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doi:10.1093/brain/awr207 Brain 2012: 135; 301–303 | 301 BRAIN A JOURNAL OF NEUROLOGY BOOK REVIEW Golgi: a life in science In 1906, the professors of the Karolinska Institute had a problem; who should get the Nobel Prize in Physiology or Medicine? In the previous 25 years, understanding of the structure of the brain and spinal cord had greatly increased. The nerve cell, or neuron, had been identified as the fundamental unit of the nervous system. The new discoveries had helped to understand the normal functions of the brain and spinal cord and to interpret the causes of neurological disease. The work of two men had led to these advances. One, the Spaniard Santiago Ramon y Cajal (1852–1934) and the other, the Italian Camillo Golgi (1843–1926). Cajal’s great contribution was recognized in his own day, and he continues to be revered. Golgi is acknowledged, but far less well known. Golgi: a biography of the founder of modern neuroscience by Paolo Mazzarello (2009) is explicit in its claims and goes some way to redressing the imbalance. Before Golgi’s work, the structure of individual nerve cells was poorly understood. The available methods for preserving and staining nervous tissue yielded an incomplete picture. The painstaking work of Otto Friedrich Karl Deiters [(1834–63); 1865] had demonstrated the continuity between the nerve cell and its dendrites and axon, but his work involved tedious dissection of hardened tissue, and was most feasible for spinal motor neurons. The many shapes of nerve cells in the brain and spinal cord could not easily be established using his methods. Golgi’s discovery changed all that. Golgi was studying blocks of brain that had been fixed in potassium dichromate and then stained with silver nitrate. In 1873, for the first time, he tentatively announced his discovery of ‘La reazione nera’, the black reaction. It was several years before the profound importance of the Golgi method was recognized. Appreciation was eventually given less to the rather diffident Golgi, than to the more outgoing Ramon y Cajal. In 1887, Cajal visited the Spanish psychiatrist Luis Simarro Lacabra (1851–1921) in Madrid. Simarro had returned to Spain after studying with Jean-Martin Charcot (1825–93) and Louis-Antoine Ranvier (1835–1922) in Paris, and was working with the Golgi method. When Cajal looked at a Golgi preparation in Simarro’s house, he was overwhelmed with the beauty and clarity with which individual nerve cells could be seen. He resolved to study every part of the nervous system with this miraculous new method. In the following years, Cajal described the structure of many of the elements of the retina, brain and spinal cord, culminating in his massive and authoritative Textura del sistema GOLGI: A BIOGRAPHY OF THE FOUNDER OF MODERN NEUROSCIENCE By Paolo Mazzarello 2009. New York: Oxford University Press Price: £45.00/$US69.95 ISBN: 978-0-19-533784-6 nervioso (1899), later translated into French and finally into English. Cajal’s magnificent work and his enthusiastic use of the Golgi stain increased the international recognition of Golgi’s great contribution. However, there was a fundamental problem of interpretation. This book traces the early disagreement that led finally to hostility between Golgi and Cajal. Golgi believed in a reticular view of the structure of the nervous system, whereas Cajal supported the neuron doctrine. According to the reticular theory, there is fusion among neural elements; axons cross and re-cross one another fusing and forming a large neural network. Golgi saw in the widespread bifurcation of axons the basis for a broad reticulum within the brain and spinal cord. In sharp contrast, according to the neuron doctrine, the brain and spinal cord are made of individual elements, called neurons. Neurons may touch, but they do not fuse. The book contrasts the personalities of the two men, perhaps a bit unfairly. Golgi emerges as reserved, and somewhat reticent; Cajal as a shameless promoter of his own work. But it was Cajal’s evidence, not his personality that made the difference. Even hardened reticularists changed sides. Within a few years of Cajal’s use of the Golgi method, nearly all neuroscientists accepted the neuron doctrine. The book makes it easy to understand the situation from Golgi’s point of view. He had invented a totally new method for the study ß The Author (2011). Published by Oxford University Press on behalf of the Guarantors of Brain. All rights reserved. For Permissions, please email: [email protected] 302 | Brain 2012: 135; 301–303 Book Review Figure 1 (A) Golgi cells of the cerebellum from Golgi’s Opera Omnia. (B) Golgi cell Purkinje cell. of nervous system structure and saw in his slides evidence for structural continuity of axons; a reticulum as the main functional unit of the brain and cord. Then, a younger man used his method to champion a radically different view. Although Cajal’s interpretations were to prove correct, it was not until the application of the electron microscope many years later that his views could finally be proven. Nerves touch one another, they do not fuse. Golgi was angered and hurt at the widespread abandonment of the reticular theory and acceptance of the neuron doctrine by most of his fellow scientists. He made defence of the reticular theory the key element in his Nobel Prize speech. Cajal seemed more embarrassed than outraged at the speech. The book highlights two types of scientist. A major contributor to science may invent a new method and spend years observing and describing their observations. However, great scientists go beyond that. Based on their work, they come up with a profound unifying principle to account for a great mass of data, and eventually prove the principle to be correct. In biology, the obvious example of the latter type of scientist is Charles Darwin (1809–82) who spent years systematically observing animals and plants, breeding pigeons and studying molluscs; and followed this by providing a profound insight that reset biology. Cajal had a similar role in neuroanatomy. Heinrich Wilhelm Gottfried von Waldeyer-Hartz (1836–1921) had coined the term ‘neuron’ for the nerve cell and was an ardent supporter of the neuron doctrine. It was Cajal who gave it the anatomical support. Cajal begrudgingly adopted the term neuron, but dismissed any claims that Waldeyer-Hartz may have made to evidence supporting the neuron doctrine. Although his reticular views are largely discredited, the book helps us to remember the great contributions that Golgi did make. One of the most important is his early description of cerebellar cell types, an example being the prominent element of the cerebellar cortex known as the Golgi cell (Fig. 1A). Golgi described for the first time the characteristic Golgi cell of the cerebellar cortex as well as the shape of cells first identified by Jan Evangelista Purkinje (1787–1869) and the characteristic right angle branching of their axons. Some of Golgi’s less successful claims are not discussed in the book (Fig. 1B). For example, in Golgi-stained sections of cerebral cortex and cerebellum, the dendrites of most neurons are covered with spines, which are the major postsynaptic site. Golgi thought that the spines were an artefact, and in his initial publications he drew smooth dendrites (Fig. 2A). Cajal (1896) eventually identified the spines using the entirely different method of methylene blue staining (Fig. 2B). As a scientific biography, the book is excellent. It documents all of Golgi’s contributions to the study of the nervous system and disease. For example, Golgi first described the characteristic receptor embedded in tendons that still bears his name; and he discovered the intracellular network of tubules, the Golgi apparatus. Golgi also served as rector of his own University of Pavia for many years, and defended it against the founding of a potentially competitive university in Milan. Golgi, the man, emerges only occasionally. He had a lifelong marriage to Lina Aletti, the niece of his friend and colleague Giulio Bizzozero (1846–1901). The couple had no children, but they took into their home and raised Carolina, the daughter of Golgi’s brother Giuseppe. A most loyal friend, Golgi remained close to Bizzozero for most of his life. A supportive teacher, he encouraged his pupils and his younger colleagues, among them Adelchi Negri (1876–1912), who discovered the characteristic inclusion bodies in rabies, and Edoardo Perroncito (1847–1936), who saw the characteristic spiral form of developing axon tips. Golgi the man is best seen in the cover photo at the age of 77. Looking up from his desk amid a sea of laboratory flasks and bottles, perhaps about to discover something new. Book Review Brain 2012: 135; 301–303 | 303 Figure 2 (A) Golgi stained pyramidal cells from the cerebral cortex. From Golgi’s Opera Omnia. (B) Cajal (1896). Cerebral cortex. Dendritic spines are clearly visible. Mitchell Glickstein University College London, Wolfson House, London NW1 2HE References Advance Access publication November 3, 2011 Cajal (Ramon y Cajal) S. Las espinas colateralesde las celulas del cerebrotenidas con el azul de metleno. Revistra trimestral micrografica; 1896. Cajal (Ramon y Cajal) S. Textura del Sistema Nervioso. Madrid: Nicolas Moya; 1899. Deiters O. Untersuchungen über Gehirn und Rückenmark des Menschen und der Säugetthiere. Braunschweig: Viewig u. Sohn; 1865. Golgi C. Opera Omnia. Vol. 1–3. Milan: Hoepli; 1903. Funding This work has been supported by a grant from the Volkswagen Foundation (to M. G., P. T. and S. B.).