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Bfm:978-3-7091-6435-8/1.Pdf Santiago Ramön y Cajal Texture of the Nervous System of Man and the Vertebrates Volume I An annotated and edited translation of the original Spanish text with the additions of the French version by Pedro Pasik and Tauba Pasik Springer-Verlag Wien GmbH Prof. em. Dr. Pedro Pasik Prof. em. Dr. Tauba Pasik Department of Neurology Mount Sinai School of Medicine New York, NY, USA This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically those of translation, reprinting, re-use of illustrations, broadcasting, reproduction by photocopying machines or similar means, and storage in data banks. Product Liability: The publisher can give no guarantee for information about drug dosage and application thereof contained in this book. In every individual case the respective user must check its accuracy by consulting other pharmaceutical literature. The use of registered names, trademarks, etc. in this publication does not imply, even in the absence of specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. © 1999 Springer-Verlag Wien Originally published by Springer-Verlag Wien New York in 1999 Softcover reprint of the hardcover 1st edition 1999 Typesetting: Thomson Press (India) Ltd., New Dehli Cover design: B. Kollmann Printed on acid-free and chlorine-free bleached paper SPIN: 10637744 With 270 (partly coloured) Figures ISBN 978-3-7091-7323-7 ISBN 978-3-7091-6435-8 (eBook) DOI 10.1007/978-3-7091-6435-8 TO THE MEMORY OF JANOS (JOHN) SZENTAGOTHAI 1912-1994 FOREWORD Most scientists agree that the 21st century will be the "Century of Biology". The revolution that started a few decades ago with the extraordinary develop­ ment of molecular biology, has provided the basis for the understanding of the mechanisms that govern life through the regulation of cell function and cell-to­ cell interactions. Neuroscience will surely be one of the biological disciplines that will profit the most from this revolution. Neuroscience comprises the study of the brain, and the more difficult relationship of brain and behavior. It is indeed a very broad field of investigation that, from molecular biology and genetics, passing through systems physiology, functional mapping of the brain, and even human behavior, has as the ultimate goal the understanding of how the brain works. The importance of this aim is immense, since it is the only scientific approach to understanding ourselves. Today, owing to the great progress made in neuroscientific research, there is a real hope that highly complex processes such as sensory perception, ICTlg­ term memory, learning and others will soon be comprehended in their entirety. This progress is timely to help in finding solutions to social and health problems emerging in our society from the growth of urban population density and its cohorts of stress and tensions, the increase in drug-addiction, the prolongation of longevity, etc. It is obvious that a better understanding of brain function, and therefore of ourselves, will improve the quality6f our life. In a sense, neuroscience can be viewed as ushering in a new humanism, ready to challenge the nature of human beings and their responsibilities. From Neuro­ science shall emerge a new vision of the world and of mankind within the world. The present revolution in Neuroscience has its roots in the work of Santiago Ramon y Cajal, who during almost half a century (1887-1934) of patient work was able to show that the nervous system is made up of billions of indepen­ dent, richly and precisely interconnected nerve cells. His studies on the architectural organization of the brain, and his prophetical predictions of its functions became the basis of neuroanatomy, neurophysiology, neuropathol­ ogy, and what he named as "rational psychology". This monumental work justifies his well deserved title of founder of modem Neuroscience. Of course, despite the outstanding accomplishments of Cajal, and his genius in unraveling the complexity of brain structure, the postulation and subsequent demonstration of the neuron doctrine, was not solely the result of his own viii Foreword investigation. As it always happens in Science, the contemporary knowledge in the field and the introduction of novel analytical and experimental methods greatly contributed to Cajal's achievements. During the second half of the 19th century, most investigators considered the nervous system as a huge syncitium, within which protoplasmic processes emerging from cell bodies, after breaking up into thinner branches, dissolved into a tight network that terminated by reconstituting the nerve fibers of the white matter. Investigators such as Wilhelm His in Germany and August Forel in Switzerland were the first to fight successfully the network, or reticularist, theory. W. His (1886), analyzing the early stage of nervous system development, was able to determine that embryonicaxons are a continuation of the first process emerging from postmitotic nerve cells, and that they grow from the free distal end of this process. Forel (1887) based his conclusions of Gudden's type of experiments by showing that, after axonal avulsion of motor cranial nerve roots, only those cells at the origin of the avulsed axons become atrophic. These observations gave indirect proof of the reciprocal trophic dependency between axons and their cell bodies. There is no doubt, however, and despite some recent unfounded attacks, that Cajal was crucial in solving the controversy about the way nerve cells communicate, and their processes terminate. Cajal's technical skills in modifying Camillo Golgi's silver stain (reazione nera), and in developing new metallic impregnation methods, as well as the successful use of the embryonic and degeneration approaches of his predecessors, provided the necessary tools for his discoveries. It is interest to recall that neuromorphologists are currently passing through a somewhat similar innovative period to that experienced by Cajal after Golgi's technical breakthrough in 1873. Thus, the longed-for wish to correlate the chemical constituents of the neuron with their structural counterparts, as a valuable approach to better appraise neuronal function and organization, is now a reality. In the last quarter of this century, great technological advances emerging from physics, chemistry and molecular biology, have revolutionized the field and breathed new life into the morphological approach. The main advances in histochemistry, which have made neuromorphology one of the most rapidly expanding fields in Neuroscience, can be summarized as follows: Histofiuorescent methods for studying monoaminergic systems, which opened up the new field of analysis of neuronal systems with identified neurotransmitters. Autoradiography and the diversity of its application, from tracing connections (axoplasmic transport), identifying neurotransmitters in neu­ rons (specific reuptake mechanisms), quantitative analysis of receptor­ binding and metabolic activity of neuronal networks, to in situ hybridization for the detection of messenger RNAs in order to study gene expression and regulation. Foreword ix - Immunocytochemistry with its many uses, such as antibodies to study the morphological distribution of small (i.e. amino acid haptens) and large molecules. Among many important discoveries, it is noteworthy that this method was the basis for the concept of the co-localization of multiple neurotransmitters in the same neuron and axon terminal. Finally, advances in physical science and informatics have been extremely useful in generating new tools to study the organization, chemical composition and dynamics of the neuron. Hence, the development of computer-assisted image-analysis has greatly promoted quantitative studies that were impossible only a few years ago. For instance, the quantification of receptors distribution, and of metabolic activity of neurons in autoradiograms; three-dimensional morphometry of neuronal bodies and their dendrites in Golgi-stained or, better yet, in intracellularly injected neurons; analysis of somatodendritic gradients of neurotransmitter receptor-protein distribution, or synaptic density in immunofluorescent preparations analyzed with confocal microscopy, etc. In addition, new tools (video-enhanced contrast microscopy, and optical recording of neuronal activity, positron emission tomography (PET), and the newly generated functional magnetic resonance imaging (MRI) techniques have added a new temporal dimension to the morphological approach. Indeed, these non-invasive imaging techniques allow the investigator to examined the same neuronal populations over extended periods of time, and even to repeat the observations weeks, months or years later. In my view, although research and technical advances contemporary with Cajal were instrumental in carrying out his work, it was his analytical power and brilliant interpretations the sole responsible for his discoveries. We are in his debt for opening up this new era in the history of Neuroscience. The best evidence for my enthusiastic assertion is the validity of Cajal's opus magnum, the "Textura del Sistema Nervioso del Hombre y de los Vertebrados". This book, published in Madrid between 1899 and 1904, and its French translation by L. Azoulay in 1909 and 1911, has kept the attention of most neuroscientists for almost a century. It is still widely cited today, not much because of its historical value but for the accuracy of its
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