Title: A Critique of the Theory of Evolution
Author: Thomas Hunt Morgan
Release date: December 17, 2009 [eBook #30701]
Most recently updated: January 5, 2021
Language: English
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| Transcriber's note: | A few typographical errors have been corrected. They appear in the text like this, and the explanation will appear when the mouse pointer is moved over the marked passage. Figures 41 and 42 have been interchanged from the printed copy in order to match the text. |
was established in 1912 with a bequest of $25,000 under the will of Louis Clark Vanuxem, of the Class of 1879. By direction of the executors of Mr. Vanuxem's estate, the income of the foundation is to be used for a series of public lectures delivered in Princeton annually, at least one half of which shall be on subjects of current scientific interest. The lectures are to be published and distributed among schools and libraries generally.
The following lectures have already been published or are in press:
1912-13 The Theory of Permutable Functions, by Vito Volterra
1913-14 Lectures delivered in connection with the dedication of the Graduate College of Princeton University by Emile Boutroux, Alois Riehl, A. D. Godley, and Arthur Shipley
1914-15 Romance, by Sir Walter Raleigh
1915-16 A Critique of the Theory of Evolution, by Thomas Hunt Morgan
OF THE
BY
PROFESSOR OF EXPERIMENTAL ZOOLOGY IN
COLUMBIA UNIVERSITY
LECTURES DELIVERED AT PRINCETON UNIVERSITY
FEBRUARY 24, MARCH 1, 8, 15, 1916
PRINCETON UNIVERSITY PRESS
PRINCETON
LONDON: HUMPHREY MILFORD
OXFORD UNIVERSITY PRESS
1916
Copyright, 1916, by
Princeton University Press
Published October, 1916
Occasionally one hears today the statement that we have come to realize that we know nothing about evolution. This point of view is a healthy reaction to the over-confident belief that we knew everything about evolution. There are even those rash enough to think that in the last few years we have learned more about evolution than we might have hoped to know a few years ago. A critique therefore not only becomes a criticism of the older evidence but an appreciation of the new evidence.
In the first lecture an attempt is made to put a new valuation on the traditional evidence for evolution. In the second lecture the most recent work on heredity is dealt with, for only characters that are inherited can become a part of the evolutionary process. In the third lecture the physical basis of heredity and the composition of the germ plasm stream are examined in the light of new observations; while in the fourth lecture the thesis is developed that chance variation combined with a property of living things to manifold themselves is the key note of modern evolutionary thought.
T. H. Morgan
| CHAPTER I | |||
| A REVALUATION OF THE EVIDENCE ON WHICH THE THEORY OF EVOLUTION WAS BASED | |||
| PAGE | |||
| Preface | v | ||
| 1. | Three Kinds of Evolution | 1-7 | |
| 2. | The Evidence for Organic Evolution | 7-27 | |
| a. | The Evidence from Comparative Anatomy | 7-14 | |
| b. | The Evidence from Embryology | 14-23 | |
| c. | The Evidence from Paleontology | 24-27 | |
| 3. | The Four Great Historical Speculations | 27-39 | |
| a. | The Environment | 27-31 | |
| Geoffroy St. Hilaire | |||
| b. | Use and Disuse | 31-34 | |
| From Lamarck to Weismann | |||
| c. | The Unfolding Principle | 34-36 | |
| Nägeli and Bateson | |||
| d. | Natural Selection | 36-39 | |
| Darwin | |||
| CHAPTER II | |||
| THE BEARING OF MENDEL'S DISCOVERY ON THE ORIGIN OF HEREDITY CHARACTERS | |||
| 1. | Mendel's First Discovery—Segregation | 41-52 | |
| 2. | Mendel's Second Discovery—Independent Assortment | 52-59 | |
| 3. | The Characters of Wild Animals and Plants Follow the Same Laws of Inheritance as do the Characters of Domesticated Animals and Plants | 59-84 | |
| a. | Sexual Dimorphism | 61-64 | |
| Eosin eye color of Drosophila | 61-62 | ||
| Color of the Clover Butterfly, Colias philodice | 62-63 | ||
| Color of Papilio turnus | 63 | ||
| Color pattern of Papilio polytes | 63-64 | ||
| b. | Duplication of parts | 65-66 | |
| Thorax of Drosophila | 65 | ||
| Legs of Drosophila | 65-66 | ||
| c. | Loss of characters | 66-68 | |
| "Eyeless" of Drosophila | 66-67 | ||
| Vestigial wings of Drosophila | 67 | ||
| Bar eye of Drosophila | 67-68 | ||
| d. | Small changes of characters | 68-70 | |
| "Speck" | 68 | ||
| Bristles of "club" | 70 | ||
| e. | Manifold effects of same factor | 71 | |
| f. | Constant but trivial effects may be the product of factors having other vital aspects | 73 | |
| g. | Sex-linked inheritance | 75-80 | |
| in Drosophila ampelophila | 75-76 | ||
| in the wild species D. repleta | 76 | ||
| in man | 77 | ||
| in domesticated Fowls | 77-78 | ||
| in the wild moth, Abraxas | 78-80 | ||
| h. | Multiple allelomorphs | 81-84 | |
| in the wild Grouse Locust | 81-83 | ||
| in domesticated mice and rabbits | 83 | ||
| in Drosophila ampelophila | 84 | ||
| 4. | Mutation and Evolution | 84-88 | |
| CHAPTER III | |||
| THE FACTORIAL THEORY OF HEREDITY AND THE COMPOSITION OF THE GERM PLASM | |||
| 1. | The Cellular Basis of Organic Evolution and Heredity | 89-98 | |
| 2. | The Mechanism of Mendelian Heredity Discovered in the Behavior of the Chromosomes | 98-102 | |
| 3. | The Four Great Linkage Groups of Drosophila Ampelophila | 103-118 | |
| a. | Group I. | 104-109 | |
| b. | Group II. | 109-112 | |
| c. | Group III. | 112-115 | |
| d. | Group IV. | 115-118 | |
| 4. | Localization of Factors in the Chromosomes | 118-142 | |
| a. | The Evidence from Sex Linked Inheritance | 118-137 | |
| b. | The Evidence from Interference | 137-138 | |
| c. | The Evidence from Non-Disjunction | 139-142 | |
| 5. | How Many Genetic Factors are there in the Germ-plasm of a Single Individual? | 142-143 | |
| 6. | Conclusions | 144 | |
| CHAPTER IV | |||
| SELECTION AND EVOLUTION | |||
| 1. | The Theory of Natural Selection | 145-161 | |
| 2. | How has Selection in Domesticated Animals and Plants brought about its Results? | 161-165 | |
| 3. | Are Factors Changed through Selection? | 165-187 | |
| 4. | How does Natural Selection Influence the course of Evolution? | 187-193 | |
| 5. | Conclusions | 193-194 | |
| Index | 195-197 | ||
A REVALUATION OF THE EVIDENCE ON WHICH THE THEORY OF EVOLUTION WAS BASED
We use the word evolution in many ways—to include many different kinds of changes. There is hardly any other scientific term that is used so carelessly—to imply so much, to mean so little.
Three Kinds of Evolution
We speak of the evolution of the stars, of the evolution of the horse, of the evolution of the steam engine, as though they were all part of the same process. What have they in common? Only this, that each concerns itself with the history of something. When the astronomer thinks of the evolution of the earth, the moon, the sun and the stars, he has a picture of diffuse matter that has slowly condensed. With condensation came heat; with heat, action and reaction within the mass until the chemical substances that we know today were produced. This is the nebular hypothesis of the astronomer. The astronomer explains, or tries to explain, how this evolution took place, by an appeal to the physical processes that have been worked out in the laboratory, processes which he thinks have existed through all the eons during which this evolution was going on and which were its immediate causes.
When the biologist thinks of the evolution of animals and plants, a different picture presents itself. He thinks of series of animals that have lived in the past, whose bones (fig. 1) and shells have been preserved in the rocks. He thinks of these animals as having in the past given birth, through an unbroken succession of individuals, to the living inhabitants of the earth today. He thinks that the old, simpler types of the past have in part changed over into the more complex forms of today.
He is thinking as the historian thinks, but he sometimes gets confused and thinks that he is explaining evolution when he is only describing it.
Fig. 1. A series of skulls and feet. Eohippus, Mesohippus, Meryhippus, Hipparion and Equus. (American Museum of Natural History. After Matthews.)
A third kind of evolution is one for which man himself is responsible, in the sense that he has brought it about, often with a definite end in view.
His mind has worked slowly from stage to stage. We can often trace the history of the stages through which his psychic processes have passed. The evolution of the steam-boat, the steam engine, paintings, clothing, instruments of agriculture, of manufacture, or of warfare (fig. 2) illustrates the history of human progress. There is an obvious and striking similarity between the evolution of man's inventions and the evolution of the shells of molluscs and of the bones of mammals, yet in neither case does a knowledge of the order in which these things arose explain them. If we appeal to the psychologist he will probably tell us that human inventions are either the result of happy accidents, that have led to an unforeseen, but discovered use; or else the use of the invention was foreseen. It is to the latter process more especially that the idea of purpose is applied. When we come to review the four great lines of evolutionary thought we shall see that this human idea of purpose recurs in many forms, suggesting that man has often tried to explain how organic evolution has taken place by an appeal to the method which he believes he makes use of himself in the inorganic world.
Fig. 2. Evolution of pole arms. (Metropolitan Museum. After Dean.)
What has the evolution of the stars, of the horse and of human inventions in common? Only this, that in each case from a simple beginning through a series of changes something more complex, or at least different, has come into being. To lump all these kinds of changes into one and call them evolution is no more than asserting that you believe in consecutive series of events (which is history) causally connected (which is science); that is, that you believe in history and that you believe in science. But let us not forget that we may have complete faith in both without thereby offering any explanation of either. It is the business of science to find out specifically what kinds of events were involved when the stars evolved in the sky, when the horse evolved on the earth, and the steam engine was evolved from the mind of man.
Is it not rather an empty generalization to say that any kind of change is a process of evolution? At most it means little more than that you want to intimate that miraculous intervention is not necessary to account for such kinds of histories.
We are concerned here more particularly with the biologists' ideas of evolution. My intention is to review the evidence on which the old theory rested its case, in the light of some of the newer evidence of recent years.
Four great branches of study have furnished the evidence of organic evolution. They are:
Comparative anatomy.
Embryology.
Paleontology.
Experimental Breeding or Genetics.
The Evidence from Comparative Anatomy
When we study animals and plants we find that they can be arranged in groups according to their resemblances. This is the basis of comparative anatomy, which is only an accurate study of facts that are superficially obvious to everyone.
The groups are based not on a single difference, but on a very large number of resemblances. Let us take for example the group of vertebrates.
Fig. 3. Limb skeletons of extinct and living animals, showing the homologous bones: 1, salamander; 2, frog; 3, turtle; 4, Aetosaurus; 5, Pleisiosaurus; 6, Ichthyosaurus; 7, Mesosaurus; 8, duck. (After Jordan and Kellogg.)
The hand and the arm of man are similar to the hand and arm of the ape. We find the same plan in the forefoot of the rat, the elephant, the horse and the opossum. We can identify the same parts in the forefoot of the lizard, the frog (fig. 3), and even, though less certainly, in the pectoral fins of fishes. Comparison does not end here. We find similarities in the skull and back bones of these same animals; in the brain; in the digestive system; in the heart and blood vessels; in the muscles.
Each of these systems is very complex, but the same general arrangement is found in all. Anyone familiar with the evidence will, I think, probably reach the conclusion either that these animals have been created on some preconceived plan, or else that they have some other bond that unites them; for we find it difficult to believe that such complex, yet similar things could have arisen independently. But we try to convince our students of the truth of the theory of evolution not so much by calling their attention to this relation as by tracing each organ from a simple to a complex structure.
I have never known such a course to fail in its intention. In fact, I know that the student often becomes so thoroughly convinced that he resents any such attempt as that which I am about to make to point out that the evidence for his conviction is not above criticism.
Fig. 4. Drosophila ampelophila. a, Female and b, male.
Because we can often arrange the series of structures in a line extending from the very simple to the more complex, we are apt to become unduly impressed by this fact and conclude that if we found the complete series we should find all the intermediate steps and that they have arisen in the order of their complexity. This conclusion is not necessarily correct. Let me give some examples that have come under my own observation. We have bred for five years the wild fruit fly Drosophila ampelophila (fig. 4) and we have found over a hundred and twenty-five new types that breed true. Each has arisen independently and suddenly. Every part of the body has been affected by one or another of these mutations. For instance many different kinds of changes have taken place in the wings and several of these involve the size of the wings. If we arrange the latter arbitrarily in the order of their size there will be an almost complete series beginning with the normal wings and ending with those of apterous flies. Several of these types are represented in figure 5. The order in which these mutations occurred bears no relation to their size; each originated independently from the wild type.