, and The Eye and Ear Hui He Department of Diagnostic Medicine/Pathobiology, Institute, Pittsburgh, PA Kansas State University, Manhattan, Kansas, USA T.G. Encarnação Program of Neurosciences, Fluminense Tatiana Helfenstein Cardiology Division, Department of Federal University, Niterói, Brazil Medicine, Federal University of São Paulo, São Paulo, Brazil Mesut Erdurmuş Abant Izzet Baysal University Medical Francisco Antonio Helfenstein Fonseca Cardiology Divi- School, Bolu, Turkey sion, Department of Medicine, Federal University of São Evangelina Espósito University Clinic Reina Fabiola, Catho- Paulo, São Paulo, Brazil lic University of Córdoba, Córdoba, Argentina Ken-ichi Hosoya Department of Pharmaceutics, Graduate Asghar Farajzadeh Tarbiat Modares University, Tehran, Iran School of Medicine and Pharmaceutical Sciences, University David T. Field School of Psychology and Clinical Language of Toyama, Toyama, Japan Sciences, University of Reading, Reading, UK Yi-Ling Huang Jean Mayer USDA Human Nutrition Research Silvia C. Finnemann Department of Biological Sciences, Center on Aging at Tufts University, Boston, MA Center for Cancer, Genetic Diseases, and Gene Regulation, Chi-Feng Hung School of Medicine, Fu Jen Catholic Univer- Fordham University, Bronx, New York, USA sity, New Taipei City, Taiwan Steven J. Fliesler Veterans Administration Western New M. Ibberson Vital-IT Group, Swiss Institute of Bioinformat- York Healthcare System; University at Buffalo/State Univer- ics, Lausanne, Switzerland sity of New York; and the SUNY Eye Institute, Buffalo, NY Hiroto Izumi School of Medicine, University of Occupa- Nicolas Froger INSERM, U968, Institut de la Vision, Paris, tional and Environmental Health, Fukuoka, Japan France; Sorbonne Universités, UPMC Univ Paris 06, UMR_S Alberto Izzotti University of Genoa, Genoa, Italy 968 Paris, France; CNRS, UMR 7210, Institut de la Vision, Riikka L. Järvinen Finnsusp Ltd, Lieto, Finland Paris, France Hua Ji Department of Nutritional Sciences, Oklahoma State P.S. Ganapathy Department of Cellular Biology and University, Stillwater, Oklahoma; Institute of Genetics and Anatomy, Georgia Health Sciences University, Augusta, GA, Physiology, Hebei Academy of Agriculture and Forestry Sci- USA; Department of Ophthalmology, Georgia Health Sci- ences, Shijiazhuang, PR China ences University, Augusta, GA, USA Yao Jin Nanjing Medical University Eye Hospital, Nanjing, V. Ganapathy Vision Discovery Institute, School of Medi- Jiangsu Province, China cine, Georgia Health Sciences University, Augusta, GA, USA; Department of Biochemistry and Molecular Biology, Corinne Joffre INRA and University of Bordeaux, Nutrition Georgia Health Sciences University, Augusta, GA, USA and Integrative Neurobiology (NutriNeuro) Bordeaux, France CONTRIBUTORS xvii Choun-Ki Joo The Catholic University of Korea, Seoul, Korea Arumugam R. Muralidharan Department of Animal Science, Sang Hoon Jung Functional Food Center, Korea Institute of School of Life Sciences, Bharathidasan University, Tiruchi- rappalli, Tamilnadu, India Science and Technology (KIST), Gangneung Institute, Gang- neung, Republic of Korea Benjamin P. Nicholson National Eye Institute, National Heikki P. Kallio Food Chemistry and Food Development, Institutes of Health, Bethesda, Maryland, USA Department of Biochemistry, University of Turku, Finland R. Paes-de-Carvalho Program of Neurosciences, Fluminense Paul Kerlin Wesley Medical Centre, Auchenflower, Qld, Federal University, Niterói, Brazil; Department of Neurobi- Australia ology, Institute of Biology, Fluminense Federal University, Niterói, Brazil Eun Chul Kim The Catholic University of Korea, Seoul, Korea J.C. Pastor IOBA: Universidad de Valladolid, Valladolid, Spain Jin Sook Kim Herbal Medicine Research Division, Korea Institute of Oriental Medicine, Daejeon, Republic of Korea Dorota Pawlik Jagiellonian University Medical College, Kraków, Poland Amar U. Kishan UC Davis Eye Center, Sacramento, CA John F. Payne Palmetto Retina Center, LLC, West Columbia, SC Caroline C.W. Klaver Erasmus Medical Center, Rotterdam, The Netherlands Daniel Petrovič Institute of Histology and Embryology, Medical Faculty Ljubljana, University of Ljubljana, Lju- Kimitoshi Kohno School of Medicine, University of Occupa- bljana, Slovenia tional and Environmental Health, Fukuoka, Japan Serge Picaud INSERM, U968, Institut de la Vision, Paris, Jean-François Korobelnik Universite de Bordeaux, Bordeaux, France; Sorbonne Universités, UPMC Univ Paris 06, UMR_S France; Inserm, ISPED, Centre INSERM U897-Epidemiologie- 968 Paris, France; CNRS, UMR 7210, Institut de la Vision, Biostatistique, Bordeaux, France; Service d’Ophtalmologie, Paris, France; Fondation Ophtalmologique Adolphe de CHU de Bordeaux, Bordeaux, France Rothschild, Paris, France Yoshiyuki Kubo Department of Pharmaceutics, Graduate Maria D. Pinazo-Duran Ophthalmology Research Unit School of Medicine and Pharmaceutical Sciences, University ‘Santiago Grisolia’, Dr. Peset University Hospital, Valencia, of Toyama, Toyama, Japan Spain; Department of Surgery, School of Medicine, Univer- Petra S. Larmo Aromtech Ltd, Tornio, Finland sity of Valencia, Valencia, Spain Ryszard Lauterbach Jagiellonian University Medical College, Adela Mariana Pintea University of Agricultural Sciences Kraków, Poland and Veterinary Medicine, Cluj-Napoca, Romania Ling-Jun Li Saw Swee Hock School of Public Health, National Jogchum Plat Department of Human Biology, Maastricht University of Singapore, Singapore University Medical Center, Maastricht, The Netherlands Dingbo Lin Department of Nutritional Sciences, Oklahoma C.C. Portugal Program of Neurosciences, Fluminense Federal State University, Stillwater, Oklahoma University, Niterói, Brazil M.I. Lopez-Galvez Universidad de Valladolid, Valladolid, Ananda S. Prasad Department of Oncology, Wayne State Spain University School of Medicine and Barbara Ann Karmanos Yi Lu Eye and ENT Hospital, Fudan University, Shanghai, Cancer Institute, Detroit, Michigan, USA P.R. China Victor R. Preedy Diabetes and Nutritional Sciences, School Zhi-Quan Lu Liaoning Medical University, Liaoning Prov- of Medicine, King’s College London, London, UK ince, P.R. China Jiang Qin Nanjing Medical University Eye Hospital, Nanjing, F. Manco Lavado Hospital Clinico Valladolid, Valladolid, Jiangsu Province, China Spain R. Roduit IRO, Institute for Research in Ophthalmology, Claudio Marcocci Department of Clinical and Experimental Sion, Switzerland; Department of Ophthalmology, Univer- Medicine, University of Pisa, Pisa, Italy sity of L ausanne, Lausanne, Switzerland Shilpa Mathew UC Davis Eye Center, Sacramento, CA Costantino Romagnoli Division of Neonatology, Catholic Cosimo Mazzotta Azienda Ospedaliera Universitaria Senese, University of Rome, Rome, Italy Siena, Italy Marie-Bénédicte Rougier Service d’Ophtalmologie, CHU de Francesca Menconi Department of Clinical and Experimen- Bordeaux, Bordeaux, France tal Medicine, University of Pisa, Pisa, Italy Dumitriţa Olivia Rugină University of Agricultural Sciences Naoya Miyamoto School of Medicine, University of Occupa- and Veterinary Medicine, Cluj-Napoca, Romania tional and Environmental Health, Fukuoka, Japan Charumathi Sabanayagam Singapore Eye Research Insti- Bobeck S. Modjtahedi UC Davis Eye Center, Sacramento, tute, Singapore; Yong Loo Lin School of Medicine, National CA University of Singapore; Office of Clinical Sciences, Duke- Lawrence S. Morse UC Davis Eye Center, Sacramento, CA NUS Graduate Medical School, Singapore Sarah W. Mount School of Psychology and Clinical Lan- Sergio Claudio Saccà St Martino Hospital, Ophthalmology guage Sciences, University of Reading, Reading, UK Unit, Genoa, Italy xviii CONTRIBUTORS José-Alain Sahel INSERM, U968, Institut de la Vision, Paris, María Fernanda Suárez Department of Clinical Biochemis- France; Sorbonne Universités, UPMC Univ Paris 06, UMR_S try, Faculty of Chemical Science, National University of 968 Paris, France; CNRS, UMR 7210, Institut de la Vision, Córdoba, Córdoba, Argentina Paris, France; Centre Hospitalier National d’Ophtalmologie H.A. Tajmir-Riahi Department of Chemistry–Biology, Uni- des Quinze-Vingts, Paris, France; Institute of Ophthalmol- versity of Québec at Trois-Rivières, Trois-Rivières, Québec, ogy, University College of London, UK; Fondation Ophtal- Canada mologique Adolphe de Rothschild, Paris, France; French Gavin S. Tan Singapore National Eye Centre, Singapore Academy of Sciences, Paris, France Vin Tangpricha Emory University, Atlanta, GA Dave Saint-Amour Département de psychologie, Université du Québec à Montréal, Montréal, Québec, Canada; Départe- Akihiko Tawara School of Medicine, University of Occupa- ment d’ophtalmologie, Université de Montréal, Montréal, tional and Environmental Health, Fukuoka, Japan Québec, Canada; Centre de recherche, Centre hospitalier P. Archana Teresa Institute of Ophthalmology, Joseph Eye universitaire Sainte-Justine, Montréal, Québec, Canada Hospital, Tiruchirappalli, Tamilnadu, India Pedro Sanz-Solana Department of Ophthalmology, Dr. Peset Philip A. Thomas Institute of Ophthalmology, Joseph Eye University Hospital, Valencia, Spain Hospital, Tiruchirappalli, Tamilnadu, India Seang Mei Saw Saw Swee Hock School of Public Health, Mehmet Tosun Abant Izzet Baysal University Medical National University of Singapore, Singapore School, Bolu, Turkey D.F. Schorderet IRO, Institute for Research in Ophthalmol- Julio A. Urrets-Zavalía University Clinic Reina Fabiola, ogy, Sion, Switzerland; Department of Ophthalmology, Uni- Catholic University of Córdoba, Córdoba, Argentina versity of Lausanne, Lausanne, Switzerland; Faculty of Life Preejith P. Vachali Moran Eye Center, University of Utah Sciences, Ecole Polytechnique Fédérale de Lausanne, Laus- School of Medicine, Salt Lake City, UT anne, Switzerland Carole Vaysse ITERG – Equipe Nutrition Métabolisme & Claudio Alberto Serfaty Instituto de Biologia, Universidade Santé, Bordeaux, France Federal Fluminense, Niterói, Brazil Sabrina Viau MacoPharma, Tourcoing, France Horacio M. Serra Department of Clinical Biochemistry, Faculty of Chemical Science, National University of Córdoba, Claire M. Williams School of Psychology and Clinical Lan- Córdoba, Argentina guage Sciences, University of Reading, Reading, UK Saumil Sethna Department of Biological Sciences, Center for Tien Y. Wong Singapore Eye Research Institute, Singapore; Cancer, Genetic Diseases, and Gene Regulation, Fordham Yong Loo Lin School of Medicine, National University of University, Bronx, New York, USA Singapore Jay Siak Singapore National Eye Centre, Singapore Chen Xi Nanjing Medical University Eye Hospital, Nanjing, Jiangsu Province, China Hüseyin Simavlı Bolu Izzet Baysal State Hospital, Bolu, Turkey Ramazan Yağcı Pamukkale University Medical School, S.B. Smith Department of Cellular Biology and Anatomy, Denizli, Turkey Georgia Health Sciences University, Augusta, GA, USA; Department of Ophthalmology, Georgia Health Sciences Jia Yan Liaoning Medical University, Liaoning Province, P.R. University, Augusta, GA, USA; Vision Discovery Institute, China School of Medicine, Georgia Health Sciences University, Baoru Yang Food Chemistry and Food Development, Depart- Augusta, GA, USA ment of Biochemistry, University of Turku, Finland S.M. Smith Biomedical Research and Environmental Sci- Ji Yong Department of Pathophysiology, Nanjing Medical ences Division (MC SK3), NASA Johnson Space Center, University, Nanjing, Jiangsu Province, China Houston, Texas, USA Vicente Zanon-Moreno Genetic and Molecular Epidemiology R. Socodato Program of Neurosciences, Fluminense Federal Unit, Department of Preventive Medicine and Public Health, University, Niterói, Brazil School of Medicine, University of Valencia, Valencia, Spain; Marco Spinazzi Laboratory for the Research of Neurodegen- CIBER Fisiopatología de la Obesidad y Nutrición, ISCIII, erative Diseases, Department of Human Genetics, University Valencia, Spain of Leuven, Belgium Xiangjia Zhu Eye and ENT Hospital, Fudan University, K. Srinivasan Department of Biochemistry and Nutrition, CSIR Shanghai, P.R. China – Central Food Technological Research Institute, Mysore, India S.R. Zwart Division of Space Life Sciences, Universities Philip Storey Wills Eye Hospital, Philadelphia, PA Space Research Association, Houston, Texas, USA Preface The eye is perhaps one of the least studied organs in aging, metabolic syndrome, obesity and liver disease, diet and nutrition, yet the consequences of vision loss iron overload, hypoglycemia, space flight, and other can be devastating. Partial or whole, vision loss can pathophysiologic conditions. The book also covers many affect not only the quality of life of the individual but micronutrients including trace elements, selenium, the family unit as well. One of the biggest contributors zinc, vitamins A, B2, B7 (biotin), B12, C, and E, folate, to complete vision loss in the western hemisphere is carotenoids, and lutein. There is also material on dietary diabetes, precipitated by metabolic syndrome. In some carbohydrates, glycemic index, lipids (including polyun- developing countries, micronutrient deficiencies are saturated fatty acids, fish oils and h yperlipidemia, and major contributing factors to impaired vision. However, cholesterol deficiency), antioxidant status, fruit and veg- there is a range of ocular defects that have their origin in etables, other natural products, quercetin, resveratrol, nutritional deficiencies or excess, or have been shown acetyl-l-carnitine, taurine, green tea extract, lycopene, to respond favorably to nutritional components in the amino acids, antiglycating phytochemicals, flavonoids, experimental (i.e., preclinical) or clinical setting. The polyphenols, plant stanol and sterol esters, wolfberry, eye, from the cornea to the retina, may be affected by sea buckthorn, and Cornus officinalis. Detailed descrip- nutritional components. As these effects may be physi- tions include all areas of molecular biology and genet- ologic or molecular, there are a great many approaches ics, biochemistry, cell biology, pathology, epidemiology, to understanding how dietary and nutritional factors and other fields within the confines and remit of diet and affect the eye. However, to date there has been no com- nutrition. For example, the molecular biology includes prehensive book on the eye and vision in relation to diet material on gene expression, adenosine A2A receptor and nutrition, especially with the new molecular sci- genes, glycoprotein IIIa gene, haptoglobin genotypes, ences. The Handbook of Nutrition, Diet, and the Eye SLC23A2 gene variation, integrins, and the nitric oxide addresses this. It contains 11 sections: system. Of course, the Editor recognizes the difficulty in ascribing chapters to a specific section, as some chapters 1. Introductions and Overviews will be comfortably placed in two or more sections, but 2. Macular Degeneration this is offset by the excellent indexing. 3. Glaucomas Contributors are authors of international and national 4. Cataracts standing, leaders in the field, and trendsetters. Emerg- 5. Other Eye Conditions ing fields of vision science and important discoveries 6. Obesity, Metabolic Syndrome, and Diabetes relating to diet and nutrition are also incorporated in 7. Macronutrients The Handbook of Nutrition, Diet, and the Eye. This 8. Micronutrients represents essential reading for nutritionists, dietitians, 9. Nutraceuticals optometrists, ophthalmologists, opticians, health care 10. Nutrigenomics and Molecular Biology of Eye professionals, research scientists, molecular or cellular Disease biochemists, physicians, general practitioners, public 11. Adverse Effects and Reactions health practitioners, as well as those interested in eye health and vision in general. Coverage includes overviews of eye diseases and vision loss, age-related macular degeneration, cataracts, Professor Victor R. Preedy, glaucoma, diabetic retinopathy, dry eye, development, King’s College London xix C H A P T E R 1 The Eye and Vision: An Overview R.A. Armstrong, R.P. Cubbidge Vision Sciences, Aston University, Birmingham, UK INTRODUCTION layer thus illustrating the weak attachment that exists between inner and outer layers of the developing retina. Vision is the sense that we rely on most to inform us Within the vitreous chamber, the hyaloid blood vessels of the state of the world. For this reason, more is known have developed that nourish the developing vitreous about the scientific basis of vision than any of our other and the crystalline lens. These vessels normally disin- senses.1 The major organ of vision, the eye, is highly tegrate before birth but remnants of them may persist specialized for photoreception. It focuses light from an into childhood. The crystalline lens has developed at object onto the light-sensitive part of the eye, the retina. this stage and the primary lens fibers have now filled the Changes in specialized neurons in the retina result in cavity of the lens vesicle. The cornea develops from the nerve action potentials that are relayed to the brain via surface ectoderm and mesoderm and the anterior cham- the optic nerve. Visual processing by the brain results in ber is formed between the developing cornea and lens. ‘visual perception’, the construction of a sensory image The fused eyelids can be seen at this stage, their skin and that is then consciously appreciated as vision2,3. All other glands developing from the surface ectoderm, while con- structures of the eye are subsidiary to this function, nective tissue and muscle develop from the mesoderm. either by facilitating focusing of light rays or supporting the tissues of the eye. This chapter is an introduction to the different parts of the eye and their various functions THE OCULAR ADNEXA in achieving a visual image. The ocular adnexa refer to the accessory or adjoining parts of the eye and comprise several structures includ- DEVELOPMENT OF THE EYE ing the eyebrows, eyelids, eyelashes, lacrimal gland, and orbit. Many of these structures are illustrated in the ante- The eyes develop from outgrowths of the brain called rior view of the eye shown in Figure 1.2. the optic vesicles (Fig. 1.1).4 Five weeks after conception, the optic vesicle has emerged from the neural ectoderm Eyebrows of the brain and begins to fold inward producing an inner and an outer layer separated by a cavity. The ret- Each eyebrow is a thickened area of skin with accom- ina and smooth muscle of the iris will develop from this panying hairs that are directed both upward and toward structure. The optic vesicle also induces the formation of the temporal side of the head. The hairs function to pre- the lens placode that develops from an invagination of vent sweat formed on the forehead from entering the surface ectoderm in front of the vesicle and ultimately eyes. In many cultures, the eyebrows are also involved will develop into the crystalline lens. In addition, the in facial expression. hyaloid artery ramifies on the back of the developing lens while the outer surface of the optic vesicle develops Eyelids a network of blood vessels in the mesoderm eventually forming the choroid. Outside this, the mesoderm forms The eyelids are moveable folds of skin that function to the sclera and the extraocular muscles. protect the eyes from particulate matter in the air. They Eight weeks after conception, the thicker inner layer also reduce the amount of light entering the eyes and of the optic cup has detached from the thinner outer provide some of the constituents making up the tears. 3 Handbook of Nutrition, Diet, and the Eye http://dx.doi.org/10.1016/B978-0-12-401717-7.00001-0 © 2014 Elsevier Inc. All rights reserved. 4 1. THE EYE AND VISION: AN OVERVIEW FIGURE 1.2 The anterior parts of the eye. The figure shows various FIGURE 1.1 Development of the eye. The figure shows the lens parts of the anterior eye including the lids, cornea, sclera, and pupil. vesicle, optic cup, and optic stalk, which will develop into the lens, Co: cornea; EB: eyebrow; EL: eyelid; La: lashes; P: pupil; Sc: sclera. retina, and optic nerve, respectively. Lv: lens vesicle, OC: optic cup; OS: Image courtesy RA Armstrong. optic stalk; SE: surface ectoderm. Image courtesy RA Armstrong. that make up the structure of the face such as the max- The ocular surface can usually resist ocular infection illa, frontal, and zygomatic. The purpose of the orbit is to both as a result of the mechanical action of the eyelids, protect the eye and to act as an anchorage point for the which physically remove potential pathogens, and the extraocular muscles and other ocular tissues. washing effect of tears.5 Extraocular Muscles Eyelashes There are six extraocular muscles that are attached to There are two or three rows of eyelashes located on the eye by tendons at the sclera (the white outer coat of the upper edge of the upper and lower lids, the lashes the eye). They function to move the eyes through 360° of numbering approximately 150 on the upper lid and 75 gaze and are coordinated so that the two eyes move in on the lower lid. They function to protect the eye against unison, thus preventing double vision (diplopia). There small particles but are vulnerable to infection, especially are four rectus muscles: medial, lateral, superior, and by bacteria, a condition called ‘blepharitis’. inferior, which are attached to a common tendon ring at their posterior ends (the annulus of Zinn), which in turn, Lacrimal Gland is attached to the posterior surface of the orbit. The pri- mary action of the medial rectus is to pull the eye hori- The lacrimal gland lies inside the eye socket above zontally in the nasal direction, whereas the lateral rectus the eye and functions to produce the tears. It is divided pulls the eye horizontally in the temporal direction. The into a large ‘orbital’ and a smaller ‘palpebral’ portion primary action of the superior rectus is to pull the eye connected by a canal. In the orbital part, ducts join with upward and the inferior rectus to pull the eye down- those from the palpebral portion entering the upper tem- ward. The two remaining muscles, the superior and poral part of the conjunctiva. Excess tears are drained inferior oblique muscles are inserted more ‘obliquely’ via the canaliculi into the lacrimal sac and ultimately into the upper and lower posterior temporal quadrants the nasal cavity. Tears contain lysozymes, lactoferrin, of the orbit. The inferior oblique, and superior, inferior, B-lysin, and immunoglobulin A, which are important in and medial recti muscles are controlled by the third cra- defense against infection. nial nerve (the oculomotor nerve) and the lateral rec- tus by the sixth cranial nerve (the abducens nerve). In addition, the superior oblique muscle is supplied by the Orbit fourth cranial nerve (the trochlear nerve). The primary The orbit is the bony socket that contains the eye. The actions of the superior and inferior oblique muscles are eye is situated in the anterior portion of the orbit closer also to pull the eyes in an upward or downward direc- to its lateral surface than the medial wall and nearer the tion, respectively. Nevertheless, only the primary muscle roof of the orbit than the floor.6 It comprises seven bones, actions have been described, and several of the muscles including the ethmoid and sphenoid bones, and bones act in concert to produce secondary and tertiary actions, 1. INTRODUCTIONS AND OVERVIEWS THE OCUlAR ADNExA 5 is a curved, transparent structure with a radius of 7.8 mm. The anterior surface is continually bathed by tears, while the posterior surface is bathed by aqueous humor. The surface of the cornea, together with the associated tear film, is responsible for most of the refractive power of the eye. Hence, the function of the cornea is to refract light rays so that they eventually come to a focus on the retina. The main thickness of the cornea is composed of regularly arranged collagen fibers, which together with the regular smooth epithelium and lack of blood vessels, is responsible for its transparency. Sclera With the exception of the cornea, the sclera forms the outermost layer of the eye. It is thickest in its posterior region and thinnest at the point of attachment of the ten- FIGURE 1.3 Regions of the eye as seen in a vertical section. The fig- dons at the ends of the extraocular muscles. It comprises ure shows the various structures of the eye as seen in a vertical section. collagen fibers, which, unlike the cornea, are irregularly Ah: aqueous humor; C: conjunctiva; Ch: choroid; CO: cornea; I: iris; L: arranged resulting in an opaque appearance. The sclera lens; O: optic nerve; OD: optic disc; Or: orbit; R: retina; VB: vitreous body. Image courtesy RA Armstrong. is highly fibrous and provides protection, support, and anchorage for structures within and outside the eye such as the musculature, and also maintains the shape of the which can move the eyes in more complex directions. globe. The study of muscle action of the eyes and the coordina- tion of eye movement is called ‘binocular vision’. Iris The Anterior Structures of the Eye The iris is a 12-mm diameter structure that func- tions to regulate the amount of light entering the eye The human eye is approximately spherical in shape, and also separates the eye into anterior and posterior 25 mm in diameter, has a volume of 6.5 mL, while mean chambers. It is analogous in action to the diaphragm axial length of the globe is 24 mm (range 21–26 mm). It of a camera. The pupil is an aperture in the center of actually comprises the parts of two spheres, which are the iris through which light rays pass on route to the represented anteriorly by the cornea, which has a greater retina. The iris also contains muscle that contracts in curvature than that represented by the curvature of the response to bright light, making the pupil smaller and posterior sclera. Hence, the eye can be divided into two reducing the amount of light entering the posterior functionally distinct regions, the anterior eye and the segment. By contrast, dim light will cause the pupil to posterior eye. The anterior segment comprises the cor- dilate thus increasing the light entering the posterior nea, iris, ciliary body, crystalline lens, aqueous humor, segment. The iris is controlled by branches of the auto- and the anterior part of the sclera (Fig. 1.3). nomic nervous system. Hence, parasympathetic stimu- lation, supplied by the oculomotor nerve, will constrict Conjunctiva the pupil while sympathetic stimulation, originating from the superior cervical ganglion, will act to dilate The conjunctiva is the outer membrane of the eye cov- the pupil. ering the white fibrous sclera. It is continuous with that of The posterior surface of the iris is covered in cells the transparent cornea and extends onto the surface of the that contain the pigment melanin and that prevent light upper and lower lids. It is a mucus membrane with a non- from entering the eye through the iris. The remaining keratinized, stratified epithelium and subepithelial layers part of the iris has varying amounts of pigment resulting that are composed of adenoid and connective tissue and is in its characteristic color. Hence, an iris with relatively a region especially vulnerable to infection (‘conjunctivitis’). little pigment appears blue and progressively more pig- ment leads successively to green, hazel, and brown eyes. The amount of pigmentation present and therefore, the Cornea resultant eye color, is genetically determined. Albinism is The cornea is the most anterior structure of the eye and a genetically determined condition that results in a lack comprises one-sixth of the circumference of the globe. It of pigment in cells in the body. In humans, pigmentation 1. INTRODUCTIONS AND OVERVIEWS

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