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Human Hand Function This page intentionally left blank Human Hand Function Lynette A. Jones Susan J. Lederman 2006 Oxford University Press, Inc., publishes works that further Oxford University's objective of excellence in research, scholarship, and education. Oxford New York Auckland Cape Town Dares Salaam Hong Kong Karachi Kuala Lumpur Madrid Melbourne Mexico City Nairobi New Delhi Shanghai Taipei Toronto With offices in Argentina Austria Brazil Chile Czech Republic France Greece Guatemala Hungary Italy Japan Poland Portugal Singapore South Korea Switzerland Thailand Turkey Ukraine Vietnam Copyright © 2006 by Lynette A. Jones and Susan J. Lederman Published by Oxford University Press, Inc. 198 Madison Avenue, New York, New York 10016 www.oup.com Oxford is a registered trademark of Oxford University Press All rights reserved. No part of this publication may be reproduced, stored in a retrievalsystem, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording, or otherwise, without the prior permission of Oxford University Press. Library of Congress Cataloging-in-Publication Data Jones, Lynette A. Human hand function / Lynette A. Jones and Susan J. Lederman. p. cm. Includes bibliographical references and index. ISBN-13 978–0–19–517315–4 ISBN 0–19–517315–5 1. Hand—Physiology. 2. Hand—Anatomy. 3. Hand—Movements. I. Lederman, Susan J. II. Title. [DNLM: 1. Hand—physiology. 2. Hand—anatomy & history. WE 830J775h 2006] QP334.J66 2006 611'.97—dc22 2005018742 987654321 To all those who have worked to unravel the mysteries of the human hand This page intentionally left blank ACKNOWLEDGMENTS The ideas and initialplanning for this book began in September 2001 during Susan Lederman's sabbaticalin the Department of Mechanical Engineering at the Massachusetts Institute of Technology. We then collaboratively wrote the book over the next three years, an experience that proved both rewarding and enjoyable. The work for this book was shared equally, and the order of authorship was decided by the toss of a coin. We wish to thank the following colleagues for providing helpful references and graphic materials, for answering the perplexing questions we posed, or for offering other valuable information: Massimo Bergamasco, Jim Craig, Roz Driscoll, Antonio Frisoli, Yvette Hatwell, Mark Hollins, Peter Hunter, Gunnar Jansson, Charlotte Reed, Hayley Reynolds, and Allan Smith. A number of individuals have provided support and assistance with the book's preparation, including Aneta Abramowicz, Catalina Tobon Gomez, and Lindsay Eng. We would also like to acknowledge the outstanding assistance provided by Cheryl Hamilton and Monica Hurt. We thank Catharine Carlin, our editor at Oxford University Press, who has enthusiastically supported this project throughout and provided us with an unusual flexibility, which made writing this book an enjoyable endeavor. We also wish to thank our anonymous reviewers, who offered thorough, thoughtful, and excellent evaluations of the manuscript. Lynette Jones acknowledges the support of the Massachusetts Institute of Technology and the National Institutes of Health's National Institute of Neurological Disorders and Stroke during the preparation of this book. Susan Lederman acknowledges the support of Queen's University in Kingston, Ontario, Canada; the Canada Council for the Arts, which generously provided a Killam Research Fellowship; the Natural Sciences and Engineering Research Council of Canada; and the Institute for Robotics and Intelligent Systems (Federal Centre of Excellence of Canada). Finally, we would like to acknowledge the support of our families. Lynette Jones thanks her husband, Ian Hunter, and children, Nicholas and Bridget Hunter-Jones, who enthusiastically supported this project, offered encouragement when needed, and were always willing to assist. Your affection and love are truly sustaining. Susan Lederman wishes to thank her partner, Les Monkman, and his children, Lindsay and Fraser. Each of you has made a profound difference in my life. Your continuing love, support, humor, and irony are gifts offered most generously, and I acknowledge them gratefully. This page intentionally left blank CONTENTS 1. HistoricalOverview and GeneralIntroduction 3 2. Evolutionary Development and Anatomy of the Hand 10 3. Neurophysiology of Hand Function 24 4. Tactile Sensing 44 5. Active Haptic Sensing 75 6. Prehension 100 7. Non-prehensile Skilled Movements 116 8. End-effector Constraints 131 9. Hand Function Across the Lifespan 150 10. Applied Aspects of Hand Function 179 11. Summary, Conclusions, and Future Directions 204 References 211 Author Index 249 Subject Index 257 This page intentionally left blank 1 Historical Overview and General Introduction [W]e must confess that it is in the human hand that we have the consummation of all perfection as an instrument. BELL, 1833 The human hand is a miraculous instrument that serves us extremely well in a multitude of ways. We successfully use our hands to identify objects and to extract a wealth of information about them, such as their surface texture, compliance, weight, shape, size, orientation, and thermal properties. We demonstrate impressive manual dexterity when reaching for, grasping, and subsequently manipulating objects within arm's reach. Manualgestures, such as those used in sign language and finger spelling, collectively offer valuable forms of communication to those who are deaf or hearing impaired. In addition, raised tangible graphic displays that are explored by hand provide sensory aids to the blind when planning how to navigate their environment without a sighted companion. Finally, the human hand serves as a formidable creative tool in a variety of aesthetic milieus that include dance, sculpture, playing music, and drawing. Depictions of the human hand in art date back to the Upper Paleolithic era (circa 22,000–28,000 years ago) and can stillbeseen today in caves at Gargas and ElCastilloin the Franco- Cantabrian region (Napier, 1993; Piveteau, 1991). There is even a museum dedicated specifically to the hand (La Musée de la Main), which was established by Claude Verdan, a Swiss hand surgeon, in Lausanne, Switzerland. Historical Roots Not surprisingly, the study of human hand function derives its historical roots from multiple sources. From archaeology come the very detailed analyses of the changes in the anatomy and structure of the hand as it gradually evolved and became capable of using and, more important, manufacturing tools (Marzke & Marzke, 2000). Zoological and anthropological studies have also focused on the structural aspects of the hand, although this work has primarily concentrated on the functional use of the hand by various primate species, including Homo sapiens (Preuschoft & Chivers, 1993). Consideration of these phylogenetic and ontogenetic changes in primate and human hands has led researchers, particularly during the latter half of the 20th century and at the beginning of the 21st century, to examine how such changes relate to the development of human hand function over the lifespan. 4 HistoricalOverview and GeneralIntroduction Philosophers and, more recently, experimental psychologists have been interested in the sense of touch for centuries, although their attention did not focus on manual sensing per se. As early as Aristotle, those interested in touch recognized that this sensory modality was indeed different from the others (i.e., vision, audition, taste, and smell), a perspective that was maintained throughout the 19th century by individuals such as Weber (1834/1978), Blix (1884), Goldscheider (1886), and Frey (1896). Two particularly noteworthy developments of the 19th century that have proved to be of significant value to those who study hand function today derive from the work of Weber and, subsequently, Gustav Fechner. It was Weber (1834/1978) who first introduced quantitative techniques to the study of sensory physiology by measuring both absolute and differential thresholds. He also documented a mathematical relation between the size of the change in stimulus magnitude and the just noticeable difference in perceived intensity, which came to be known as Weber's law. Fechner (1860/1966), both a physicist and a philosopher, was interested in offering an alternate interpretation, monism, to Descartes' mind-body dualism. To do so, he argued that the relation between mental and physical events could be described by a general quantitative “psychophysicallaw ” that assumed that all just noticeable differences are subjectively equal and that Weber's law is true. Toward this end, he developed several classical psychophysical methods for measuring sensory thresholds, all of which remain in common use today. Early tactile concerns centered on issues related to determining the appropriate number of human sensory modalities and submodalities and to the varying sensitivity and spatialacuity of the skin across the entire human body when stimulated by mechanicaland thermalforms of energy. During the 19th century, anatomists whose names are well known today (e.g., Meissner, Pacini, Krause) showed much interest in using the new histological techniques of their day to identify the various receptors within the skin. The results of such studies served as the basis for addressing a third important issue, namely, the links between the receptor populations known at that time and the specific human cutaneous sensibility (or sensibilities) that each served. Although the hand was frequently included as a site of stimulation,