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Regeneration

Regeneration
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Title: Regeneration
Release Date: 2018-05-22
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REGENERATION

Columbia University Biological Series.
EDITED BY
HENRY FAIRFIELD OSBORN
AND
EDMUND B. WILSON.

1. FROM THE GREEKS TO DARWIN.
By Henry Fairfield Osborn, Sc.D. Princeton.

2. AMPHIOXUS AND THE ANCESTRY OF THE VERTEBRATES.
By Arthur Willey, B.Sc. London Univ.

3. FISHES, LIVING AND FOSSIL. An Introductory Study.
By Bashford Dean, Ph.D. Columbia.

4. THE CELL IN DEVELOPMENT AND INHERITANCE.
By Edmund B. Wilson, Ph.D. J.H.U.

5. THE FOUNDATIONS OF ZOOLOGY.
By William Keith Brooks, Ph.D. Harv., LL.D. Williams.

6. THE PROTOZOA.
By Gary N. Calkins, Ph.D. Columbia.

7. REGENERATION.
By Thomas Hunt Morgan, Ph.D.

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COLUMBIA UNIVERSITY BIOLOGICAL SERIES. VII.

REGENERATION

BY
THOMAS   HUNT   MORGAN,   Ph.D.
PROFESSOR OF BIOLOGY, BRYN MAWR COLLEGE



New York
THE MACMILLAN COMPANY
LONDON: MACMILLAN & CO., Ltd.
1901
All rights reserved
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Copyright, 1901,
By THE MACMILLAN COMPANY.
Norwood Press
J. S. Cushing & Co.—Berwick & Smith
Norwood, Mass., U.S.A.

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To My Mother

PREFACE

This volume is the outcome of a course of five lectures on “Regenerationand Experimental Embryology,” given in Columbia University in January,1900. The subjects dealt with in the lectures are here more fullytreated and are supplemented by the discussion of a number of relatedtopics. During the last few years the problems connected with theregeneration of organisms have interested a large number of biologists,and much new work has been done in this field; especially in connectionwith the regenerative phenomena of the egg and early embryo. Thedevelopment of isolated cells or blastomeres has, for instance, arousedwidespread interest. It has become clearer, as new discoveries have beenmade, that the latter phenomena are only special cases of the generalphenomena of regeneration in organisms, so that the results have beentreated from this point of view in the present volume.

If it should appear that at times I have gone out of my way to attackthe hypothesis of preformed nuclear germs, and also the theory ofnatural selection as applied to regeneration, I trust that theimportance of the questions involved may be an excuse for the criticism.

If I may be pardoned a further word of personal import, I should like toadd that it has seemed to me that far more essential than each specialquestion with which the biologist has to deal is his attitude toward thegeneral subject of biology as a science. Never before in the history ofbiology has this been more important than at the present time, when weso often fail to realize which problems are really scientific and whichmethods are legitimate for the solution of these problems. The custom ofindulging in exaggerated and{viii} unverifiable speculation bids fair to dullour appreciation for hypotheses whose chief value lies in thepossibility of their verification; but those who have spent their timeand their imagination in such speculations cannot hope for long to holdtheir own against the slow but certain advance of a scientific spirit ofinvestigation of organic phenomena. The historical questions with whichso many problems seem to be connected, and for which there is norigorous experimental test, are perhaps responsible for the loose way inwhich many problems in biology are treated, where fancy too oftensupplies the place of demonstration. If, then, I have tried to use mymaterial in such a way as to turn the evidence against some of theuncritical hypotheses of biology, I trust that the book may have a widerbearing than simply as a treatment of the problems of regeneration.

I wish to acknowledge my many obligations to Professor H. F. Osborn andto Professor E. B. Wilson for friendly criticism and advice; and inconnection with the revision of the text I am greatly indebted toProfessor J. W. Warren, to Professor W. M. Wheeler, to Professor G. H.Parker, and to Professor Leo Loeb.

Bryn Mawr College, Pennsylvania,
June 11, 1901.
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CONTENTS

CHAPTER I
General Introduction
 PAGE
Historical Account of the Work on Regeneration of Trembley, Bonnet, and Spallanzani1
Some Further Examples of Regeneration6
Definition of Terms19
CHAPTER II
The External Factors of Regeneration in Animals
The Effect of Temperature26
The Effect of Food27
The Effect of Light29
The Effect of Gravity30
The Effect of Contact33
The Effect of Chemical Changes in the Environment35
General Conclusions36
CHAPTER III
The Internal Factors of Regeneration in Animals
Polarity and Heteromorphosis38
Lateral Regeneration43
Regeneration from an Oblique Surface44
The Influence of Internal Organs at the Cut-surface52
The Influence of the Amount of New Material54
The Influence of the Old Parts on the New62
The Influence of the Nucleus on Regeneration65{x}
The Closing in of Cut-edges69
CHAPTER IV
Regeneration in Plants
Regeneration in Flowering Plants71
Regeneration in Liverworts, Mosses, and Moulds84
Hypothesis of Formative Stuffs88
CHAPTER V
Regeneration and Liability to Injury
Examples of Supposed Connection between Regeneration and Liability to Injury92
Regeneration in Different Parts of the Body97
Regeneration throughout the Animal Kingdom103
Regeneration and the Theory of Natural Selection108
CHAPTER VI
Regeneration of Internal Organs. Hypertrophy. Atrophy
Regeneration of Liver, Eye, Kidney, Salivary Glands, Bones, Muscles, Nerves, Brain, and Cord of Vertebrates111
Examples of Hypertrophy115
Theories of Hypertrophy118
Atrophy123
Incomplete Regeneration125
CHAPTER VII
Physiological Regeneration
Supposed Relation between Physiological Regeneration and Restorative Regeneration128
Regeneration and Growth131
Double Structures135
CHAPTER VIII
Self-division and Regeneration. Budding and Regeneration. Autotomy. Theories of Autotomy
Review of Groups in which Self-division occurs142{xi}
Division in Plane of Least Resistance144
Review of Groups in which Budding occurs. Relation of Budding to Regeneration149
Autotomy150
Theories of Autotomy155
CHAPTER IX
Grafting and Regeneration
Examples of Grafting in Hydra, Tubularia, Planarians, Earthworms, Tadpoles159
Grafting Pieces of Organs in Other Parts of the Body in Higher Animals178
Grafting of Parts of Embryos of the Frog182
Union of Two Eggs to form One Embryo188
CHAPTER X
The Origin of New Cells and Tissues
Origin of New Cells in Annelids190
Origin of the New Lens in the Eye of Salamanders203
The Part played by the “Germ-layers” in Regeneration207
The Supposed Repetition of Phylogenetic and Ontogenetic Processes in Regeneration212
CHAPTER XI
Regeneration in Egg and Embryo
Introduction216
Regeneration in Egg of Frog217
Regeneration in Egg of Sea-urchin228
Regeneration in Other Forms: Amphioxus, Ascidian, Ctenophore, Snail, Jelly-fish, Fish236
CHAPTER XII
Theories of Development
Theories of Isotropy and of Totipotence of Cells242
Theory of Qualitative Division of Nucleus243
Theory of Equivalency of Cells244
Theory of the Organized Structure of the Protoplasm246{xii}
Theory of Cells as Units250
Further Analysis of Theories of Qualitative Nuclear Divisions and of the Equivalency of Blastomeres252
Driesch’s Analytical Theory, Criticism, and Later Theories of Driesch253
Conclusions256
CHAPTER XIII
Theories of Regeneration
Pre-formation Theory260
Comparison with Growth of Crystal263
Completing Theory264
Theory of Formative Stuffs265
Conclusions269
Theory of Tensions controlling Growth271
CHAPTER XIV
General Considerations and Conclusions
Organization277
Machine Theory of Development and of Regeneration283
Teleology283
“Action at a Distance”284
Definition of Terms: Cause, Stimulus, Factor, Force, Formative Force, Organization287
Regeneration as a Phenomenon of Adaptation288
Literature293
Index311

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REGENERATION

CHAPTER I
GENERAL INTRODUCTION

Although a few cases of regeneration were spoken of by Aristotle and byPliny, the subject first attracted general attention through theremarkable observations and experiments of the Abbé Trembley. Hisinterest was drawn to certain fresh-water polyps, hydras, that were newto him, and in order to find out if the organisms were plants or animalshe tried the effect of cutting them into pieces; for it was generallyknown that pieces of a plant made a new plant, but if an animal were cutinto pieces, the pieces died. Trembley found that the polyp, if cut intwo, produced two polyps. Logically, he should have concluded that thenew form was a plant; but from other observations, as to its method offeeding and of movement, Trembley concluded that the polyp was ananimal, and that the property of developing a new organism from a partmust belong to animals as well as to plants. “I felt,” he says,“strongly that nature is too vast, and too little known, for us todecide without temerity that this or that property is not found in oneor another class of organized bodies.”

Trembley’s first experiments were made in 1740, and the remarkableresults were communicated by letter to several other naturalists. Itcame about in this way that before Trembley’s memoir had appeared, in1744, his results were generally known, and several other observers hadrepeated his experiments, and extended them to other forms, and had evenpublished an account of their own experiments, recognizing Trembley,however, as the first discoverer. Thus Réaumur described, in 1742, anumber of other forms in which regeneration takes place; and Bonnet, in1745, also described some experiments that he had made during the fourpreceding years. Widespread interest was aroused by these results, andmany different kinds of animals were experimented with to test theirpower of regeneration. Most important of these new discoveries werethose of Spallanzani, who published a short preliminary statement of hisresults, in 1768, in his Prodromo.{2}

Trembley found that when a hydra is cut in two, the time required forthe development of the new individuals is less during warm than duringcold weather. He also found that if a hydra is cut into three or fourparts, each part produces a new individual. If these new hydras are feduntil they grow to full size, and are then again cut into pieces, eachpiece will produce a new polyp. The new animals were kept in some casesfor two years, and behaved in all respects as do ordinary polyps.

Trembley also found that if the anterior, or head-end, with itstentacles, is cut off, it also will make a new animal. If a hydra is cutlengthwise into two parts, the edges roll in and meet, and in an hour,or less, the characteristic form may be again assumed. New arms mayappear later on the new individual. If a hydra is split lengthwise intofour pieces, each piece will also produce a new polyp.

If the head-end only of a hydra is split in two, each half becomes a newhead, and a two-headed hydra results. If each of the new heads is splitagain, a four-headed hydra is produced; and if

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