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The Brain in Hominid Evolution v\useum o/ ,\. %,'*/ * \ 1869 THE LIBRARY j? THE BRAIN IN HOMINID P EVOLUTION PHILLIP V. TOBIAS The brain in hominid evolution g COLUMBIA UNIVERSITY PRESS Q NEW YORK & LONDON i 9 yi Phillip V. Tobias is Professor of Anatomy at the University of the Witwatersrand, Johannesburg. This book is based on the author's James Arthur Lecture, delivered at the American Museum of Natural History, New York City, April 30, 1969; it was the thirty-eighth in this series of lectures. Copyright © 1971 Columbia University Press International Standard Book Number: 0-231-03518-7 Library of Congress Catalog Card Number: 78-158458 Printed in the United States of America LIBRARY OFTHE AMERICAN MUSEUM JAMES ARTHUR LECTURES ON THE EVOLUTION OF THE HUMAN BRAIN March 15, 1932 Frederick Tilney, The Brain in Relation to Behavior April 6, 1933 C. Judson Herrick, Brains as Instruments of Biological Values April 24, 1934 D. M. S. Watson, The Story of Fossil Brains from Fish to Man April 25, 1935 C. U. Ariens Kappers, Structural Principles in the Nervous System; The Development of the Forebrain in Animals and Prehistoric Human Races May 15, 1936 Samuel T. Orton, The Language Area of the Human Brain and Some of Its Disorders Apnl 15 R. W. Gerard, Dynamic Neural Patterns Franz Weidenreich, The Phylogenetic Development of the Hominid Brain and Its Connection with the Transformation of the Skull G. Kingsley Noble, The Neural Basis of Social Behavior of Vertebrates John F. Fulton, A Functional Approach to the Evolution of the Primate Brain Frank A. Beach, Central Nervous Mechanisms Involved in the Reproductive Behavior of Vertebrates George Pinkley, A History of the Human Brain James W. Papez, Ancient Landmarks of the Human Brain and Their Origin James Howard McGregor, The Brain of Primates K. S. Lashley, Neural Correlates of Intellect Warren S. McCulloch, Finality and Form in Nervous Activity S. R. Detwiler, Structure-Function Correlations in the Develop- ing Nervous System as Studied by Experimental Methods Tilly Edinger, The Evolution of the Brain Donald O. Hebb, Evolution of Thought and Emotion Ward Campbell Halstead, Brain and Intelligence Harry F. Harlow, The Brain and Learned Behavior Clinton N. Woolsey, Sensory and Motor Systems of the Cere- bral Cortex May 81, 1953 Alfred S. Romer, Brain Evolution in the Light of Vertebrate History May 5, 1954 Horace W. Magoun, Regulatory Functions of the Brain Stem April si, 1955 *Fred A. Mettler, Culture and the Structural Evolution of the Neural System v5£ igr,6 Harman, Paleoneurologic, Neoncurologic, April 26, Pinckney J. and Ontogenetic Aspects of Brain Phylogeny April 25, ic)57 'Davenport Hooker, Evidence of Prenatal Function of the Central Nervous System in Man May 8, 1958 *David P. C. Lloyd, The Discrete and the Diffuse in Nervous Action May 6, 1959 'Charles R. Noback, The Heritage of the Human Brain May 26, i960 *Ernst Scharrer, Brain Function and the Evolution of Cerebral Vascularization May 16, 1961 Paul I. Yakovlev, Brain, Body and Behavior, Stercodynamic Or- ganization of the Brain and of the Motility-Experience in Man Envisaged as a Biological Action System May 29, 1962 H. K. Hartline, Principles of Neural Interaction in the Retina May 28, 1963 Harry Grundfest, Specialization and Evolution of Bioelectric Activity June 3, 1964 *Roger W. Sperry, Problems Outstanding in the Evolution of Brain Function ' May 6, 1965 *Jos£ M. R. Delgado, Evolution of Physical Control of the Brain May 10, 1966 'Seymour S. Kety, Adaptive Functions and the Biochemistry of the Brain May 25, 1967 Dominick P. Purpura, Ontogenesis of Neuronal Organizations in the Mammalian Brain April 2, 1968 'Kenneth D. Roeder, Three Views of the Nervous System April 2, 1969 fPhillip V. Tobias, The Brain in Hominid Evolution / April 23, 1970 *Karl H. Pribram, What Makes Man Human * Published versions of these lectures can be obtained from The American Museum of Natural History, Central Park West at 79th St., New York, N. Y. 10024. f Published for The American Museum of Natural History by Columbia University Press. 5* vi U FOREWORD J^ It can scarcely be denied that the development of the brain is a central, even a determining, aspect of human evolution. For, as we learn more about this process, it becomes increasingly evident that the changes leading to Homo sapiens are intimately linked in a kind of feedback situation with the brain. Even if we knew nothing else about it, its threefold increase in size during hominid evolution would merit, if not demand, a special interest. And yet this is one of the least known of the adaptive changes that have occurred in human evolution. One obvious reason, but not the only one, for this is that the brain does not fossilize and thereby provides us, as do the bony and dental structures, with an anatomical record that would enable us to trace in some detail its development. All that we do have in the way of fossil evidence is the inner surface molding and the capacity of the braincase. And the latter can be inferred by indirect methods only for the many cases where the fossil specimen is fragmentary. But because our evidence is so limited, what we do have, or can reason- ably be inferred, takes on an enhanced importance, and it is essential that it be properly and meticulously assembled and appraised. For much can be learned and certain deductions of great importance may be drawn from such data. This necessary task has been admirably carried out by Professor Tobias in his James Arthur Lecture, now expanded in the following book. His broad experience with hominid fossil material has richly equipped him for this task. We are all deeply indebted to him for this valuable survey, long overdue, which I predict will be a standard reference for a long time. Harry L. Shapiro vii j£ U PREFACE J^ This little volume grew out of the thirty-eighth James Arthur Lecture on the Evolution of the Human Brain, which I delivered at the American Museum of Natural History, New York City, on 30 April 1969. There are many ways to approach brain evolution, including the electrophysiological, comparative, ontogenetic, and neurochemical. My choice of a direction, conditioned by my own background as an anatomist and palaeoanthro- pologist, fell on "Some Aspects of the Fossil Evidence on the Evolution of the Hominid Brain." This choice confined me to two phases: the gross external morphology of the brain (as reflected by the modeling of, and markings on, the interior of the braincase and by endocranial casts, natural and artificial) and the tangible evidence of what hominids could do with their brains, in the form of artefacts and other cultural objects. The study of gross external morphology was further circumscribed: it was confined to the measurable. Of all the quantifiable aspects of the brain, its overall size is perhaps the parameter that can be determined most readily and most objectively. Moreover, brain size is highly relevant for such a study as it shows evidence of dramatic change during hominid evolution. Of course, in fossil studies we are unable to determine brain size as such; we are able to measure only the size of the braincase, the endocranial capacity (often loosely called the cranial capacity). Although much tissue and fluid intervene between the brain and the bony walls of the cranial vault, there is obviously a relationship between the size and shape of the two, variable though it may be. Following normal development, big brains lie in big cranial vaults and small brains in small vaults. In selecting size as my yardstick of change in gross encephalic mor- phology, I was, of course, aware of the work of numerous investigators who have addressed themselves to other external morphological features, (e.g. Le Gros Clark, Cooper, and Zuckerman 1936; Edinger 1948; Connolly 1950; Simon 1965; Bauchot and Stephan 1967). Such features have included the lobes and areas of the cerebrum, the positions of certain key fissures, the detailed pattern of convolutions and sulci—and what has been rather inelegantly described by some as "the degree of sulcification" or "the degree of fissuration." I was aware, too, of the considerable subjective element that has often entered into such studies, especially when applied to inter- racial comparisons; of the fact that it is most difficult, and for some characters virtually impossible, to express such observations in precise metrical form; and of the comment by the distinguished neuroanatomist, Dr. Gerhardt von Bonin, after a lifetime of studies on brains and endocasts: "It should at last be admitted that most of what has been said and written on the sulci of the brain as they have been seen on endocasts is worth very little" (von Bonin 1963, p. 50), a view that is shared by not a few of those mentioned above. These are among the reasons why I have chosen to confine this study to brain size and endocranial capacity. It started, as I have said, as a single lecture and, as such, possesses a single unifying theme. Briefly stated, it is this: nothing is more striking and more sustained in the whole of human evolution than the twofold trend towards increase in brain size on the one hand, and, on the other, towards cultural activities, cultural mastery, and, indeed, utter dependence on cul- ture for survival. These two sets of changes are indissolubly linked. The chain between them may be set forth simply as follows: increase in brain size ?± gain in intricacy of neuronal organization ^± rise in complexity of nervous function ^± ever more diversified and complicated behavior re- sponses ?± progressively amplified and enhanced cultural manifestations. This essay explores the first and last steps of this causal chain—the steps that are directly manifest in the fossil record.
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