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THE MECHANISTIC CONCEPTION OF LIFE .

ILLINOIS Bgcnts THE BAKER & TAYLOR COMPANY NEW YORK THE CAMBRIDGE UNIVERSITY PRESS LONDON AND EDINBURGH .THE UNIVERSITY OF CHICAGO PRESS CHICAGO.

D. M. PH.THE MECHANISTIC CONCEPTION OF LIFE BIOLOGICAL ESSAYS BY JACQUES LOEB.D.. ILLINOIS . MEMBER OF THE ROCKEFELLER INSTITUTE FOR MEDICAL RESEARCH THE UNIVERSITY OF CHICAGO PRESS CHICAGO. SO..D.

A. . U.Copyright 1912 By The University of Chicago All rights reserved Published July 1912 Composed and Printed By The University of Chicago Press Chicago. Illinois.S.

but in view of the technical difficulties presented by some of the topics this may gave J. Professor Albert Putnam's Sons. repetition work was unavoidable. The Rockefeller Institute FOR Medical Research April 4. Lippincott Company. to The author wishes thank the editors and publishers who Professor their consent to the reprinting of these essays: Cattell. G. J.PREFACE The essays contained in this volume were written on differ- ent occasions mostly in response to requests for a popular presentation of the results of the author's investigations. England.. title of The the volume characterizes their general tendency as an life attempt to analyze point. from a purely physico-chemical view- Since they deal to a large extent with the personal of the author. Philadelphia. of Charles Seward. and the B. serve to facilitate the understanding of the subject. New York and London. Ginn & Co. of McKeen Columbia University. P. of the University of Cambridge. 1912 . of Boston.

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TABLE OF CONTENTS PAGE I. On Some ogy _----------------Facts and Principles of Physiological Morphol- 79 85 113 VI. The Prevention of Fertilization IX. The Role of Salts in the Preservation of Life X. VII. The Mechanistic Conception of Life - _ - - 3 IL The Significance of Tropisms for Psychologj'- - - 35 IIL Some Fundamental Facts and Conceptions concerning the Comparative Physiology of the Central Nervous Sys- tem ----^-----65 IV. Pattern Adaptation of Fishes and the Mechanism of Vision V. On On the Nature of the Process of Fertihzation the Nature of Formative Stimulation (Artificial Par- thenogenesis) - - - -127 155 169 VIII. Experimental Study Animals Index __--__------------------of the Death of the Egg tlii-ough the Act - - - of the Influence of Environment on 195 229 .

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I. THE MECHANISTIC CONCEPTION OF LIFE .

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or we succeed in finding the numerical relation between the conditions of the experiment and the biological result (e. 1911. January. the sum of all life phenomena. scientific basis and ethical life will have to be and our rules of conduct must results of scientific be brought into harmony with the It is seemingly often taken for granted by laymen that is 'Hruth" in biology.. that everything rests on argument or rhetoric and that what is regarded as true today may be expected with some probaIt bility to be considered untrue tomorrow. LIFE^ INTRODUCTORY It is the object of this paper to discuss the question whether our present knowledge gives us any hope that ultimately life.. by coiu-tesy of Professor J.g. but always two forms: it is either possible to control a life phenomenon to such an extent that we can produce it at desire (as.. i. that hypotheses are forwarded. of the same order is as ''truth" in certain of the mental sciences. 3 . reprinted from Popular Science Monthly. happens in sci- ence. September 10. the contraction of an excised muscle). ZSIcKeen CatteU.e. and then abandoned. e. biology is It should. 1912. discussed. be remembered that modern fundamentally an experimental and not a descripof tive science.THE MECHANISTIC CONCEPTION OF I.g. and that its results are not rhetorical. that to say. assume one 1 Address delivered at the First International Congress of jSIonists at Hamburg. especially in the descriptive sciences like paleontology or zoology. in the affirmative our social If on the basis of a serious survey this question can be answered put on a biology. however. or science in general. can be unequivocally explained in physico-chemical terms.

investigators began with primitive means has been completed by more recent investigators Pettenkofer and Voit. must advance. This progress connected pre-eminently with the names of Berzelius and Wilhelm Ostwald.4 The Mechanistic Conception of Life Biology as far as it is Mendel's law of heredity). Zuntz and Atwater. This discrepancy was done away with by the physical chemists.. What these two completely to physico-chemical terms. but II. life a — — — not easily oxidized. based on these two principles cannot retrogress. and proteins are substances which at ordinary temperature are cases are identical. They require the temperature of the flame in order to undergo rapid oxidation through the oxygen This discrepancy between the oxidations in the in the laboratory manifests itself also e. the so-called catalyzers. the beginning of scientific biology with the Scientific biology. who demonstrated is that the same acceleration of brought about by a high temperature can also be accomplished at a low temperature with the chemical reactions which aid of certain specific is substances. namely. These investigations left a gap. living in body and those other chemical processes. digestion or hydrolytic which were at first found to occur outside the living body rapidly only under conditions incompatible with reactions. of the air. defined in this sense. provided that the quantities of carbon dioxide formed in both This was the first attempt to reduce phenomenon. Rubner. life. no matter whether it takes place in the living body or outside. The substances which undergo oxidations in the animal body starch. the formation of animal heat. The oxidation of a food-stuff always furnishes the same amount of heat. begins attempt made by Lavoisier and Laplace (1780) to show that the quantity of heat which is formed in the body of a warm-blooded animal is equal to that formed in a candle. The specific substances . fat.g.

we must either succeed in producing . germs. the 'Griddle of life" By same the ''riddle of life" not everybody will understand the thing. salts. to know how life originates and what death since our ethics must be influenced to a large extent through the answer to this question. desire is. III. The gap in our knowledge which we feel most keenly catalyzers is the fact that the chemical character of the (the enzymes or ferments) is is still unknown. I Biology will certainly retain but believe that we must also follow out the other problem: namely. life. at present that the artificial production of living matter beyond the possibilities of This view does not stand in opposition to the idea of Arrhenius that germs of sufficiently small dimensions are driven by radiation-pressure through space. and that these if they fall upon new cosmic bodies possessing water. Nothing science. however.The Mechanistic Conception of Life 5 which accelerate the oxidations at body temperature sufficiently to allow the maintenance of life are the so-called ferments of oxidation. We all. not the basis. also touches the core of the if for it seems that oxidations form a part. and the proper temperature. We how are not yet able to give an answer to the question life is as to originated on the earth. give rise to a new evolution this idea. however. and we formed also know that not only the compounds which are artificially. of all life phenomena in higher organisms. and oxygen. The work problem of of Lavoisier and Laplace not only marks the it beginning of scientific biology. indicates. of organisms. We know that every living being able to transform food-stuffs into living matter. in the animal body can be produced but that chemical reactions which take place in living organisms can also be repeated at the same rate and temperature in the laboratory.

e. and so on. divides again into eggs. and the egg is now transformed Then an intestine develops one region of the blastula through the growing in of etc. reduced to physico-chemical If The egg it is in the unfertilized condition a single cell with only one nucleus. as to The question shrouded how a spermatozoon can cause an egg to still develop into a new individual was twelve years ago in that of life in general. Every animal originates from an egg and in the majority of animals a new individual can only then in general. in some animals in a few hours. originate. . and later such an egg forms a hollow sphere whose sists of a large number of cells. enters into the egg.g. that is. THE activation OF THE EGG Although we are not yet able to state how life originated more modest problem. how the egg is caused by the sperm to develop into a new individual. the latter begins to develop.. If.. vascular system. enters into the egg. it. . no spermatozoon enters into perishes after a comparatively short time. a spermatozoon i. Embryologists had noticed that occasionally the unfertilized eggs of certain animals. On the outer surface of this hollow sphere cilia are formed into a free-swimming larva. the nucleus begins to divide into two nuclei and the egg which heretofore consisted of one cell is divided into two cell cells.. however. IV. develop if a male sex-cell. or this is why impossible. cells in and gradually the other organs. problem of mystery which today surrounds the origin But today we are able to state that the the activation of the egg is for the most part terms.6 The Mechanistic Conception of Life we must find the reasons living matter artificially. another. a spermatozoon. skeleton. sea-urchins. Subsequently each nucleus and each two. has been solved. in others in a few days or weeks.e. the These cells have. in many tendency shell con- to remain at the surface of the egg or to creep to the surface.

The eggs were put solution the osmotic pressure of which of substituting physico-chemical agencies for the action of the living spermatozoon. 2. 1. These experiments proved the possibility of a salt or sugar. — The same egg immediately surrounded by a larger raised through the addition of a small but definite quantity for two hours into a had been raised to a certain height. Fig. which is formed tlxrough the action of the spermatozoon. but did not yet explain how the in spermatozoon causes the development of the egg. The egg is circle. Fig. 2 tails — Unfertilized egg of the sea-urchin surrounded by spermatozoa. or even birds.The Mechanistic Conception of Life worms. of But the cell divisions in these cases never led to the development of a larva. Mead. since . succeeded twelve years ago in causing the unfertilized eggs of the sea-urchin to develop into treating them with sea-water. show a tendency to a nuclear or even a cell division. The same result was obtained in the eggs of other animals. after the entrance of the spermatozoon. worms. since at the magnification used the were not visible. Only the heads of the spermatozoa are drawn. star-fish. and R. When the eggs were put back into normal sea-water they developed into larvae and a part of these larvae formed an intestine and a skeleton. 1 Fig. and Morgan had succeeded in inducing one or more cell divisions artificially in such eggs. and moUusks. the fertilization membrane. Hertwig. but at the best to the formation an abnormal mass of I cells which soon perished. swimming larvae by the concentration of which was P C7 Fig. This formation of a fertilization membrane can be induced by a purely chemical treatment of the egg.

This membrane formation or rather the modification of the surface of the egg which underlies the membrane formation starts the development.water containing oxygen they all develop. It does not allow it. and that of these the treatment resulting in the formation of the membrane is the more important one. When a spermatozoon enters into the egg it causes primarily a change in the surface of the egg which results of fertilization. Here we have a choice between two methods. all form a fertilization membrane in exactly the same way as if a spermatozoon had entered. or we can put them for about three hours into sea-water deprived of oxygen.8 The Mechanistic Conception of Life upon the these experiments the action of the spermatozoon egg was very incompletely imitated. Strongylocentrotus purpuratus. and in a large number the development is as normal as if a spermatozoon had entered. to proceed very far at room temperature. We can either put the eggs for about one half-hour into a hypertonic solution (which contains free oxygen) . I causing the unfertilized egg to form a injuring the egg. the development to go farther it is In order to allow necessary to submit the eggs after the butyric acid treatment to a second operation. in the formation of the so-called membrane of This phenomenon of membrane method of formation which had always been considered as a phenomenon minor importance did not occur in my original treating the egg with hypertonic sea-water. The essential feature is therefore the fact that the development is caused by two different treatments of the egg. If after that time the eggs are brought back into normal sea-water. If the eggs are then returned to normal sea. however. Six years ago while experimenting on the Californian sea-urchin. for succeeded in finding a method of membrane without This method consists in treating the eggs from one to two minutes with sea-water to which a definite amount of butyric acid (or some other monobasic fatty acid) has been added. This is proved .

3 Fig. 5). 6 respectively. and 5. 4. The changes from Fig. 3. This segmentation occurs after fertilization or after the chemical treatment of the egg described in the text. — Fig. Segmentation of the sea-urchin egg. Fig. — The sea-urchin egg divided into four and eight cells . 6 Figs. 5 Figs.The Mechanistic Conception of Life 9 Fig. 3 to Fig. resulting in the formation of two cells (Fig. 7 and 7. 4 Fig. 5 occur in about one minute or less time.

An analysis of the Fig. arises. cortical layer destroyed. as long as their action to the surface layer of the of the egg cell. as for instance the star-fish. Fig. 9 of the cells cilia of the water. 8 Fig. The intestine begins to form tion of the skeleton appears in the form of fine crystals. larvae. At the surface are formed and the larva begins to swim and reaches the surface — and the first indica- process and of the nature of the agencies which cause it yielded the result that the unfertilized egg possesses a superficial cortical layer. as R. although here. agencies which cause a definite type of cell destruction —the so-called cytolysis is cause also the egg to develop. cortical layer can be destroyed mechanically by shaking the . First larval stage of the sea-urchin egg. the second treatment increases not only the number of found. 8. It is is which must be destroyed before the egg immaterial by what means this superficial All can develop. too.— 10 The Mechanistic Conception of Life fact that in certain forms. limited The butyric acid treatment mentioned above only serves to induce the destrucIn the eggs of some animals this tion of this cortical layer. Lillie The question now how the membrane formation can start the development of the egg. 9. —Blastula. but also improves the appearance of the larvae. artificial by the the causation of the suffice for membrane formation may the development of normal larvae. Fig. Gastrula stage.

therefore. may be Fig. intestine.g. mentioned parenthetically that foreign blood sera have also a cytolytic effect.The Mechanistic Conception of Life egg.^ The mechanism by is which development is caused all apparently the same in these cases. Recently Shearer has succeeded in Plymouth in causing a number of parthenogenetic plutei produced by my method to develop beyond the stage of metamorphosis. and Delage has reported that he raised two larvae of the sea-urchin produced by artificial parthenogenesis to the stage of sexual maturity. This can be caused generally by certain chemical means which play a role also in bacteriology. as A. consequence the develit opment of the sea-urchin egg by treating with the blood of various animals. locentrotus purpuratus. as Bataillon found in his beautiful experiments a year ago. or the rabbit.. it but can also be caused in special cases by mechanical means. P. the de- struction of the cortical layer of the eggs. state that the complete imitation of the developmental spermatozoon by certain physico- chemical agencies has been accomplished. effect of the We may. In the case of the eggs of the frog to pierce the cortical layer with a needle. 11 Mathews found it suffices in the case of star-fish eggs and I in the case of the eggs of certain worms. of cattle. I succeeded in showing that the spermatozoon causes the development of the sea-urchin egg in a way similar to that 1 This method does not work with the eggs of in its applicability as the causation of fish and is apparently as limited development by mechanical agitation. . 10. and I succeeded in membrane formation and e. in causing D Strongy—Pluteus stage Sof skeleton. It such as agitation or piercing of the cortical layer. namely.

while ing two substances into the egg. begins. prove this for the sea-urchin egg foreign sperm. In order to e. e. F. In many cases the spermatozoa which had begun to penetrate into the egg were thrown off again. artificial butyric acid. fish If. They begin If to develop. the action of which corresponds to the treatment of the egg with the hypertonic solution. of cases. but soon show signs of disintegration. He mixed the sperm and eggs of Nereis and centrifuged the mass. that of The sperm of the sea-urchin penetrates so rapidly into the sea-urchin egg that almost always both substances get into the egg. by which the further the egg was prevented. the star-fish. In consequence of the is membrane formation the spermatozoon thrown out.g. Lillie has recently confirmed the of a same fact in the egg worm. in a large number membrane formation before the spermatozoon has found time to penetrate entirely into the egg.12 in The Mechanistic Conception of Life my method of artificial parthenogenesis. must be used. In touching the egg contents the spermatozoon had a chance to give off a substance which liquefied the cortical layer and thereby caused the membrane formation entrance of the spermatozoon into If. one of which acts like the butyric acid and induces the the other acts like the treatment with a hypertonic solution and enables the full development of the larvae. treated with a hypertonic solution they develop into larvae. the star-fish sperm enters completely into the egg before the membrane formation carries also the second substance into the egg.g.. Such eggs behave as if only the membrane formation had been caused by some agency. by carryformation. The consequence was that only a membrane formation resulted without the spermatozoon penetrating into the . membrane namely.. however. Nereis. the spermatozoon however. In this case the egg can undergo complete development into a larva. staroccurs sperm is used for the fertilization of the sea-urchin egg.

The solution of these problems must be reserved for further investigation. and similar experiments I concluded that the spermatozoon causes the development by accelerating the oxidations This conclusion was confirmed by experiments in the egg. if the oxygen is again promptly as soon as oxygen this completely withdrawn the development stops. con- clude that the spermatozoon causes the development of the way similar to that which takes place It carries first a in the case of parthenogenesis. at present is the way in which the destruction of the cortical layer of the egg accelerates the It is possible that the cortical layer acts like a solid crust and thus prevents the oxygen from reaching the surface of the egg or from penetrating into the latter sufficiently rapidly. Years ago I had found that the fertilized sea-urchin egg can only develop in the presence of free oxygen. substance into the egg which destroys the cortical layer of the egg in the same way which as does butyric acid. and secondly a substance egg after the membrane the destruction of the corti- corresponds in its effect to the influence of the hyper- tonic solution in the sea-urchin formation. egg in a artificial This membrane formation led only to a beginning but We may. therefore. Warburg was able to show that the mere membrane formation by the butyric acid treatment has the same accelerating effect upon the oxidations causation of the as fertilization. . but begins From is again admitted. What remains unknown oxidations.The Mechanistic Conception of Life egg. This question leads us to the process of oxidation. Warburg and by Wasteneys and myself in which it was found that through the process of of oxidations in the egg is fertilization the velocity its increased to four or six times original value. 13 not to a complete development. The question cal layer arises as to how can cause the beginning of the development of the egg. by O.

individual of the sea-urchin Death was assumed to be principle'' from the body. The answer assumed the anthropomorphic form characteristic of all explanations of nature in the prescientific period. the surface films of the contain enough water and high. which twelve years ago was shrouded in complete darkness. It is. Life was assumed to begin with the entrance of a individual life '^life principle" into the body. NATURE OF LIFE AND DEATH The nature of life and of death are questions which occupy the interest of the layman to a greater extent than possibly any other purely theoretical problem. due to the departure of this ''life Scientifically. however. As soon ceased for some time.t has been accomplished in this problem will occur in rapid succession in those problems which today still appear as riddles. if is tion of oxidation in the body. and we can well understand that humanity did not wait for experimental biology to furnish an answer.14 The Mechanistic Conception of Life We therefore see that the process of the activation of the egg by the spermatozoon. therefore. The problem of the beginning and end of individual life is physico-chemically clear. and the body is destroyed by micro-organisms. have they the temperature sufficiently become permeable for bacteria. is today practically com- pletely reduced to a physico-chemical explanation. life begins (in the case and possibly in general) with the accelerathis acceleration tion of the rate of oxidation in the egg. unwarranted to continue the statement that in addition to the acceleration of oxidations the beginning of . and begins after the destruction of its cortical layer. that begins with the egg was of course unknown to primitive or prescientific man. V. Life of warm-blooded animals —man if included —ends with the cessaas oxidations cells. Conright sidering the youth of experimental biology we have a to hope that whd.

Hagedoorn. We can cause an egg to develop into a larva without sperm.The Mechanistic Conception of Life individual life is 15 determined by the entrance of a meta- physical and that death is the from cessation of oxidations. the difference between spermatozoon and effects of the spermatozoon cease and activator to the egg. however. This fact has some bearing upon the further investigation of heredity. a star-fish. VI. far as the transmission of paternal characters is con- we can say today that the view of those authors was correct who. localized this transmission not only in the cell nucleus. by the determined. the result is a seaurchin larva which possesses no paternal characters. If the egg of a sea-urchin is fertilized with the sperm from a different species of sea- form has distinct paternal characters. but we cannot cause a spermatozoon to develop into a larva without an egg. but in a special constituent of the nucleus. Kupelwieser. In the case of ''life principle" into the egg. egg exceeds a certain limit the hereditary it acts merely as an As cerned.. and Baltzer were able to confirm. The spermatozoon can influence the form of the offspring only when the two forms are rather closely related. while urchin. e. with Boveri. inasmuch as it shows that the egg is the main instrument of heredity. the larval apparently the spermatozoon is restricted in the transmission If of characters to the offspring. as I found and as Godlewski. the evaporation of water we are satisfied with the explanation given by the kinetic theory of gases and do not of the demand that —to repeat a well-known jest of Huxley—the disappearance "aquosity" be also taken into consideration. aside departure of this ''principle" from the body.g. . HEREDITY is It may be stated that the egg the essential bearer of heredity. If. the eggs of a sea-urchin are fertilized with the sperm of a more remote species.

the other to the pure maternal type.16 The Mechanistic Conception of Life The proof for this the chromosomes. if it is a male. Assuming it to be true that the chromatin is the important part of the cell in the matter of heredity. along the lines of Mendelian investigations. investigation of the heredity of those qualities which occur mainly in one sex. and one upon wliich almost is all investi- gators are united in opinion. Considering the importance of this idea we may render it in his own words: A most significant fact. The proof view was furnished through the e. simplest form be expressed forms which in the following way.. that Henking's Montgomery afterward showed an accessory chromosome. was given by facts found The essential can in If its law of Mendel. two kinds of spermatozoa The one kind corresponds to the pure paternal.g. form of insects {Pyrrhocoris) two kinds of spermatozoa exist which differ in the fact that the one possesses a nucleolus while the other does not. nucleolus was McClung first expressed the idea that this accessory chromosome was connected with the determination of sex. . its essential Each animal has a definite number of chromosomes Henking had found that in a certain in its cell nucleus. two kinds of eggs if it is a female. Nine years ago McClung published a paper which solved the problem of sex determination. that the element is apportioned to but one-half of the spermatozoa. at least in feature. the law of segregation. The investigation of the structure and behavior of the nucleus showed that the possibility for such a segregation of the sex-cells in a hybrid can easily be recognized during a given stage in the formation of the sex-cells. we cross two differ in only one character every hybrid resulting from this union forms two kinds of sex-cells in equal numbers. color blindness which occurs pre- eminently in the male members of a family. if the assumption is made that the chromo- for the correctness of this somes are the bearers of the paternal characters.

H. animals. A careful consideration will suggest that notliing but sexual characters thus divides the members of a species into two well-defined groups. B. correct for many While in of eggs many animals there are two kinds of sperma- tozoa and only one kind of eggs. We expect. have amply confirmed the correctness of this ingenious idea and cleared up the problem of sex determina- and others. which differ by one chromosome. I must here also point out a fact that does not seem to have the recognition it deserves. is e. The eggs are all equal in these More recent investigations. therefore. in other animals two kinds and only one kind of spermatozoa are formed. especially the work of E. T. Wilson.. according to Guyer. that exhibit marked differences in structure. tion in its main features. that if there is a cross-division of the must be two kinds of spermatozoa regardless of the presence of the accessory chromosome. B. and we are logically forced to the conclusion that the peculiar chromo- some has some bearing upon the arrangement. . especially those of is E. sea-urchins and certain species of birds and of butterflies (Abraxas). According to McClung each animal forms two kinds of spermatozoa in equal numbers." The brilliant researches of the following years. in the It is of interest that. imder normal conditions. One kind of spermatozoa produces male animals..The Mechanistic Conception of Life 17 then it follows that we have two kinds of spermatozoa that differ from each other in a vital matter.g. there a preponderant maleness would attach to one-half of the spermatozoa. in man. human being two kinds of sperma- tozoa exist and only one kind of eggs. have shown that this view animals. viz. therefore. Wilson. to find in the offspring two sorts of individuals in approximately equal numbers. due to the ''qualitative division of the tetrads. In these animals the sex predetermined in the egg and not in the spermatozoon. Miss Stevens. the other female animals. It is thus possible that even in the absence of any specialized element chromosomes in the maturation mitoses. Morgan. sex is determined by the spermatozoon.

According to Wilson all unfertilized eggs contain in this case one so-called Diagrammatic presentation of sex deterFigs. one with There are two kinds and one without an X-chromosome. namely. Hence one-half of the eggs will contain after fertilization two X-chromosomes each and one-half only one X-chromosome . e represents a fertilized egg after a spermatozoon of the type c (without a sex chromosome) has entered it. This egg contains after fertilization only one sex chromosome X and hence will give rise to a male. Each contains one sex cliromosome marked X. B.18 The Mechanistic Conception of Life How is sex determination accomplished ? Let us take the case which according to Wilson is true for many insects and according to Guyer for human beings. that there are two kinds of spermatozoa and one kind of eggs. one-half of the eggs will be fertilized by spermatozoa with and one-half by spermatozoa without an X-chromosome. sex sex chromosome. Given a sufficiently large number of eggs and of spermatozoa. b containing a sex chromosome X. This egg now has two — chromosomes and therefore wiU give rise to a female. Wilson). mination in an insect (Protenor). 11-16 (after E. the other six dark spots are the chromosomes which are supposed to transmit hiereditary characters not connected with sex. b and c represent the two different types of sperm. c being without such a chromosome. the X-chromosome. d represents the constitution of the egg nucleus after it is fertilized by a spermatozoon of the type b containing a sex chromosome. of spermatozoa. a a are the nuclei of im^fertUized eggs.

while in the male are found two kinds of spermatozoa. those containing two X-chromosomes give rise This seems as Wilson and others have proved. These observations show for example.g. It is stated that the number slight percentage. in the why it is impossible to influence If. since the cells of both twins have the same number X-chromosomes.The Mechanistic Conception of Life each. If. one with and one without a sex chromosome. in these animals an egg is of males bom exceeds that of the females by a statement is correct it must be due to a secondarycause. was known that from eggs in these animals only females develop. series of other difficulties.^ These discoveries solved also a tion. the sex of a developing embryo by external influences. the egg will give rise to a a spermatozoon with an X-chromosome girl. If this . e. therefore. a greater motility or greater duration of life of the male spermatozoon. 19 The eggs containing only one X-chromosome give rise to males. as we should of expect. bees. Further researches will be needed to clear up this point. In plant lice. human being a spermatozoon without an X-chromosome boy. males never. but if enters into an egg. to females — to be a general law for those cases in which there are two kinds of spermatozoa and one kind of eggs. and the same is probably true for bees 1 and ants. and ants. the eggs It may develop with fertilized and without It fertilization. Certain types of twins originate from one egg after fertiliza- Such twins have always the same sex. gets into the egg the latter will give rise to a Since always both it is kinds of spermatozoa are given off by the male girl a matter of chance whether a boy or a it originates.. For Phylloxera and Aphides it has been proved with certainty by Morgan and others that the spermatozoa which contain no sex chromosome cannot live. was found that in these animals the eggs contain only one sex chromosome. mere and is agrees with the law of probability that in a large popula- tion the number of boys and girls born within a year approximately the same.

In the male the one X or have from the begin- ning only one X-chromosome and each primordial sperma- tozoon divides into two new spermatozoa.g. It had been known males only. two pairs of) an X-chromosome while the other is without such a chromosome. is That this not a mere assumption can be color blindness sho"v\Ti in those cases in which the hereditary character appears only. a division of the chromosomes occurs. If a color-blind individual . during the critical. time that in bees and ants the unfertilized eggs can also develop. What can be observed here directly in the male animal takes place in every hybrid. found a simple solution. insects and in man the cells of the female have chromosomes.20 The Mechanistic Conception of Life always done by a spermatozoon which contains fertilized it is an X-chromosome. whereby only one-half of the sex-cells receive the hereditary substance in regard to which the two original (in reality into one of which contains pure forms differ. but such eggs give these animals contain only one due to the fact that the eggs of X-chromosome and from eggs with only one chromosome only males can arise (at least in the case of animals in which the male is heterozygous for sex). therefore. retains only cells sex chromosome. therefore. in these animals The egg has. and simultaneously Mendel's law of segregation also finds its solution. which appears is mated with an individual with normal color vision the heredity of color mostly in the male. after fertilization always two X-chromosomes and from such for a long eggs only females can arise. in one sex as.. or pre- eminently. rise to This is The problem of sex determination has. Each egg. In a certain stage of the history of the two sex egg one-half of the chromosomes leave the egg (in the form of the '^ polar-body") and it keeps only half the number of In many chromosomes. so-called maturation division of the sexual cell in the hybrid. therefore. e.

In the next generation color blindness cannot appear. since each fertilized egg contains the factor for color perception. The factor for color vision is obvi- ously transmitted through the sex chromosome. since all flies had the factor for pigment in their sex-cells. however. white eyes.The Mechanistic Conception of Life 21 blindness in the next two generations corresponds quantitatively ^vith what we must expect on the assumption that the chromosomes. and especially of Morgan. We say pre-eminently. Doncaster. which behave the eyes. In man these conditions cannot in a fly (Drosophila) T. . we may con- clude with certainty that the sex chromosomes are the bearers of those hereditary characters which appear pre-eminently in one sex. in the second generation all females and exactly one-half of the males had red eyes. the other half of the males. like blindness. Morgan has observed tion with white eyes. pigment in These a muta- have normally red eyes. which occurs in the male. When he crossed a white-eyed with a red-eyed female first all flies of the generation were red-eyed.. Morgan has found lack of a number color flies of similar sex-limited characters e. H. as ]Mendel already intimated. the theory In the second of the demands that one-half males should be color blind. generation.g. From these and numerous similar breeding experiments of Correns. since theoretically we can predict cases in which color blindness or white eyes also in the female. must appear law Breeding experiments have is justified. In the color-blind individual some- chemical substances determining color vision are contained in the sex thing is lacking which can be found in the individual with normal color perception. sho^\TL that this theoretical prediction The riddle of Mendel's of segregation finds its solution through these experiments incidentally also the problem of the determination of is and sex which only a special case of the law of segregation. as the theory demands. be verified. however.

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Here the ground has already been broken. for absorption and secretion. however.The Mechanistic Conception of Life The main task which is left 23 is here for science to accomplish the determination of the chemical substances in the chromo- somes which are responsible for the hereditary transmission of a quality. all where facts cannot only be predicted qualitatively. and the determination of the mechanism by which these substances give rise to the hereditary character. of fertilization selected only the phenomena phenomena are specific for living We have and heredity.. therefore. but also quantitatively. as. While until twelve years ago the field of heredity was the stamping ground for the rhetorician and metaphysician it is today perhaps the most exact and rationalistic part of biology. may. settle a question which offers itself not only to the layman but also to every biologist. e. It is knowTi that for the formation of a certain black pigment the cooperation of a substance — —tyrosin—and of a ferment We of oxidation —tyrosinase is required. since organisms and without we can convince our- analogues in inanimate nature.g. and if selves that these processes can be explained physico-chemically we may safely expect the same of such processes for which there exist a-priori analogies in inanimate nature. THE HARMONIOUS CHARACTER OF THE ORGANISMS not possible to prove in a short address that will yield to all life It is phenomena these a physico-chemical analysis. VII. The hereditary transmission of the black color through the male animal must occur by substances carried in the chromosome which determine the formation of tyrosin or tyrosinase or of both. must. say that the solu- tion of the riddle of heredity has succeeded to the extent that further development will take place purely in cytological and physico-chemical terms. how we shall conceive that wonderful " adaptation of each part to the We . namely.

24

The Mechanistic Conception of Life
In the answer

whole " by which an organism becomes possible.

to this question the metaphysician finds an opportunity to put

above the purely chemical and physical processes somethingspecific

which is characteristic

of

life

only: the ''Zielstrebigkeit,"

the ''harmony" of the phenomena, or the "dominants" of

Reinke and similar things.

With
I

all

due personal respect for the authors

of

such terms
all

am

of the opinion that

we

are dealing here, as in

cases of

metaphysics, with a play on words.
structed that
it

serves the ''whole"
is

is

That a part is so cononly an unclear expression

for the fact that a species

only able to live
if it is

expression

—or to use Roux's

is

only durable,

provided with the automatic

mechanism for self-preservation and reproduction. If, for instance, warm-blooded animals should originate without a circulation they could not remain alive, and this is the reason why we never find such forms. The phenomena of "adaptation" cause only apparent
difficulties since

we

rarely or never

become aware
which appear
that the

of the

numerous

faultily constructed

organisms

in nature. of species

I will illustrate

by a concrete example
is

number

which we observe

only an infinitely

small fraction of those which can originate and possibly not
rarely do originate, but which

we never
it is

see since their organiza-

tion does not allow

them
fish

to continue to exist long.

Moenk-

haus found ten years ago that
of each

possible to fertilize the egg

marine bony

with the sperm of practically any

other marine bony

fish.

His embryos apparently lived only
I

a very short time.
over a month.
practically

This year

succeeded in keeping such
fish alive for

hybrid embryos between distantly related bony
It
is,

therefore, clear that

it is

possible to cross

any marine teleost with any other. The nmnber of teleosts at present in existence
If

is

about

10,000.

we accomplish all possible hybridizations 100,000,000
Of these
teleosts only a very small of 1 per cent,

different crosses will result.

proportion,

namely about one one-hundredth

The Mechanistic Conception of Life
can
live.

25

It

turned out in

my experiments that the heterogenefins,

ous hybrids between bony fishes formed eyes, brains, ears, and pulsating hearts, blood and blood-vessels, but could

live

only a limited time because no blood circulation was established

in spite of the fact that the heart beat for
if it

weeks
fish

—or that the

circulation,

was estabhshed at

all,

did not last long.

What
nants"?
of faulty

prevented these heterogeneous

embryos from

reaching the adult stage?
Scarcely.
I

The

lack of the proper "domi-

succeeded in producing the same type

embryos

in the pure breeds of a

bony

fish

{Fundulus

heteroclitus)

by raising the eggs in 50 c.c. of sea-water to which was added 2 c.c. 1/100 per cent NaCN. The latter substance retards the velocity of oxidations and I obtained embryos which were in all details identical with the embryos produced by crossing the eggs of the same fish with the sperm of remote These embryos, which teleosts, e.g., Ctenolahrus or Menidia.
lived about a

month, showed the peculiarity

of possessing

a

beating heart and blood, but no circulation.

This suggests

the idea that heterogeneous embryos show a lack of ''adapta-

tion" and durability for the reason that in consequence of the
chemical difference between heterogeneous sperm and egg the
chemical processes in the fertilized egg are abnormal.

The

possibility of hybridization goes

much

farther than

we

have thus

far assumed.

We
;

can cause the eggs of echinoderms

to develop with the sperm of very distant forms, even mollusks

and worms (Kupelwieser)
It
is,

but such hybridizations never lead

to the formation of durable organisms.

no exaggeration to state that the number of species existing today is only an infinitely small fraction of those which can and possibly occasionally do originate, but
therefore,

which escape our notice because they cannot five and reproduce. Only that limited fraction of species can exist which possesses

no coarse disharmonies
tion

in its

automatic mechanism of preserva-

and reproduction.

Disharmonies and faulty attempts in

26

The Mechanistic Conception of Life
rule,

nature are the
exception.

the harmonically developed systems the rare

But

since

we only

perceive the latter

we

gain the

erroneous impression that the ''adaptation of the parts to the

plan of the whole" is a general and specific characteristic of animate nature, whereby the latter differs from inanimate nature. If the structure and the mechanism of the atoms were

knowTi to us we should probably also get an insight into a world
of wonderful

harmonies and apparent adaptations of the parts

to the whole.

But

in this case

we should quickly understand

that the chemical elements are only the few durable systems

among a large number of possible but not durable combinations. Nobody doubts that the durable chemical elements are only
the product of blind forces.

There

is

no reason

for conceiving

otherwise the durable systems in living nature.
VIII.

THE CONTENTS OF LIFE
from the cradle to the bier are wishes
struggles,

The contents
and hopes,

of life

efforts

and

and unfortunately
this inner life

also dis-

appointments and

suffering.

And

should be

amenable to a physico-chemical analysis? In spite of the gulf which separates us today from such an aim I believe that
it is

attainable.

As long
lifted
first

as a

life

phenomenon has not yet found
usually appears inexplicable.

a physico-chemical explanation
If

it

the veil

is

once

we

are always surprised that
it.

we

did

not guess from the

what was behind

That
tion
is

in the case of our inner life a physico-chemical explanais

not beyond the realm of possibility

proved by the fact

that

it is

already possible for us to explain cases of simple mani-

festations of animal instinct
basis;

and

will
I

on a physico-chemical
in former

namely, the phenomena which

have discussed

papers under the
simple example

name

of

animal tropisms.

As the most

we may mention

the tendency of certain ani-

mals to

fly or

creep to the light.

We

are dealing in this case

with the manifestation of an instinct or impulse which the

The Mechanistic Conception of Life
animals cannot
these animals
resist.

27

It

appears as

if

this blind instinct

which

must

follow, although

it

may
of

might be explained by the same law
which explains the photochemical

them their life, Bunsen and Roscoe,
cost

effects in

inanimate nature.

This law states that within wide limits the photochemical effect
equals the product of the intensity of light into the duration
of illumination.
It is

not possible to enter here into

all

the

details of the reactions of these animals to light;

we only wish

to point out in which

way

the light instinct of the animals

may

possibly be connected with the Bunsen-Roscoe law.
positively hehotropic animals

The

go instinctively to a source of light

— —have

i.e.,

the animals which
in their eyes

(and

occasionally also in their skin) photosensitive substances which

undergo chemical alterations by
indirectly,
is

light.

The products formed

in this process influence the contraction of the muscles

—mostly
is

through the central nervous system.

If

the animal

illuminated on one side only, the mass of photochemical

reaction products formed on that side in the unit of time
greater than on the opposite side.

Consequently the develop-

ment of energy in the symmetrical muscles on both sides of the body becomes unequal. As soon as the difference in the masses
of the photochemical reaction products

on both sides of the
it

animal reaches a certain value, the animal, as soon as
is

moves,
as

automatically forced to turn toward one side.

As soon

it

has turned so far that

its

plane of symmetry

is

in the direction

of the rays, the symmetrical spots of its surface are struck

by

the light at the same angle and in this case the intensity of light

and consequently the velocity of reaction of the photochemical processes on both sides of the animal become equal. There is no more reason for the animal to deviate from the motion in
a straight line and the positively heliotropic animal will
in this line to the source of light.
(It

move
in

was assumed that

these experiments the animal

is

under the influence of only

one source of light and positively heliotropic.)

18 Fig. e. The which new polyp-bearing stems grow the direction of the rays of Figs. certain hydroids and worms.. e. Motile plant organs. 18 and 19.g. 19 —Positive heliotropism of the polyps of Eudendrium. all in light is indicated by an arrow in each figure. the swarm spores of plants. (From nature. their stems to the source of light until the axis of their tip is symmetry I of in the direction of the rays of light. move to the source of light (or if they are negatively .. It was knowTti that sessile heliotropic plants bend Fig. found the same phenomenon in sessile animals.g.) These animals bend in the same way to the light as the stems of positively heliotropic plants kept under similar conditions.28 The Mechanistic Conception of Life In a series of experiments I have shown that the heliotropic reactions of animals are identical with the heliotropic reactions of plants.

The direct in regard to the applicability of Bunsen's law to the phenomena of animal heliotropism have not yet been made. Positive heliotropism of a marine worm (Spirographis). The on our had already been proved measurements for the action of light retina. and the same result It by another botanist.) The light fell into the aquarium from one side only and the worms all bent their heads toward the source of light. The first of these facts is the identity of the light reactions of plants and animals. In plants only the and the same is observed in motile more refrangible rays from green to and the same is blue have these heliotropic effects. It has been shown by Blaauw for the heliotropic curvatures of plants that the product of the intensity of a source of light into the time required to induce a heliotropic curvature is a Fig. was obtained simultaneously is thus proved that the Bunsen- Roscoe law controls the heliotropic reactions of plants. as the stems of positively heliotropic plants would do under the same conditions. The second is at least a rough observation which harmonizes mth the Bunsen-Roscoe law. As long as the intensity of light or the mass of photochemical substances at the surfaces of the .The Mechanistic Conception of Life heliotropic 29 away from it). same fact Froschl. But a number of data point to the probability that the law holds good here also. animals. 20. while the red and yellow rays are little or less effective. (From nature. — constant. true for the heliotropic reactions of animals.

since according to this law the photo- chemical effect is equal to the product of the intensity of the If. The need its of and the its struggle for food. the inten- sity of the light is strong or the active mass of the photochemical substance great. if photosensitive substances on the surface of the animal) small the animals go in irregular lines. . the sexual instinct with poetry and chain of consequences. The behavior If of the animals agrees with this assumption. since the directing force not It will. from the mechanistic point of view." Our wishes and hopes. the muscles through the influence of physical term 'Svill" and for the meta- we may in these instances safely substi- tute the chemical term "photochemical action of light. But we may already safely state that the apparent will or instinct of these animals resolves itself into a modification of the action of light. is it must take a comparatively long time until the animal oriented automatically by the light. disappointments and their source in instincts instinct of sufferings have light which are comparable to the the heliotropic animals. however. however. For some of these instincts the chemical basis is at least sufficiently indicated to arouse the hope that their analysis. is only a question of time.30 The Mechanistic Conception of Life is animal small. the instinct of workmanship. according to the law of Bunsen. but at last they also is land at the source of entirely abolished. light. and some other instincts are the roots from which our inner life develops. be necessary to ascertain in by direct measurements to what extent these phenomena animals are the expression of Bunsen-Roscoe's law. sufficiently strong the the light is animals go in an almost straight line to the intensity of light (or the mass of is the source of light. light into the duration of illumination. the maternal instincts with the felicity and the suffering caused by them. difference in the it will require only a very short time until the mass of photochemical reaction products is on both sides of the animal reaches the value which for the necessary automatic turning to (or from) the light.

social.The Mechanistic Conception of Life ix. pelled to be so by processes in our central nervous system. The mother loves and cares for her children. we are compelled to do so. ditions or ignorance faulty or low development of ethics. isms we ourselves are only chemical mechan- —how can there be an ethics for us? The answer is. because we are com- are just as hereditary as drink. If 31 ethics our existence is based on the play of blind forces and only if a matter of chance. machine-like. life life compatible \vith ethics it seems the only con- which can lead to an understanding of the source . We struggle and truth since we are instinctively compelled to see Economic. We eat. arise in Individual mutants is lost. and as long as human beings are not economic slaves the instinct of successful work or of workmanship determines the direction of their action. and numerous enough. but because. not because metaphysicians had the idea that this was desirable. the ethical status of a community. that our instincts are the root of our ethics and that the instincts is the form of our body. and political conand superstition may warp and inhibit the inherited instincts and thus create a civilization with a our fellow beings happy. but because the instinct of taking care of the young body. may which one or the other desirable instinct just as individual mutants without pigment may arise if the offspring of such mutants may. is inherited just as distinctly as the morphological characters of the female We seek and enjoy the fellowship of human beings because hereditary conditions compel us to do for justice so. We are active. in animals. lower is Not only : the mechanistic conception of ception of of ethics. and reproduce not because mankind has reached an agreement that this is desirable.

.

II. THE SIGNIFICANCE OF TROPISMS FOR PSYCHOLOGY .

.

phenomena which the metaphysician would classify under the term of animal 'Svill. namely.) Reprinted by courtesy of Professor James McKeen Cattell. 35 . Five years of experiments with extirpations in the cerebral cortex proved to me without doubt that Munk had become the victim of an error and that the method of cerebral operations may give data concerning the path of nerves in the central it nervous system but that brain processes. analysis of these Some authors hold that even if a complete physico-chemical phenomena were possible today it would leave the 'Hruly psychical" imexplained. von Hartmann I became interested in the problem of vrill. Watkinson. When as a student I read Munk's investigations on the cerebral cortex I believed that they might serve as a starting-point for an experimental analysis of will. We are at least able to show for a limited group animal reactions that they can be explained unequivocally on a purely physico-chemical basis.II THE SIGNIFICANCE OF TROPISMS FOR PSYCHOLOGY^ I A mechanistic conception of life is not complete miless it includes a physico-chemical explanation of psychic phenomena. into a discussion of such We do not need to enter still an objection since we are too far from the of goal." Through the writings of Schopenhauer and E. (After a translation in Popular Science Monthly by Miss Grace B. teaches little about the dynamics of A of the 1 better opportunity seemed to offer itself in the study comparative psychology of the lower animals in which Lectiire delivered at the Sixth International Psychological Congress at Geneva. 1909. Munk stated that he had succeeded in proving that every memory image in a dog's brain is locahzed in a particular cell or group of cells and that any one of these memory images can be extirpated at desire.

Experimentally. which determine the movements and in discovering the laws according to which these forces act.36 The Mechanistic Conception of Life memory is the mechanism for all. by external agencies. developed but sHghtly or not at it It seemed to me that some day must become possible to discover the physico-chemical laws underlying the apparently random movements of such animals. and that the word ''animal will" was only the expression of our ignorance of the forces which prescribe to animals the direction of their apparently spontaneous movements just as unequivocally as gravity movements of the planets. But if one part only of the animals moves in this definite direction and the other does not. For if a savage could directly observe the movements of the planets and should begin to think about them. we speak of an act of will. Only if we succeed in this have we the right to assume that we know the force which under certain conditions seems to a layman to be the ^\ill of the animal. We is. In the latter case purely phj^sical forces are concerned. If a sparrow flies down to a seed lying on the ground. while in the former chemical reactions are also taking place in the sense-organs. we have not succeeded in finding the force which unequivocally determines the direction of their movement. nerves. The scientific solution of to consist in finding the forces of animals. plays its part also. that the natural movement of locomotion. only when this latter complex. and muscles of the animal. but if a dead sparrow falls upon the seed this does not appear to us as such. One other point should be observed. he would probably come to the conclusion that a ''will action" guides the movements of the planets prescribes the is just as a chance observer today inclined to assume that "will" the problem of will seemed then causes animals to move in a given direction. any number of individuals of a given kind of animals to move in a definite direction by means of their locomotor apparatus. speak of an act of will. the solution of the problem of will must take the form of forcing. and .

If is directed toward the source they remain there. psychology of the II Some experiments on winged direction of their progressive plant lice may serve as an introduction to the methods of prescribing to animals the movements.g. fifty suitable perhaps twenty-five or subjects for the experi- With these animals it may be demonstrated that the direction of their movement toward the light ment. the aphids (plant into winged insects. change After this metamorphosis the animals leave the plants. In this manner a sufficiently large number. which causes the animals to drop into the test-tube. The animals place themselves with their heads toward the source of light and move toward it in as straight a line as the imperfection of their locomotor apparatus allows. fly to the window. in a the test-tube is crowded mass. . lice). They can then easily be collected upward by holding a test-tube underneath and touching one animal at a time from above with a pen or scalpel. is and that the light is not too weak. stationary. turned 180"^ the animals again go straight toward the source of light until the interference of the glass stops their further progressive move- ments. artificial light.. 1906. If the plants are allowed previously wingless. is definitely determined —provided that the animals are healthy e. 1889. and there creep on the glass.^ It can be demonstrated in these animals that the 1 Loeb.Significance of Tropisms for Psychology it is 37 only with this sort of reactions that we have to deal in the will. used. potted rose bushes or Cinerarias infected with plant lice are brought into a room and placed in front of a closed window. The experiment is so arranged that only a single source of light. may be obtained. Wurzburg. When closely they reach that end of the test-tube which of light. approaching as near to the source of light as their prison (the test-tube) permits. In order to obtain the material. to dry out. Translated in Studies in General Physiology. Der Heliotropismus der Tiere und seine Uebereinstimmung mit dem Heliotropismus der Pfiamen.

^ justified in many is chemical reactions of organic Especially is bodies are accelerated by light.g. New York. this true of oxi- The mass is of facts already so great that we are assuming that the determining action of in its last analysis light upon animals and plants due to the fact that the modified rate of certain chemical reactions in the cells of the retina or of other photosensitive regions of the organisms is by light. as also of Wolfgang Ostwald (Biochem..38 The Mechanistic Conception of Life movement is direction of their progressive just as unequivocally governed by the source of light as the direction of the move- ment of the planets is determined by the force of gravity. 1905. by which I mean that s^inmetrical regions of the body are chemically identical and have the same metabolism. and the second the photochemical action of light. as well as in an anatomical. We will consider the light. 1906. two separately. chemical reactions. oxidation. have an identical metabolism. see the work of Ciamician. Xlll. Biochem. which are certainly symmetrical. 1907).. 1 The individual points on Luther. while a region of the skin which is not symmetrical with the retina has a different metabolism. 1908. Loeb. C. Zeitschr. while non-symmetrical regions of the body are chemically different. difference it is In order to illustrate this only necessary to point out that the two retinae. . But it is my belief that such a symmetry exists in a chemical sense. The second factor is the symmetrical structure of the animal. Neuberg. and in general have a quantitatively or qualitatively different metabolism. In addition. Die Aufgaben der Photochemie.. The theory of the compulsory movements of aphids under the influence of light gressive is as follows : Two factors govern the pro- movements of the animals under these conditions. Leipzig. In regard to the photochemical action of that a great we know today dations. one is is the symmetrical structure of the animal. 305. The Dynamics of Living Matter. increases. with increasing intensity of light the rate of certain e. As expressed in the gross anatomy of the animal. Zeitschr. the right and left halves of the body are symmetrical.

that the rate of the nerve impulse has a temperature coefficient of the order of magnitude which reactions. take place in the two optic nerves (or in the sensory nerves of the This inequality of chemical processes two illuminated points) the motor nerves and eventually to to sensory the passes from the muscles connected with them. that with equal illumination of both retinae the symmetrical groups of muscles of both halves of the body will receive equal chemical stimuli and thus reach equal states of contraction. are photosensitive. We conclude from this . as my experiments and those of Parker have shown. is characteristic for chemical From this we must conclude that when two retinae (or other points of symmetry) are illuminated with unequal intensity. and accordingly more intense chemical changes will take place in one optic nerve than in the other.Significance of Tropisms for Psychology the retina are also chemically unlike. the chemical reactions will also be more accelerated in the one retina than in the other. The same naturally holds true for every other pair of sym- metrical photosensitive surface elements. Maxwell and CD. when the rate of reaction is unequal. In planarians. and the peripheral parts of the retina indicate that the points of sjnmmetry of the two retinae are chemically alike. upon the other. S. but also in other places on the surface of many animals. the symmetrical into stronger action than muscles on one side of the body come . 39 The observations upon visual purple. chemical processes. S. not only the eyes. while. retina than Snyder have demonstrated. if Now more an unequal amoimt of light falls upon the two retinae the photochemical reactions in the one which receives light will also be more accelerated than those in the other. independently of each other. the non-symmetrical points chemically unlike. For it should be mentioned that photochemical substances are not found in the eyes only. also of unequal intensity. the differences in the color sensitiveness of the fovea centralis. but also parts of the But if more light falls upon one skin.

40 The Mechanistic Conception of Life The result of such those on the other side. Loeb. ' If two sources of light of equal intensity are at an equal distance from the animal. it will move in a direction at right angles to a line connecting the two sources of light. As soon as this happens. direction of movement can result either in consequence. the movement in the latter case not being determined by heliotropism.. and with it the whole body of the animal.As a result the head. is turned toward the source of light. that a single source of light is the animal from one side. It is thus automatically guided toward the source just as of light. or in a turning of the head and the animal in the opposite direction. 1900. action of an inequality of the is symmetrical muscles of the two sides of the body in the a change in the direction of The change movement on the part of the animal. it is gravity in the case of a falling stone or the action direct. Comparative Physiology of the Brain and Comparative Psychology. whole animal toward the source of nervous system light. The of gravity upon the movement of the falling stone is while the action of light upon the^direction of movement is of the aphids indirect. New York and London. the machine-like heliotropic reaction of animals differs from the movement of a human being toward one of two sources of light. . In the winged aphids the relations are as follows: Suppose present and that the light strikes As a consequence the activity of those muscles which turn the head or body of the animal toward the source of light will be increased. movement of a planet. as Bohn has rightly said. inasmuch as the animal is caused only by means of an acceleration of photochemical reactions to move 1 in a definite direction. the two retinae become illuminated equally. is The structure of the central segmental and the head segments generally determine^ the behavior of the other segments with their accessory parts. In this instance the light determines the direction of is the ''will" of the animal which its movement. of the in a turning of the head and. There is therefore no longer any cause for the animal to turn in one direction or the other. Herein. because in this base both eyes are similarly influenced by the light.

and as negatively heliotropic those animals which are oriented or compelled to move in the opposite direction. vertebrates as well as We cannot. The same phenomena of positive heliotropism may be demonstrated with equal precision in a great invertebrates. In the presence of only the axis of sym- one source of light this condition is fulfilled if metry of the animal moves in the direction of the rays of light. We do not yet know whether is or not Bunsen's law holds good for the If it does. Experiments on the heliotropism of plants as well as on the perception of light by our retina have shown that the effect of light equals the product of the intensity into the duration of illumination.. unusually common among the larvae of marine animals and insects. of course. Whether an animal Why . all many who animals. This law is identical with the general law of Bunsen and Roscoe which of light is states that the chemical effect within wide limits equal to this product. but also not lacking in sexually mature individuals. photometric Heliotropic animals are therefore in reality machines.^ The aphids serve here only as an example. give here an account of these cases. with the cosine of the angle The animal is oriented its by the light in such a way that symmetrical elements of photosensitive surface are struck at about the same angle. 1 we shall have to substitute 5\(^^ positively or negatively heliotropic depends upon the fact whether the light causes an increase or a decrease in the tension of the muscles light should have these opposite effects is as yet imknown. light varies According to photometric laws the intensity of with the sine of the angle at which the light strikes (or a surface element of the animal of incidence). The reader is interested in them must look Heliotropism into the voluminous literature is upon this subject. In this case the velocity of photochemical reactions on both sides of the animal is the same and there its is no reason why it should deviate from this direction in progressive motions. Significance of Tropisms for Psychology 41 We will now designate as positively heliotropic those animals which are forced to turn their head or move toward the source of light. heliotropic animals.

heliotropic line as the imperfection of their toward the source of light.c. Many animals show no heliotropism it at all . namely. move toward the light. to go more toward the source of light.42 this The Mechanistic Conception of Life law for what the metaphysician Ill calls the will of these animals. and the majority perhaps pay no attention to the light. of the fresh water a few cubic centimeters of water charged with carbon dioxide are slowly added. many show only a slight reaction. or that the exception must be satisfactorily explained. crowded together on the illuminated side is added all the individuals and move in as straight swimming movements perand remain there closely is of the vessel. show no positive a fresh-water pond or illuminated from one heliotropism. which easily penetrates the the vessel pretty cells of much Some seem the animal. preferably carbonic acid. science it is serve as an example. Every other acid and . others in the opposite direction. because they all fulfil the above-mentioned requirement. The problem therefore presented itself of producing heliotropism artificially imder natural conditions. To every 50 c. it is in an aquarium which at is often found that these animals move about in random and distribute themselves irregularly. as was to be expected. If the correct amount become actively positively a mits. If the vessel turned 180°. The winged aphids without exception. all It was soon found. If small crustaceans of lake are taken with a plankton net at noontime or in the after- noon and placed side only. that individuals. while others show in a degree as pronounced as do the winged aphids. This condition changes instantly if we add to the water some acid. they go directly back again to the lighted acts like carbonic acid side of the vessel. For mechanistic a methodological postulate that the same law acts without exception. in animals which. that not organisms in their natural condition are equally suitable for these experiments.

according to Stieglitz. especially in the eye. is the mere addition of positively heliotropic.i How that it We will assume a sensitizer. complete slaves does the acid produce this result? acts as of the light. not sufficient to make Daphniae 564. . CXV. But if we add an acid this could act as a catalyzer. sufficient to turn the ani- mal toward the source In the catalysis of of light. under carbonic acid treatment. because if now the animal is struck on one side only by the light. Pflugers Archiv. on the surface of the animal. the difference in the chemical changes still on the two sides of the body remains too small to call forth a difference in tension or action in the muscles of the two sides of the body. esters. oxidation. as was suggested in the case of the aphids.Significance of Tropisms for Psychology alcohol acts in the 43 same manner. ence in the action of the muscles of both sides of the body and a In certain forms. Loeb. in the catalysis of esters. The mass the light of photochemical substance which is is often relatively small. a decrease acid 1 in temperature also increases the If tendency to positive heliotropism. for instance. The light produces chemical changes. only more weakly and much more light slowly. the acid acts. the difference in the reaction products in both retinae differ- becomes rapidly large enough to cause automatically a turning of the head toward the source of light. In this way the same intensity of light which before produced no heliotropic reaction now may cause a very pronounced positively heliotropic reaction. Animals which were previously indifferent to become. for instance. so that even acted upon by when the light strikes the crustacean (copepod) on one side only. 1906. in Daphnia and in certain marine copepods. as. for instance. only to the extent of increasing the active mass of the substance which undergoes a chemical change. provisionally In order to fix our ideas we will assume that the acid makes the animal more strongly positively heliotropic by increasing the active mass of the photosensitive substance.

But if the intensit}^ of the light or the photosensitiveness of is the animal lessened the animal may deviate for a longer period from the direction of the light rays. some zoologists seem to have found difficulty in explaining the behavior of those animals which come between the two extremes. For instance. one writer has asserted that ^^ith greater intensity of light the laws of heliotropic orientation hold good. for instance. in other words. turns the head even for the fraction of a second from the direction of the light raj's. the fraction of a second.44 this The Mechanistic Conception of Life may often be accomplished by simultaneously reducing the temperature. while with a lessened light intensity the animals react to light by the method be understood. of "trial and error. to deviate from the direction of the light rays. are strongly positively helio tropic all and diffi- those animals which do not react at to light offer no Nevertheless. Such animals but they no moving instead in zig-zag lines . before the difference in the mass of the reaction products formed on the two sides light. it requires only a short time. vessel. If the intensity of the light is sufficient and the active mass of photochemical reactions in the substance in the animal great enough. it of the animal reaches the value necessary for the compulsory In this case the has hardly time for if it turning of the head toward the source of animal is a slave of the light. do eventually reach the lighted side of the longer go straight toward it. a com- pulsory turning of the head toward the source of light occurs only when the difference in the rate of certain photochemical two eyes reaches a certain value." From a chemical standpoint the behavior of is is animals at low intensity tively heliotropic animal easily to If a posi- illuminated from one side. the difference in the photochemical reaction is products in the two retinae becomes so great that the head at once turned back automatically toward the source of light. IV The animals which culties to the observer.

difference in the behavior of heliotropic animals under greater or lesser illumination. Every animal is continually producing acids in its especially carbonic acid and lactic acid. Still another point is to be considered. It is 45 therefore not a case of a qualitative. If he is forced to find an . few seconds directly toward the source of Later the production of carbonic acid decreases and the animal again becomes indifferent to light and can move in any direction. If. can naturally give no explanation of their significance. It probably produces also substances which could have the Fluctuations in the rate of the production of opposite effect and which decrease the heliotropic sensitiveness of the animals.Significance of Tropisms for Psychology or very irregularly. for a short time. cells. finally gather at the lighted side of the vessel because the algebraic sum of the movements in the other directions becomes zero according to the law of chance. a temporary increase to in the production of carbonic acid sufficiently can increase the photosensitiveness of the animal to cause it move for the period of a light. Then the production Such animals of car- bonic acid increases again and the animal goes again. We have seen that acid increases the sensitiveness of certain animals to light. the active mass of the photosensitive substance in a copepod is relatively small. and fore it is there- erroneous to assert of animals that heliotropism determines the movement toward the source of light only under strong illumination. but of a quantitative. toward the light. for instance. A \^Titer who is not trained to interpret the variations in the behavior of such an animal chemically and physiologically. and such acids increase the tendency in certain animals to react hehotropically. these substances will also produce fluctuations in the heliotropic sensitiveness of the animal. possibly by increasing the active mass of the photochemical substance. but that under weaker illumination an essentially different condition exists. But it is plain that such animals do not reach the source of light by a straight path.

is the ovary filled if with mature larvae placed in a watch crystal with sea-water in the dark. also to the formation of acid.46 The Mechanistic Conception of Life ''trial explanation he will wind up with the suggestion of error" which is and no more chemical or have. it scientific than the explana- tions of metaphysicians in general. is. they react much more This might be interpreted as a case of ''learning. worked only with animals which were not pronouncedly heliotropic and whose photosensitiveness wavered about the threshold of stimulation in the manner described above. probably rightly. therefore. The larvae Balanus perforatus develop filled entirely in the dark. it may be brought about by the carbonic or lactic acids produced through the muscular activity . I also believe that observations upon animals which are not sufl&ciently photosensitive have caused in the line of the rays of light.^ many first writers to assert that heliotropic animals do not place themselves directly but that they have to learn the right orientation. however." as it is In so far not a case of a lessening of the stickiness of the feet or the removal of some other purely mechanical factor which retards the rate of movement. this A If very striking experiment contradicts of assertion.^ 1 Provided that only a single source of light is present. I In experiments with winged aphids after having often found that gone through the heliotropic reactions a few times quickly to light than at the beginning. that is. They were. Some authors seems. comparable to or identical with the increase in the rate of reactions in the experiments described here. the larvae emerge at once and. so-called " staircase " phenomenon of stimulation of a muscle is ascribed. This phenomenon. their photosensitiveness if Such animals are not suitable it is for experiments in heliotropism and necessary to first increase the laws of the action of light upon them are to be investigated. pronouncedly positively heliotropic before they came under the influence of the light. the increase of the amount of contraction with every new stimulus. light. 2 The . they are brought into the they move at once to the side of the watch crystal nearest to the window.

^ In this case it is clear that the chemical changes directly or indirectly connected with nutrition lead to a permanent dimiIn nution or disappearance of the photochemical reaction.Significance of Tropisms for Psychology 47 V As far back as 1889 I pointed out that the photosensitiveness of an animal is different in different physiological conditions and that. at the time maturity. op. The ant workers show no heliotropic of sexual reactions. I in certain have already mentioned that in the aphids distinct heliotropic reactions may only be when the animals have developed wings and have left The influence of the chemical changes which take place in animals upon heliotropism is much more distinct in the larvae of Porthesia chrysorrhoea. The larvae hatch from expected the plant. while in the males and females.) . cit. 24. the intensity of which continues to increase. rising temperature in the spring drives them out of the also be driven out of the nest in winter and they can by I an increase in temperature. When driven out of the nest in this condition they are strongly positively hehotropic and in natural surroundings any animals whose have never found heliotropic sensitiveness was more pronounced than it is in the young larvae of Chrysorrhoea. heliotropism is found often only in certain developmental stages. ants and bees the influence of substances from the sexual organs seems to be the determining factor in the production of positive heliotropism. therefore. p. hibernate in a nest. as young larvae. (This latter fact has been overlooked by writers. But as soon as the animals have once eaten. under natural conditions. It is a well-known fact that during sexual maturity special substances are formed which influence various organs. 1 For several Loeb. or physiological states of an animal. the positive heliotropism disappears and does not return even if they are again allowed to become himgry. a distinct positive heliotropism develops. The nest.. the eggs in the fall and.

but them to creep straight upward to the top of a tree or branch. The 1 fact that ants at the time of and bees become positively sexual maturity plays an important is heliotropic role in the noticeable in The physico-chemical cause insects many and crustaceans of this "restlessness" at present unknown. set free tiveness in the non-pregnant uterus. could cause the formation of any way he nonsub- number of deciduae in follicle pregnant a uteri. When yoimg larvae of Chrysorthe young larvae are awakened from their winter of the spring they are positively helio- sleep by the sunshine tropic. In this way they their heliotropism guides them to their food. I will not discuss these facts further in this place. cal significance. so that every mechanical stimulus causes the latter to form a decidua. which is . is of great biologiit I pointed out in former papers that serves to save the lives of the above-mentioned rhoea. Their positive heliotropism leaves them no freedom forces of movement. the odor or tactile stimulus of which stops the pro- gressive movement of the machine sets their eating activity again in motion. while in others they are not sensitive to light or are even negatively heliotropic. while without the circulation of stance in the blood the uterus did not react in this manner. Should now remain positively heliotropic they would be held fast of the twigs on the ends and would starve to death. Here they find the first buds. but refer to my readers my earlier papers.48 The Mechanistic Conception of Life Leo Loeb has found that the substances which are by the bursting of an egg foUicle cause a special sensiIn this instance. and the animals^ forces restlessness having eaten they once more lose their positive heliotropism. after But we have already mentioned that ward. It is common phenomenon that animals in certain larval stages are positively heliotropic. They can now is creep down- which characteristic of so many them to creep downward and until they reach a new leaf. produced by certain metabolic products in the animal body. This change in the heliotropic sensitiveness.

according to these obser- vations the bees at the time of the nuptial flight are positively heliotropic machines. only true that species possessing tropisms which would make reproduction and preservation I of the species impossible must die out. that this nuptial flight is I gained the impression merely the consequence of a very The case seems to be similar among the bees according to the following experihighly developed heliotropic sensitiveness. Radl.^ and myself. These observations may serve as examples of the way in which the analysis of the vital phenomena of certain animals shows tropisms to be elements of these phenomena. The heliotropic sensitiveness of the animals was so great that they crept upward within the box. is proper value. ment described by Kellogg. Der Phototropismus der Tiere. Parker. . Radl. following the direction of the light rays. Leipzig. Many observations of a similar nature are found in the papers of VI Under the influence of the theory of natural selection the view has been accepted by many zoologists is and psychologists that everything which an animal does for its best interest. Thus. of founded by Mendel and It advanced to the position reduces this idea to its a systematic science in 1900. I found that among the male and female ants of a nest the heliotropic sensitiveness increases steadily up to the time of the nuptial flight and that the direction of their flight follows the direction of the rays of the sun.Significance of Tropisms for Psychology vital 49 economy of these creatures. The bees were ready to out of the opening of the box used for the experiment swarm when he suddenly removed the dark covering of the box so that the light now entered it from above. the mating of these insects takes place during the so-called nuptial flight. and were not able to make the nuptial flight. The exact doctrine of heredity. As is well kno^\Ti. 1903. Georges Bohn.

In my opinion this difference the result of the physical difference in the action and of the electric current. however. it can be observed that an orientation in relation to the direction of the current takes place in many of the animals. no animal has ever had the chance to come under the is influence of an electric current. 1 a remarkably common cell is reaction And yet among A more direct contradiction of the of the ions view that the Or where the movement withm the retarded. and animals are placed in this trough. also demonstrable in the as. which have in recent years fallen into the hands of physiologists who happened to be working on galvanotropism. purely a laboratory product. With the exception of a few individuals.50 The Mechanistic Conception of Life Galvanotropism illustrates this fact in is a striking manner. This phenomenon we call galvanotropism. The number is of species which show typical galvanotropic reactions not so great as the number of light of those is showing typical heliotropism. . In galvanotropism the current lines or the current curves play the same role as the light rays in heliotropism. galvanotropism animals. retina is involved. For this reason. Thus the action of the current upon the skin becomes complicated and modified by its simultaneous effect upon the nerve branches and upon the central nervous system. The result is thus much more complicated than that of the action of light is where essentially only the effect upon the skin and a simple structure. for instance. Galvanotropism is. At those points where the current curves enter the influences cells^ a collection of ions takes place which the chemical reactions. although it is latter. Light acts essentially upon the free surface of the animal. and that they move in the direction either of the positive or of the negative current. in unicellular than in vertebrates. filled Tvith If a galvanic current passed through a trough water. a dis- tinct galvanotropism found more often in organisms with organisms. while the electric current affects all the cells and nerves.

" Journ. as well as those of Bancroft indicate that the upon the galvanotropism of these organisms mechanism of these reactions in Volvox is in Volvox is the same and the degree of determinism of the heliotropic and galvanotropic reactions Claparede raises the are governed equally great. electric current because the does not exist in nature. moreover. gations The investi- made by Holmes and myself upon heliotropism. the larvae of the goat moth. XIII. I found. while galvanotropism has no biological significance. And this expectation is fulfilled. and we should expect that galvanic and heliotropic reactions would more nearly resemble one another in this case. however. while the heliotropic reactions by the ''interest of the animal. as The reason whjis may we have seen. be animals shown also that heliotropism just as useless to I many as galvanotropism. Such a view. "^ facts. which mider the bark of trees. provided that we work with organisms possessing both forms of sensitiveness. led to suppose that galvanotropism and heli- otropism are not comparable. while the electric current influences cells of the the body. "Les tropismes devant la psychologic. Cuma Rathkii. The algae of the species Volvox show heliotropism and galvanotropism. the disturbing comfrom this latter circumstance disappear for plications arising the most part when we work with unicellular organisms. which lives in the mud of 1 Claparede. objection that the galvanotropic reactions are purely compulsory. is It can. /. .Significance of Tropisms for Psychology reactions of animals are determined selection could hardly be found. heli- however. Psychologie und Neurologie. same category with the exception of that light acts generally only upon the all surface of the skin. 51 by their needs or by natural One might be of fact. as a matter phenomena of the the aforementioned fact. pointed out twenty years live in the light live ago that some varieties of animals which do not at all. 150. that the crab. however. otropism is not supported by the occasionally. As already mentioned. They are. for instance. For instance. be of use. may show positive heliotropism. 1908.

I believe best suited that this is also a suggestion forced by the extreme application I have made experiof the theor}' of natural selection. according to which the animal is to be looked upon of in- as an aggregate dependent hereditary Arrangement to prove that posiFig. but. free ourselves at once from the overvaluation selection of natural and accept of of the consequences theory Mendel's heredity. therefore.52 The Mechanistic Conception of Life when brought into the Ught It the harbor at Kiel. 21. Only the dayUght D can reach this part of the table. tively hello tropic animals move toward the source of light even if by so doing they go from the simis & window through light into the shade. but instead of remaining at b they keep on moving from the direct svmhght into the shade toward the sovu"ce of light until they all reach the end of the tube c near the — qualities.. which smihght S falls into the room. By a piece of board d e the sunlight 5 is prevented from striking the region 6 c of a table near the window and this part of the table is in the shade. The animals move at once toward the window. from the mud shows positive heUotropism. there- fore. with a clear upon the investigators . test-tube a c is put on this table at right angles to the plane of the window. the winged aphides) are all at a.g. At the beginning of the experiment the animals (e. We must. WW VII The attempt has been made to prove that A organisms are attuned to a certain intensity of light and window nently. (in the shade) where they remain perma- so regulate is their heliotropism that they invariably reach that intensity of light which to their well-being. just as incorrect to assert that the hehotropic reactions are governed by the bio^ ' logical interests of the animal as that this is true for galvanotropism. ments upon a large number of animals. and removed is.

immediately on in the direction of the source of light.^ have never observed a selection" of a suitable intensity of light. heliotropic larvae For these experiments the larvae should be placed only in a very shallow depth of sea-water. which very rich in ultra-violet rays. has been sho\\Ti that positively heliotropic animals are positive to any intensity of light above the threshold. from the sky or weak lamp provided that the (threshold) value of the intensity of light required for the reaction is exceeded. which have been made heliotropic is by acids. What probably lies behind these interpretations of the is "selection of a suitable intensity of light" the fact that under certain conditions reaction products formed by the photonewly hatched (of chemical action of light I may inhibit the positive heliotropism. If they are placed in the light of a quartz mercury lamp is Heraus). with the larvae of Polygordius. sometimes possible to cause positively heliotropic animals to heliotropic. this sort in the found a very clear instance of larvae of Balanus perforatus. I have been able to show that positively heliotropic animals also light move toward I the source of even if the arrangement is such that by so doing they go from the *' light into the shadow. .Significance of Tropisms for Psychology arrangement of the physical conditions of the experiment. The reader will find a further account of this experiment in my book on The Dynamics of Living Matter. In animals sensitive to differences (see next chapter) a stopping occurs in this experiment in the passing from the light into the shadow. Even in a strong light which is not so rich in ultra-violet it is rays as the light of the mercury lamp. which are positively heliotropic. But would be wrong in this case to an adaptation of the animal to a certain intensity of light. become negatively speak of This is it the case. the positively soon become negatively heliotropic. Indeed. nevertheless. for instance. but they go. Thus winged plant lice or wingless larvae of Chrysorrhoea or copepods. 1 Quite often without even stopping for a moment. 53 I have never found a In every case it single indication of such an adaptation. go toward the light whether the source of light reflected light the direct sunlight or light.

in 1 (''Unterschiedsempfindlichkeit") sensi- order to distinguish them from tropisms. of There is a second class phenomena which is determined by a sudden change in the rate of chemical reactions in the tions to a clearly in If same surface elements.54 The Mechanistic Conception of Life is In my opinion it alters merely a case where a metabolic product either the photochemical action or so influences the central nervous system that the excitation of the retina by the light weakens the tonus of the muscles. and the writers that class of errors which fall victims to we have already pointed out in light. because they used a hollow prism filled with ink in order to produce a gradual In the semidarkness thus pro- decrease in the light intensity. " Ueber die Umwandlung positiv heliotropischer Tiere." Bull." Pflugers See also the recent investigations of Georges Bohn. mals gather chiefly in parts of the space where the intensity of light is relatively small. whose are exposed to light. gills Reac- sudden change in the intensity of light are sho^vn most marine tube-worms. for instance. Paris. instead of strengthening it. with planarians. A sudden increase in the intensity of light has no such effect. La naissance de V intelligence. . speaking of the influence of lesser intensities of VIII Heliotropic phenomena are determined by the relative rates of chemical reactions occurring simultaneously in sym- metrical surface elements of an animal. Archiv.. "Les essais et les erreurs ctiez les etioles de mer. is the intensity of the light in the aquarium suddenly dimin- ished the worms withdraw quickly into their tubes. duced.w. u.s. I have designated such reactions as differential the expression of sensitiveness to changes in the intensity of a •stimulus bility. 1893. 1909. a sudden decrease in the intensity of the light causes a decrease in movement. With other Such ani- forms. the intensity of light often remains beneath or near the threshold of stimulation.^ Loeb. for instance. Some of the other mistakes have perhaps also arisen because the writers worked with complicated experimental conditions instead of with simple ones.

The Behavior of Lower Organisms. Paris. d.^ Inst. return to the former direction of moveit. 1906. but deviate sideways from to movement. psychol. franc. "Intervention des reactions oscillatoires dans les tropismes." 1911. 3 * Paris. can complicate and entirely obscure the heliotropic phe- nomena. 1900. " Bibliotheque de philosophic contemporaine. . and like the tubeworms they thereupon draw back very quickly. New York and London. they do not. But Jennings^ has with on such organisms to argue against the theory of tropisms. "Bibliothequedephilosophiescientifique. easily when they are very marked.Significance of Tropisms for Psychology 55 It is hardly necessary to point out here that the effects of rapid changes in intensity. 1 Jennings. and it is therefore easy understand that such animals do not furnish the best mademonstrating the laws of heliotropism. "associative reader is referred to my book^ as well as to the more recent works of Bohn. gen." 1909. Since their locomotor organs are not symmetrical. especially special preference used observations terial for since they possess only a slight photochemical sensitiveness. as the action of a constant current in muscles different Just from that of and nerves is an intermittent current. The upon the call reactions. but are arranged in a peculiar unsymmetrical manner. 1907. analogous case in the action of If we wish to trace all animal reactions back to physico-chemical laws we must take into consideration besides the tropisms not only the facts of the differential sensibility but also all other facts which exert an influence which we gory. sensibility In Hypotricha and other infusoria this differential is very pronounced in response to sudden touch or sudden alteration of the chemical medium. 2 Comparative Physiology of the Brain and Comparative Psychology." Ass. after the next progressive ment. La naissance de V intelligence^ and La nouvelle Let us bear in mind that "ideas" also psychologie animale. 1907.. The influence of that mechanism memory" also belongs in this catebut we cannot discuss this further at this place. Sciences. so we find an light.

a reaction takes place Every enlargement of the latter inat the surfaces of contact. instance. If in the geotropically sensitive elements two masses of different specific gravity are present. creases the mass of reacting molecules. Finally. because in the former the specifically heavier substances are directed toward the center of the stem. two fluid substances which are not at solid or not easily. of the reacting surfaces If it is formed the such geo- assumed that (for in tropically sensitive cells two phases all. the cells of the upper side of a stem which is laid horizontally will ac- quire a different rate of reaction from those of the lower side. flowing sap in the center or the periphery of the stem. or one and one fluid substance) of different specific gravities are which react upon one another. can much as acids do for the hehotropism of certain animals. a third possibility remains which could perhaps be realized in plant roots and stems.^ In the frog's egg. namely. IX Besides light and the electric current. only one of which reacts to the present. hence the geo tropic bending. A shifting of the surfaces would act in the same manner. we can actually demon- strate directly the existence of 1 two substances of Living Matter. one side will grow faster than the other. to increase the sensitiveness to certain stimuli. of different Chapter on "Tropisms" in Dynamics . but it has been suggested that an enlargement or shifting essential connecting link. while in the latter the specifically lighter ones are directed toward the center. The majority of such animals are forced to turn their heads away from the center of the earth and to creep upward. It was uncertain for a long time how the orientation of cells in relation to the center of gravity of the earth could influence the rate of the chemical reactions within. and thus can lead to tropism-like movements or actions directed toward a goal. miscible. Con- sequently. the force of gravity also has an orienting influence upon a number of animals.56 The Mechanistic Conception of Life act.

since in this case they are of different In animals has been observed that orientation toward the center of gravity of the earth often becomes less compulsory when the inner ear has been removed. Lyon carried out these experiments. indeed. it their behavior. no right to maintain that the orientation of is animals in relation to the center of gravity of the earth regu- by the pressure but that this selves. which showed that no disturbance of the orientation resulted from this operation. large otolith that can easily be removed from the ear. The problem was solved by experiments on flounders. regulation takes place in the nerve endings themof the otoliths of and probably. therefore. upon the nerve endings. Mach first pointed this. We may conclude. We have. out the possibility that the otoliths are responsible for He believed that they might press upon the end-organs It is of the sensory nerves and every change of pressure might cause a correction of the position of the animal. Through the change of orientation of the cells in relation to the center of gravity of the earth. because in so doing the nerve endings in the ears were injured. which have only a single E. that in my experiments of scraping out the otoliths a disturbance of the orientation occurred. but if the otoliths are simply washed out from the of sea-water the orientation does inner ear by a weak current suffer. the two . P. as a result of the existence there two different phases of different specific gravity which react upon one another.Significance of Tropisms for Psychology specific gravity 57 and can study color. generally assumed I that this view has been verified by experiment but entirely agree with it cannot although I once described experiments I which seemed to support Mach's that otolith theory. it is not so easily In the latter case. had found when the otoliths of the inner ear of the shark are scraped out with a sharp spoon the normal orientation of the animal suffers. lated therefore. doubtful whether all the otolith powder has been removed from the ear.

and F. these writers ought to base the law of tion of the existence of the molecules and ions.58 The Mechanistic Conception of Life phases undergo a shifting by means of which a change in the rate of reaction is brought about according to one of the ways Since then I have looked through the htera- described above. tend toward progress to assert that this is a case of functions of more than two variables. tendency has lately been gaining influence." I believe cells of plants and France even attributes to the latter a ''soul" and that in order to be consistent. Haberlandt. without furnishing sufficient proof for this assertion. no dis- turbance of the orientation occurred. size mass action upon the assumpsense-organs. souls. of deriving the tropic reactions of plants as directly as possible from the law of mass action or the law of Bunsen and Roscoe. in In the only case which a removal of the otoliths without tearing or other injury of the nerve endings can be justifiably assumed. While in plants my own finally work I have aimed to trace the complex laws. in my opinion. Darwin. . and have reached the conclusion that all writers who assert that the removal of the otoliths disturbs the geotropic orientation of animals have been victims of the same fallacy as myself. Nemec. Some namely. presentation of a "explanation" consists solely in the as phenomenon it is an unequivocal function of if the variables by which determined. and intelligence in It is probably unnecessary to emphato the fact that it is better for the progress of science to derive the more complex phenomena from simpler components than For all do the contrary. endeavor to show that the relatively simpler reactions of plants may be traced Instead back to the more complex relations found in animals. ture on the function of the otoliths or statoliths. reactions of animals back to simpler reactions like those of and to physico-chemical the opposite botanists. and it in nature we find a function of two variables. They have injured or removed the nerve endings. does not. they try to show that ''sense-organs" exist in the "intelligence.

Mendel's laws have been brilliantly In some cases of apparent it confirmed in a number of cases. they know that an apparent exception does not necessarily overthrow the law. an unnecessary complication of simple relations. my opinion.Significance of Tropisms for Psychology 59 These writers explain the geotropic reactions of plants by saying that in certain cells starch grains are present which serve the purpose of the otoliths in animals. grouped according to their attitude toward such simple laws or rationalistic elements. When is such investigators have discovered a simple law which generally applicable. I is then supposed to give rise to the geotropic curva- have no opposition to offer to the assumption that the starch grains change their position with a change in the position of the cells. find that there are We two classes of investigators in biology. One seems to aim at the denial of the existence of such simple laws and every new case which does not for fall at once imder such a law offers an opportunity them to point out the inadequacy of the latter. But no necessity for assuming besides this in the existence of intracellular sense-organs which perceive the pressure of the starch grains. This is. These starch grains are believed to press upon the sense-organs or nerve endings in the plant cells concerned and the ''pressure-sense" of the plant ture. X The progress of natural science depends upon the dis- covery of rationalistic elements or simple natural laws. has not always been One group of investi- gators has recognized that these deviations do not indicate the incorrectness of Mendel's laws. but that they are merely the . deviations (from these laws). but that possibly an opportunity is offered for a new discovery and an extension of the old law. however. possible at once to recognize the cause. The other group of investigators aims to discover and not to disprove laws. and I I am see also willing to pass over for the present the view that the starch grains form one of the two phases in the cell.

Jemiings draws from this fact the following conclusions: is ''The symmetrical position its an incident of the reaction.60 The Mechanistic Conception of Life and often minor complications. however. therefore. Tropisms and tropism-like reactions are elements which pave the way for a rationalistic conception of the psychological reactions of animals and I believe. The case is similar in the realm of tropisms. comes about as we have already described. to assert that is not permissible to speak of tropisms as being governed in other words. the latter result of secondary investigators have from this standpoint discoveries. Consequently the heliotropic orientation. such as the fiddler crab. often The fact that in an electric current the same animal moves differently from what it does under the influence of light finds its explanation for the observer conversant with physical chemistry in the fact that the electric current causes changes in the concentration of ions within. however. not essence. for instance. as well as upon the surface. certain crabs. he uses these observations of Holmes to . in an attempt to minimize the importance of Mendel's laws and thus to retard the progress of science. primarily. leave this difference of consideration in the action of the two agents out and make it use of the difference in the behavior of certain organisms in response to light and to the electric current. made further fruitful The role of the other group of investigators in this case has consisted. that it is in the interest of the progress of science to develop further the theory of animal tropisms. There are animals. Certain A\Titers. for instance. Holmes has found that these crustaceans also go sideways toward the light. while the chemical action of light is essentially limited to the surface. which move sideways. Animals in general are and the motor elements of the right and left sides of the body usually act symmetrically. by general laws. built they say that tropisms are symmetrically without significance." In other words.

however. as. but also chemically.Significance of Tropisms for Psychology indicate that the role ascribed to for the theory of tropisms. secondly. furthermore. however. and that. for in- stance. I expect. application of the law of mass action to this set of reactions The is thus made *' possible. To me it is a question of making the facts of psychology accessible In this way it is to analysis by means of physical chemistry. that they will only confirm still more the laws of heliotropism. that in the fiddler crabs in the first place there retina is an entirely different connection between the and the locomotor muscles from that in other crustaceans." but am of the opinion that the contents of comparative psychology will under the influence of the above-mentioned endeavors be different from the contents of speculative psychology. is This orientation automatically regulated according to the law of mass action. I 61 symmetry has no importance am. namely. or chemical substances. are obliged to orient their bodies in a certain way in relation to certain centers of force.^ XI These data may suffice to explain my point of view. However. the course of light. I believe that a new discovery may be made here. which I cannot. the center of gravity of the earth. . their Many animals. already possible to reduce a set of reactions. because body structure is not only morphologically. the tro- pisms to simple rationalistic relations. inclined to draw another conclusion. probably no one will maintain that the existence of the pleiironectids invalidates all laws in regard to the symmetrical body structure. symmetrical. This expectation is based upon analogous relations in the pleiironectids. for it is in general hardly 1 From which. But I believe also fall that the further development of this subject will more to the lot of biologists trained in phj^sical chemistry than to the specialists in psychology or zoology. discuss further here. I consider it unnecessary to give up I the term comparative psychology. there is a special peculiarity in regard to the function of the two retinae whereby they do not act like symmetrical surface elements. namely. an electric current.

for instance. we have given. the con- dition that human beings are willing to sacrifice their lives for an idea comprehensible neither from the utilitarian standIt point nor from that of the categorical imperative. within which the analogies necessary for psychiatry can be called forth experimentally and can be investigated. These experiments ethics. I believe that the investigation of the conditions which produce tropisms may be of importance for psychiatry. it .62 The Mechanistic Conception of Life and psychologists who lack a to be expected that zoologists physico-chemical training will feel attracted to the subject of tropisms. In closing let me add a few remarks concerning the possible application of the investigations of tropisms. in the dog by means and acoustic signals. might be possible that under the influence of certain ideas chemical changes. may also attain a similar value for The is highest manifestation of ethics. Since Pawlow have succeeded of optic in causing the secretion of saliva no longer seems strange to us that what the philosopher terms an ''idea'* is a process which can cause chemical changes in the body. are produced which increase the sensitiveness to certain stimuli to such an unusual degree that such people become slaves to certain stimuli just as the copepods become slaves to the light when carbon and his pupils dioxide is added to the water. I believe. internal secretions within the body. If we by can call forth in an animal otherwise indifferent to light means of an acid a heliotropism which drives if it irresistibly into a flame. the same thing can be brought about by means of a secretion of the reproductive glands. a group of facts. namely.

III. SOME FUNDAMENTAL FACTS AND CONCEPTIONS CONCERNING THE COMPARATIVE PHYSIOLOGY OF THE CENTRAL NERVOUS SYSTEM .

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If we ask what the elementary com1. By J. example. reflex. upon an ponents are in the physiology of the central nervous system. destruction of the spinal cord would. is called a There has been a growing tendency in physiologj^ to make reflexes the basis of the analysis of the functions of the central nervous system. and consequently much importance has been attached to the processes underlying them and the mechanisms necessary for reflex. In each of these cases. Putnam's Sons of New York and London. 1 make the reflex impossible. 65 . is directed to a class of processes which are called A reflex is a reaction which is caused by an external stimulus. and back again to the peripheral muscles. when the conjuctiva touched by a foreign body. according to this.. tive Psychology (1899). the is closing of the eyelid. and which results in a coordinated for movement. Comparative Physiology of the Brain and Comparacourtesy of G. Reprinted from Loeb. This passage from the stimulated part to the central nervous system. producing there a contraction. The name reflected reflex suggests a comparison between the spinal cord and a mirror.Ill SOME FUNDAMENTAL FACTS AND CONCEPTIONS CONCERNING THE COMPARATIVE PHYSIOLOGY OF THE CENTRAL NERVOUS SYSTEM i The understanding of complicated phenomena depends analysis by which they are resolved into their simple elementary components. P. Sensory stimuli were supposed to be from the spinal cord to the muscles. or the narrowing of the pupil under the influence of light. passes from there to the motor nerves. This change travels to the central nervous system. changes in the sensory nerve endings are produced which bring about a change of condition in the nerves. our attention reflexes. and terminates in the muscle-fibers.

however. A prominent psychologist The ganglion- has maintained that reflexes are to be considered as the mechanical effects of acts of volition of past generations. This comparison. merely as conductors.66 The Mechanistic Conception of Life might prevent the reflection just as the destruction of the mirror of Hght. of strong light. its however. in such reflexes has also been The movements which it are produced are so well intelli- planned and coordinated that seems as though some gence were at work either in devising or in carrying them out.^ cell seems the only place where such mechanical It effects could be stored up. the purposeful character reflex of many movements. the nerve-fibers being regarded. for the cornea is thereby protected from hurtful contact with foreign bodies. The closing of the eyelid and the narrowing of the pupil are eminentlj' purposeful. Both the authors who emphasize the purposefulness of the reflex act and those who see in it only a physical process have invariably looked upon the ganglion-cell as the principal bearer in of the structures for the reflex action. of the reflex process in the central nervous system with the reflection of light has. become meaningless. another feature in the conception of the term reflex has gained prominence. The with fact. complex coordinated movements inclined as any other physiologist to doubt the correctness of this conception had not the establishment of the identity of the reactions of animals and plants to light proved the untenability of this view and at the I should have been as little 1 A statement for which no trace of experimental proof exists. and at present few physiologists in using the term reflex think of its original significance. . that even a decapitated frog its skin. long since. In- stead of this. essential has therefore been considered the most element of the reflex mechanism. will brush foot a drop of acetic acid from is suggests that some other explanation necessary. namely. and probably correctly. effects and the retina from the injurious Another striking characteristic emphasized.

and the moth is caused to the flame. body and the muscles. Wurzburg. 1 A further objection has been raised. and from there to the muscles fly into the wings. A preliminary note on these experiments appeared January. If the ganglion-cells or the central nervous system. . This argument. 1890. as in the case of plants which possess no nerves. 67 flight The moth into the flame is a typical reflex process. is not sound. that although these Loeb. this identity of animal with plant heliotropism can these heliotropic effects must depend upon conditions which are common to both animals and plants. for the nervous reflex arc in higher animals forms the only protoplasmic bridge between the sensory organs of the surface of the body and the muscles. we destroy we interrupt no longer the continuity of the protoplasmic conduction between the surface of the possible. 1888. unless we find that the phenomena cannot be explained by means of the latter alone. in the case of animals which possess nerves." On the other hand. and depend in the same way upon the external form of the body. The of light stimulates the peripheral sense-organs. ganglia. ganglion-cells are nothing 2. consequently. the stimulus passes to the central nervous system. cannot depend upon force but one inference specific qualities of — the central nervous system. however.. This reflex process agrees in every point ^vith the heliotropic effects Since plants possess no nerves and no of light on plant organs. the objection has been raised that destruction of the ganglion-cells interrupts the reflex process. we are justified in seeking in them only general protoplasmic qualities. J. At the end of my book on heUotropism^ I expressed this view in the following words: ''We have seen that.Physiology of Central Nervous System same time of the offered a different conception of reflexes. Der Heliotropismus der Tiere und seine Uebereinstimmung mit dem Heliotropismus der Pflanzen. the movements of orientation toward light are governed by exactly the same external conditions. These heliotropic phenomena. and a cylinders of reflex is Since the axis the nerves and the more than protoplasmic formations.

yet in reflexes occur in plants possessing animals where ganglion-cells are present the very existence of the ganglion-cells necessitates the presence in reflex them of special if mechanisms. This is a reflex comparable to the closing of the eyelid if we touch the cornea. and the arrangement of character of the reflex act. all and conductibility are the only qualities and these are both common qualities of irritable structures at the surface of protoplasm. 2 This animal closes the oral opening when we touch it. the 1 majority of investigators explain in this way only a II. for instance. While some authors explain J. ail reflexes on a psychical basis. Wurzburg. Their value the fact that they are quicker and more sensitive conductors than undifferentiated protoplasm.^ A study. in Such a phenomenon could be expected only in forms which a direct transmission of stimuli from the skin to the is muscle or direct stimulation of the muscle possible. lies in to conclude that the nerves are superfluous and a waste Certainly not. then. 1892. 3. The the body. Such power of adaptation absolutely necessary for free animals. When the latter is removed the oral opening still closes upon mechanical stimulation. case. the muscles determine the the central The assumption that But have we now ? nervous system or the ganglion-cells are the reflex specific bearers of mechanisms cannot hold. Loeb. in addi- tion to the transmission through the reflex arc. of comparative physiology brings out the fact that irritability essential to reflexes.68 The Mechanistic Conception of Life no nervous system. Untersuchungen zur physiologischen Morphologic der Tiere. I succeeded^ in demonstrating in Ciona still intestinalis that the complicated reflexes continue after removal of the central nervous system. in This is the worms and in ascidians. The central nervous sytem of the animal consists of one ganglion. . It was therefore necessary to find out still there w^ere not animals in which coordinated reflexes con- tinued to exist after the destruction of the central nervous system.. Because of these qualities of the nerves. itself an animal it is better able to adapt if it to changing conditions than possibly could is had no nerves.

To this class of reflexes belongs the habit possessed by certain insects of laying their eggs on the material which the larvae will afterward require for food. and . chemotropism. the tropisms (heliotropism. These circumof stimu- stances force an animal to orient lation in such a of the itself toward a source In this way that symmetrical points on the surface body are stimulated equally. How can the action fly of such an insect be determined if not by mysterious structures ? which can only be contained in the ganglion-cells explain the inheritance of such instincts if How can we it we believe to be a fact that the ganglion-cells are only the conductors of stimuli ? It was impossible either to develop a mechanics of instincts or to explain their inheritance in a simple point. geotropism. Those nearer the oral pole possess greater than that of those near the aboral pole. way from the old stand- but our conception makes an explanation possible. upon the relations of body. upon the specific irritability of certain elements of the bodyotropism) play an important part. way the animals of are led without will of their own it. Among the elements which compose these complicated instincts. Symmetrical elements at the surface symmetry of the of the body have an irritability the same irritability. we cannot cease to wonder at the seeming care which nature takes for the preservation of the species. surface. second. unsymmetrical elements have a different irritabiUty. and. but no matter is how the definition phrased the meaning seems to be that they are inherited reflexes so purposeful and so complicated in character that nothing short of intelligence and experience could have produced them. Instincts are defined in various ways. When we consider that the female pays no attention to her eggs after laying them. stere- These tropisms are identical The explanation of them depends first for animals and plants. either toward the source the stimulus or away from for the ganglion-cell to Thus there remains nothing do but to conduct the stimulus.Physiology of Central Nervous System certain group of reflexes 69 —the so-called instincts.

For the inheritance of instincts only necessary that the egg will contain certain substances tropisms of the —which determine the different symmetry —and the conditions for producing bilateral embryo. to their unknowable structures giving in the ganglion-cells served formerly as all a convenient terminus for thought in this direction. especially in the more complicated ones. and the significance of the ganglion-cells confined. We find that authors prefer to speak of reflexes in cases where the reaction of single parts or organs of an animal to external stimuli is concerned. that a stretched muscle shows an increased Since irritability while the contracted muscle shows a decreased irritability. The assumption of complicated but unkno\\Ti and perhaps reflex processes. The mystery with which the ganglion-cell has been surroimded led not only to no definite insight into these processes.70 this The Mechanistic Conception of Life may be accomplished by protoplasm it is in any form. but has proved rather a hindrance in the attempt to them. If the mechanics of a of the tropisms number of instincts is explained by means if common to animals is and plants. find the explanation of It is evident that there is no sharp line of demarkation between reflexes and instincts. while they speak of instincts where the reaction of the animal as a whole is involved (as is the case in tropisms). 4. we upon to show what conditions are able to determine the coordinated character of reflex movements.^ 1 Since this was written von UexkueU found a law which will go far in explaining the mechanism of coordination. . This perhaps creates a rational basis for the further investigation of coordinated movements. we are what circumstances determine the coordinated movements in reflexes. Experiments on galvanotropism of animals suggest that a simple relation of certain may exist between the orientation nervous system and is motor elements in the central the direction of the forth movements of the body which called by the activity of these elements. are called In up this assumption. namely. as in all forced to ask power of conducting stimuli.

71 We must also deprive the ganglion-cells of all specific significance in spontaneous movements. Recent investiga- however. presence of Ca ions in the blood which prevents the muscles of our skeleton from beating rhythmically it is is in our body. ions be increased If in the muscles of the skeleton the Na and the Ca ions be reduced. no are apparently deterStrictly internal conditions of the living system. K. By spontaneous movements we mean movements which mined by speaking. the muscles are It is only the able to contract rhythmically Hke the heart. Respiration and the heart beat belong in this category. namely. and from this the con- clusion has been drawn that automatic movements are due to specific structures of the ganglion-cells. tions. for atmospheric oxygen and a certain range of temperature are always necessary in order to preserve the activity beyond a short period of time. aperiodic spontaneous processes and rhythmically spontaneous or automatic processes. .Physiology of Central Nervous System 5. in definite proportions. Ca. As the power of muscles contain no ganglion-cells. We mical processes are of importance for our consideration. and others) portions. must discriminate between simple and conscious sponIn simple spontaneity we must consider two kinds of processes. certain that the rhythmical spontaneous contractions not due to the specific the contraction of one group of muscles necessitates the stretching of their antagonists the coordinated character of locomotive action seems to become intelligible (1912). have transferred the problem of rhythmical field of spontaneous contractions from the of physical chemistry. respiratory The movements seem all to indicate that automatic activity can arise in the ganglion-cells. just as we have done in the case of simple reflexes and instincts. By changing these proit we can impart to a tissue properties which does not ordinarily possess. morphology into that The pecuHar it qualities of each tissue are partly due to the fact that contains certain ions (Na. The rhythtaneity. movements of animals are exclusively determined by internal conditions.

. for the tropisms depend 1 Loeb. Thus far we have not touched upon the most important namely. Some physiologists and psychologists consider the purposefulness of the psychic action If as the essential element. show that the coordination of automatic movements is caused by the fact that that a specific element which beats most quickly forces the others to beat in its own rh^'thm. Plants must therefore have a psychic life. following the argument. which possess no ganglion-cells. Observations in lower animals. we must ascribe it to machines also. are looked upon as psychic functions. American Journal of Physiology. We must find out the ele- mentary physiological processes which underlie the complicated phenomena of consciousness. Aperiodic spontaneity is still less function of the ganglion-cell than rhythmical spontaneity. spores of algae. The show spontaneity equal to that of animals having ganglion-cells.72 The Mechanistic Conception of Life morphological character of the ganglion-cells. and. 327 and 383. the case of phenomena which are called psychic or Our method of procedure must be the same as in instincts and reflexes.^ necessarily confined to The coordinated often been explained tion. J. 1900. We shall see in the follow^ing chapters that many of these reactions are merely tropisms which may occur in exactly the same form in plants. we phenomenon the of In this way many instincts especially. to that complex of conscious. but to definite chemical conditions which are not ganglion-cells. because its many of functions are purposeful. swarm 6." character of automatic movements has by the assumption see that they of a '^ center of coordina- which is supposed to keep a kind of police watch on the different elements and move in the right order. Conscious- ness has been ascribed even to the spinal cord. these authors declare that consciousness. rational an animal or an organ reacts as a are dealing with a reflexes. . however. which mechanisms give rise problem in physiology. III. human being would do under the same circumstances.

physiologists who have appreciated the untenable character of such metaphysical speculations have held that the only alternative is to drop the search for the mechanisms underlying consciousness and study exclusively the This would be throwing results of operations on the brain. Consciousness are is only a metaphysical that mechanism by effects term for phenomena which determined by associative I memory. or of association. according to my opinion. the errors of the metaphysician must be avoided and explanations must rest upon facts. namely. This process. If brain physiology gives up its fundamental best possi- problem. We can dispose of this view by the mere fact that the phenomena of embryological development and of organization in general show a degree of purposefulness which we would arrive at molecules may even surpass that of any reflex or instinctive or conscious act. 73 In the last analysis. but that they employ the wrong methods of investigation and substitute a play on words for an explanation by means of facts. The mistake made by metaphysicians is not that they devote themselves to fundamental problems. then. out the ^^heat with the chaff. the discovery of those elementary processes which make consciousness bilities. it abandons its But to obtain results. and atoms endowed with mental qualities. It should consist in the right understanding of the fundamental process which recurs in all psychic phenomena as the elemental component. not words. By associative memory mean which a stimulus brings about not only the but by which which its nature and the specific structure of the irritable organ it call for. The method should be the same for animal psj'chology that it is for brain physiology. On the other hand. brings about also the effects of other stimuli which formerly acted upon the organism almost or quite . is the activity of the associative rnemory. possible.Physiology of Central Nervous System only on simple mechanical arrangements. And yet we do not consider the phenomena of development to be dependent upon consciousness.

LVI. all Our criterion puts an end to the metaphysical ideas that matter. and present in these only after a certain development has been reached. 7. We are brought to the theory that only certain species of animals possess associative sciousness. . ^ an animal can be trained. developed the shark or frog) do not or differ but slightly. "Beitrage zur Gehirnphysiologie der Wiirmer. support this view. The fact that only certain animals possess the necessary mechanical arrangements for associative memory. at all or only slightly differ... in their reactions after losing the cerebral hemispheres. 1894. and worms do not possess a trace of associative memory. Coelenterates. and which do not possess. It is a comparatively easy task to find out which representatives of the various classes of animals possess. 247. associative memory.g. 1 The J. ants.74 The Mechanistic Conception of Life If simultaneously with the stimulus in question. if it can learn. bees. and hence the whole animal world. the existence of associative memory can be proved. possesses con- sciousness. liquefaction of gases an example sudden Loeb." PflUgers Archiv. Our criterion therefore might be of great assistance in the development of comparative psychology. criterion it possesses associative memory. and therefore for consciousness. is not stranger than the fact that only certain animals possess the mechanical arrangements for uniting the rays from a luminous point is in one point on the of a retina. The fact that certain vertebrates lose all after the destruction of the cere- power of associative memory bral hemispheres. memory and have con- and that it appears in them only after they have reached a certain stage in their ontogenetic development. and the fact that vertebrates in which the associative memory either is not developed (e. Among certain classes of insects (for instance. This is apparent from the fact that associative memory depends upon mechanical arrangements which are present only in certain animals. By means of this it can be shown that Infusoria. and wasps).

Which peculiarities make the phenomenon of assoFor the solution of this memory possible? problem the experience of physical chemistry and of the physiology of the protoplasm must be combined. the process of conduction of brain stimuli. and when we consider It so many variables subject to that colloids easily change their state of matter. namely. But same or time it is evident that this mechanism cannot be unraveled by or histological methods. We have to remember that by all life phenomena are ultimately due to motions or changes occurring in colloidal substances.Physiology of Central Nervous System change of condition which is 75 may be produced when one variable changed. by operations on the brain. . it is not surprising that there should be sudden changes in the ontogenetic and phylogenetic of organisms development when there are change. becomes evident that the unraveling of the mechanism of associative field of memory is the great discovery to be made at the in the brain physiology and psychology. of the colloidal substances can ciative The question is. measuring reaction times. From the same sources we must of expect the solution of the other fundamental problems physiology.

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IV.PATTERN ADAPTATION OF FISHES AND THE MECHANISM OF VISION . .

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Minkowski was able to confirm Henschen's conclusions through experi lents on dogs. Henschen has recently. 79 . No. The image possibility that vision is is based on the formation of an in the brain supported by a group of facts which to in this my knowledge have never received any consideration connection. furnished the proof. 1912. on the basis of excellent pathological observations on man. From results obtained in experiments on dogs Munk stated years ago that there existed a projection of the retina on a part of the cortex which he had designated as the visual sphere and I that the extirpation of definite parts of this sphere caused blindness in definite parts of the retina. XXV. 1 Reprinted from Physiologisches Centralblatt.IV PATTERN ADAPTATION OF FISHES AND THE MECHANISM OF VISION^ The mechanism of the action of the brain is entirely unknown to us. in the area Munk had believed it to be. This note is given merely as a suggestion concerning the mechanism underlying certain brain processes. cortex. that such situated in another a projection after all exists. 22. We are unable to look into the active brain and the objective action that results of brain action are in general so differ- ent in their nature from the external stimulus which leads to the we are prevented in most cases from drawing any conclusions concerning the nature of the processes occurring in the brain. but that it is part of the cortex from where namely. however. repeated these his experiments but was not able to confirm statements. These observations and experiments suggest the possibility that in vision an image is formed not only on the retina but also on the striata.

the transmission of the retinal image upon the skin takes place through the follows that the image optic nerve. Pouchet many years ago showed that the adaptation of fishes to the ground ceases as soon as their eyes are tion of retinal images refractive is removed or as soon as the forma- prevented through the turbidity of the This fact (confirmed by color is media of the eye. An image consists of a number of points of different intensity of light. cutting of and the optic ganglia in the brain acts like the extirand finally it has been proved that the the sympathetic fibers which go to the pigment cells skin. that the so-called adaptation of fishes to their surroundings mission of the retinal image to the skin. fish is an image of the object on the retina since it is proved that the image on the skin of the since. many observers) proves that the harmony between and pattern transmitted of the skin of fishes with their surroundings through the retinal image. is only the trans- moreover. adapt their color and pattern to the ground upon which they happen to be. for of which is definite and charSumner has shown that certain fishes are able to reproduce on their skin rather complicated . It seems to me that the same facts furnish also the proof that an image of the objects is formed in the brain. of the skin also prevents the formation of a picture of the ground on the retinal is image is transferred to the skin of fishes. Hence we know the path by which the One station Since the ending of the optic fibers in the brain. in other words.80 It The Mechanistic Conception of Life has been some time that many animals. been shown that the destruction of the optic fibers pation of the eyes. moreover. it must pass the central stations of the optic nerve during the transmission to the skin. It has. we are able to prove the existence of of fishes. This fact has been for known extensively utilized for the theory of natural selection. a picture of the retinal image but not of the object (in this case the ground) itself. the acteristic mutual arrangement the object. especially certain fishes.

which form the bottom of the is aquarium. we cannot well imagine the transmission of the retinal image upon the skin in any other way than by assuming that the relative arrangement of the individual points of the retinal image is preserved in the optic fibers and the end ganglia of the optic nerve. .. Under this assumption a relative distribution of the stimulating intensities must occur in the primary optic ganglion which corresponds to the distribution of the image points on the retina and which again can be called an image. 81 a chess board). We may consider each point of the retinal image as a luminous or a stimulating point which produces a corresponding image point in the fiber primary optic ganglia through the action of the nerveit is through which connected with the ganglia. therefore. definite arrangement of the images of the different luminous points of the ground on the retina and a similar arrangement of the images of the luminous points on the skin of the fishes. Every image point in the primary optic ganglia may be considered again as a luminous or stimulating point which through the mediation of a special nerve-fiber influences an individual chromatophore or a small group of chromatophores of the skin. These observations in fish and the conclusions drawn note suggest the idea that vision is in this a kind of telephotography. This reproduction of the pattern if somewhat imperfect. exists. is Considering the fact that the retina a mosaic.g.Pattern Adaptation of Fishes patterns (e. but we deduct the secondary disturbing factors the is fact remains that the pattern on the skin There a tolerably true a picture of the pattern of the ground.

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V. ON SOME FACTS AND PRINCIPLES OF PHYSIOLOGICAL MORPHOLOGY .

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Antennularia antenFrom a nina. In case this could be done. from the hips legs. Organbildung und AVachsthum. or polyps originating on the under side of a branch. HETEROMORPHOSIS- The various organs of the higher animals have a definite arrangement. which carry polyps on upper sides. In observing these phenomena the question arose shall originate at certain places in the : What are the circumstances which determine that only one kind of organ body? It occurred to me that the answer to this question might be obtained by finding out it were possible to make any grow at any desired place. Wtirzburg. but we never see legs growing out from the shoulders hips. or in place of a branch. the question to be decided was whether the same circumstances by which the arrangement of organs can be changed experimentally also determine the arrangement of first of all whether or not desired organ of an animal 1 Reprinted from Biological Lectvires delivered at the Marine Biological Laboratory of Woods Hole. morphosis. of height to a bundle of roots or stolons a straight stem arises six inches or more. short their From this main stem originate. or arms from the Fig. 85 . by courtesy of Ginn & Co. 1892. which is quite common in the Bay of Naples. from the shoulders arms originate. 1893. Translated in Studies in General Physiology. in regular and slender branches. succession. arrangement of organs 22 gives a diagram of a hydroid. 2 Untersuchungen zur physiologischen Morphologie der Tiere. In the lower animals the same definite exists. II. 1891. In this animal we never find a root originating at the apex.ON SOME FACTS AND PRINCIPLES OF PHYSIOLOGICAL MORPHOLOGY ^ I. HeteroI. Wurzburg.

24. i. is the apical organs end a forms roots W. of heteromorphic regeneration in Antennularia. mentioned. organs in the natural development. The old normal arrangement of thus restored through the process of regeneration. W.86 The Mechanistic Conception of Life \ \/ Fig. In the root this case — end a. The piece forms roots at the lower end b and a new stem »S at the upper Diagram Fig. piece ab (Fig. of the normal stem of Antennularia antennina. end b upward and the stem end a downward. which laria A . A piece a b cut out of the stem is hung up in an inverted position. —A piece solve this problem an attempt to and the following simple experiments were performed. 23) of an Antennuwas cut out and hung up vertically in the water of the aquarium. seemed to afford a suitable subject for experimentation in Fig. S S. It was found that after a few days the root end b had formed little roots. t/ The above Antennularia antennina. 23. Approximately natural size.e. and the basal end b forms a new stem S which grows upward. of normal regeneraif a piece a 6 of is liung vertically in the — Diagram W tion Antennularia up water. stolons or roots.. 22. hydroid. stem. & hydroid of the Bay of Naples. W. the apical end a above and the root end b below.

87 had formed a new A similar piece was cut out from another specimen and was hung upside dowTi in the aquarium (Fig. ^\ . This could be proved by their physiological reactions. e. and these downward growing elements were no longer branches but roots (Fig. for the roots piece of an Antennularia attach themselves to the surface of solid bodies.. From this we see that it was possible to substitute a root for a stem and an apex for a root. grow downward as stolons and attach themselves to solid bodies. b .: Fig.g. Regeneration of a piece a b cut out from the stem of Antennularia and put horizontally into the water. ^ ::^. the glass ... From nature. :n> . .. The root end b.. On the upper side a new stem c d grows vertically upward. W. something still more remarkable occured. the branches which were growing . a. which was below. The branches on the lower side which had ceased to grow. 25). If the excised was placed horizontally instead of vertically in the aquarium. 25. S. *S. and the apical end.Physiological Morphology grew downward. which was now above. formed roots. formed a new stem. namely. stem.„. — upward bore polyps on their upper side. and the apical end a.. In the newly formed stem the arrangement of the organs was the same as in the normal animal. This phenomenon of the substitution of obliquely one organ for another I termed heteromorphosis. namely. 24). the branches on the lower side suddenly began to grow vertically downward.

the stem a b new stems S. they grew toward the center of the earth). with the root If the stem were placed in an oblique end above. 26). solid bodies with which they came geotropic (that Moreover. from every element of the main stem new stems and roots from its roots originated. The branches into on the upper side were not roots. as a rule. but at parts surface of of the the origi(c d. piece o b of a stem of Antennularia put obOn the upper side of liquely into the water. usual. 26. they were positively while the branches never is. not in other forms of Antennularia. while the stems never These new parts growing out from the branches of the under side of the stem attached themselves to the in contact. transformed They either per- ished or gave rise to long. of the aquarium. S„ S. . upper nated Fig.„ grow vertically upward. — new 25).88 The Mechanistic Conception of Life show such a reaction. position. were too slender to bear branches. These stems. the branches on the mider and at the upper end a stem arose as side grew out as roots. If W„ produced little typical branches bearing polyps. we brought the stem into an obhque position (Fig. that stems always originated from the upper side of an element and lower side.„ grow out. but with this difference. while at the lower end of the piece a b opposite the new stems roots W. with the apex a upward. perfectly straight stems. slender. showed any positive geotropism. stems which grew vertically upward the and W. This influence of gravitation is foimd only in Antennularia antennina. which grew vertically upward. main stem there Diagrammatic regeneration in a Fig.

the stem grows vertically upward. which Mr. if not in contact with a solid body.Physiological Morphology 89 What circumstances had all these experiments in common ? The stems always element. but gives a great number of small polyps which appear at the end of long stems. as a root. namely. and place it in a dish from every motion. ask whether the action of this force. Now we may in this form. Even the main root. Fig. a curious change takes place in the organism. drawing of an example at of heteromorphosis a hydroid common Woods Hole. roots to arise at Gravitation does not permit any place except at the under side of the organs. 27 gives a in Margelis. and thus the general arrangement of organs is brought about by gravitation. is also responsible for the natural arrangement of the organs I believe namely. under normal conditions. no longer rise to grows as a root. or a small piece of a stem of this hydroid. originated from the upper end or side of an element. In this way the apical end of a stem may continue to grow as a totally different organ. . But how does gravitation determine that stems grow on the upper and roots at the under side? This is a question to which we shall return later. gravita- tion. that this is the case. Almost all. Every containing sea-water. and roots always from the lower side or end of the same These facts can be explained only on the assump- tion that in this case gravitation determines the place of origin of organs. The same force determines that polyps can originate only on the upper side of branches. it If we cut off a stem. By reason of its negative geotropism. Fig. that roots appear only at the base of the stem and never at the apex or in the place of a branch. upon which another set of experiments was carried on. of the stems which touch the glass give rise to roots that spread out and very soon cover a large area of the glass. protecting carefully organ not in contact with some solid body gives rise to polyps. at the base of the stem. and in some cases all. 27. and that is.

place which was in contact with sea. which was not in contact with the Fig. new roots or stolons grow out. A piece a b of this hydroid was cut out and put into a jar with sea-water. At a and 6. 27.90 The Mechanistic Conception of Life to Tower was kind enough formed roots at its draw for me. Heteromorphosis in Pennaria. with solid bodies gave was in contact rise to roots. rise to roots. Heteromorphosis in Margelis. shows a branch which its apex and polyps at roots in this manner. glass. a hydroid. where the points of stems touch the ground of the aquarium. 28. — The stem touched the bottom a and h. The ends a and b touched the bottom of the jar. All these ends gave From the upper side of the original root. on small polyps grew out. Every place which and every rise to polyps. — At both points new roots grew later out. of the dish with the apical ends. Fig.water gave This is not the only species of hydroid found at Woods Hole .

and have nothing to do with natural selection. as the circumstances that determine the forms of animals and plants are only the different forms of energy in the sense in which this word is used by the physicist. way the contact may have an influence shall be mentioned later on.Physiological Morphology in 91 which such forms of heteromorphosis can be produced. the nature of the contact not only determines the place of origin of the various organs. gravitation not only determines the place of origin of the various organs. that the formation of roots in Pennaria is. if they grow at all. Pennaria. is negativelj' geotropic. is positively geotropic. . although as a rule they do.^ In these experiments on Margelis and Pennaria organs brought into contact wdth solid bodies continue to grow as roots. In Antennularia. one may say that these roots in Pennaria may give rise later on to polyps. In the special case that I observed they did not. absolutely necessary. in which polyps also arise from the roots. We 1 In a Tubularian I was able to produce the opposite result. causal or physiological morphology would not gain thereby. In Pennaria I succeeded repeatedly in producing roots at both ends of a small stem that bore no polyps (Fig. that the latter result is to be explained by the principle of natural selection. contact with a solid body plays the same role as did gravitation in the case of Antennularia. but the same is the case in Tubularia. is Another form. but here one more point may be mentioned. the polyp begins to grow away from the body. the stem. in his Germ Plasm. for some reason. they grow at In what In Margelis. inasmuch as an animal without polj-ps could not continue to live. perhaps. the root. It might be said. In Pennaria. If we bring an outgrowing pol^-p of Pennaria into contact with a solid body. growing downward. but it is just as easy to produce polyps at both ends. to get an animal that ended at both ends in a polyp and had no root. but also the direction of their growth. but also the direction of their gro\\'th. namely. Even if it were possible to reconcile these facts with the principles of natural selection. 28). growing upward. Weismann seems to assume. and the new stem I is very soon nearly' perpendicular to the part of it the surface with which came into contact. In Pennaria this supposed impossibility was realized. and hence it would be impossible to produce roots at both ends. all. Organs surrounded on if all sides by water continue to grow in the form of polyps. just as favorable. have called this form of irritability stereotropism.

bodies. Here. that in observed of many animals I was unable to produce any kind in heteromorphism. every piece has its north pole on that side which in the unbroken magnet was directed toward the north. a magnet regard to the formation of organs. a solitary We find similar cases in ascidian. If is broken into pieces. has Tunicates. believe that such is the case. POLARIZATION I While the foregoing experiments were in progress. Negative stereotropism forces the polyps to grow away from the ground into the water. there are animals every piece of which produces. and in others we find more complicated may appear from the foregoing that such cases of heterois morphosis are confined to hydroids. These animals showed. Bardeen. It light. we make an incision. under ordinary circumstances. Likewise. but such not the case. Herbst found that in to be formed in the place of an excised observed phenomena of heteromorphosis of heteromorphosis have been produced crustaceans an antenna could be caused eye. Ciona intestinalis. other hydroids the place of origin of the different organs determined by relations. Van Duyne. at either end.92 The Mechanistic Conception of Life speak of positive stereotropism in the case of the root. . find roots only at the base in this we where it touches the ground. Positive stereotropism forces roots in contact with the ground to hold to it. a phenomenon with which in a we are familiar magnet.^ incision eye-spots are formed on both sides of the II. eye-spots If around the two openings into the pharyngeal cavity. and hence parts surrounded by water form polyps only. we may ask whether in these the contact with foreign which experiments determines the arrangement of the various organs. and Morgan in Planarians and so on (1912). it Thus animal In is happens that. hence parts in contact with the ground give rise to roots only. too. that organ toward 1 Since this was written phenomena In many animals. may not have the same effect in the I natural development of the organism. and may of negative stereotropism in the case of the polyp.

Fig. differently we make an incision in the stem. or or f d. I tried in every possible way to produce tentacles in the aboral end of a piece which had been cut out. — C^J an incision. in The c clearest example of an actinian. Cerianthus membranaceus. DiagramIf an incision matic. into the body-wall of the actinian. The same polarization is shown in the following place before variation of the preceding experiment.Physiological Morphology which found If it 93 was directed in the normal condition. while the c. but only from the side ef (Fig. curious to see out only from lower edge c h. Diagrammatic. of The productakes tion tentacles any other regeneration begins. formed. only the h c. We may speak this I in such cases of polarization. Nothing d c of the sort oce. from Cerianthus. 30. which was directed toward the oral end of the animal. They endeavor to force them into the new oral disc. upper lip. but without success. The two that one- ends heal together in such a way It \J Fig. acb lower lip. 30). 29). a . 31). where the mouth should be. a produces none. a cb is made into the body of Cerianthus new tentacles grow half of a mouth. 29. we d ef. b (Fig. new tentacles are formed only at the upper cut ef. produces tentacles. curs in the side c . and only after a struggle futile of some minutes do they give up the attempt. cut a rectangular piece. a sea anemone. as regards polarization. If a piece c d ef is cut out from the wall of Cerianthus. If we make (Fig. the how these tentacles behave if we offer them bits of meat. a If little Hydra behaves. out tentacles of the body-wall of Cerianthus new soon begin to grow out of this piece. with is its surrounding tenis — tacles.

therefore. reproduce only the lost organ. but if a piece of meat is offered to the group of tentacles at b they seize it and press it to the place where a mouth ought to be. in others the most definite polarization exists. corresponding to the fraction of the periphery laid bare by the incision. In this case we must assume that unkno^Mi internal conditions determine the arrangement of limbs. Formation of a second head in Cerianthus after a Fig. From natvire. that while in — some animals we are able to produce heteromorphosis. but otherwise polarization. sho\ving the purely machine-like character of all these reactions. Only a fraction of the normal number of tentacles are formed lateral incision at b. 31. we find cases in which both phenomena . In addition to examples of heteromorphosis or polarization occurring separately. No new mouth is formed.94 The Mechanistic Conception of Life it whole new oral pole grows out. and we are able to produce regeneration of lost parts only in the arrangement which exists in the normal animal. but never show any heteromorphism. We see. so far as we know. too shows A good many animals.

such cases the aboral polyps were produced almost as quickly as the oral polyps. . produces a polyp. 28 an animal was represented which ended at both ends in a stolon or foot. true case of heteromorphosis cases the polyp at the oral In all end is formed first. by inhibiting or retarding the formation of the oral pol3rp. and place it in the carefully protected rior bottom of a dish of water. and we have a (Fig. If we cut out a sufficiently large piece of the stem of Tuhularia mesemhryanthemum. at tlie other in a head or polyp. the anterise to end of the piece gives a new if polyp. Thenormal Tubularia THE MECHANICS OF GROWTH IN ANIMALS ends at one end in a stolon. This could be done readily by diminishing the supply of oxygen at the oral end. while in Fig. and sufficiently provided with oxygen. Heteromorphosis in Tuhularia. In Fig. and a relatively long time (one or more weeks) the aboral polyp is elapses before formed. Under one con- dition. however. We can thus produce an animal wliich terminates in a head at both ends of its body. namely. from jarring. the posterior end to a root. too. this end. It ligature was found later independently by both Godlewski and myself that if we the stem of a Tubularian the polyps at both ends are formed simultane- ously (1912).^ III. 32. but we hang up posterior the stem in such a way that the of end does not touch the surface is the glass. If a piece o 6 is cut out and suspended in water a new liead or polyp c and d is formed In order to arrive at an explanation of the ask phenomena of organization we must what the physical forces are that determine the formation of a new organ. — From nature. 32). I could cause the stem to form a polyp at the aboral as quickly as at the oral end.Physiological Morphology are exhibited 95 by the same animal. We know energy for 1 that all the ultimate sources of the functions of living bodies at both ends.

in the germinating seed. The becomes stretched and. The result is when substances cell. possesses peculiar osmotic properties. The result is that water enters the by the pressure of the water within the cells their walls The chemical and are stretched out and the seed grows.96 are The Mechanistic Conception of Life chemical processes. seed. the same of as the conditions for growth in plants. temperature in perhaps. A brief outline of the : manner it growth in plants is as follows Before the grows forms substances which attract the physicist expresses it. . How can these chemical forces be brought into relation with the visible changes which take place to this question in the is formation of a to be obtained It is new organ ? The answer by a knowledge of the in the history of mechanics of growth. we have cell the beginning of a basis for work. subject has been worked out very carefully in plants. in conse- quence of which molecules of that it allows molecules of water to pass through freely while remaining resistant to the passing through of the many salts dissolved in the water. thus rendering growth growth is most conspicuous. of very remarkable that the mechanics animal can refer here only to the few but fortunately the growth forms almost an empty page I experiments morphology and physiology. and as my experiments show that the conditions for growth in animals are. the}' originate. to a certain extent at least. as forms substances which determine a higher osmotic pressure cell within the than did the substances from which cell. according to Traube. of higher osmotic pressure are formed cell-wall is inside the water from the outside passes in until the pres- sure within again equals the pressure without. The rising spring produces in the seed substances of higher osmotic pressure (with greater attraction for water) than the substances from which they originate. water from the surroundings. new material permanent. This method of precipitated in the enlarged interstices. it or. I have made on this subject. The question is. The walls of the or rather the protoplasmic layer that lines the cell-wall.

that within was greatest in the certain limits of concentration the increase in the length of the stems during the same period most diluted and It is least in the most concenof trated sea-water. is formed and which causes the swelling may be an found that acids cause a swelling of muscles and it has since been shown that this a general phenomenon. a limit effect. These and similar experiments. are represented on the axis of the abscissa. The following curve (Fig. In some of the dishes the concentration had been raised by adding sodium chloride.^ is 97 needed In order to ascertain whether I could determine what are I the mechanical causes of growth in animals. we should expect finally. that the more diluted the sea-water the more rapid would be the growth in the Tubularian stem. though this of course may not be the case in all animals. the amount being greatest in the most diluted solution and least in the most concentrated solution. remarkable that the maximum growth took place not in sea-water of normal concentration. indeed.Physiological Morphology osmotic changes are the sources for the energy which to overcome the resistance to growth. I chose long stems belonging to the same colony and distributed them in a series of dishes containing sea-water of different concentrations. The values for the amount of sodium chloride. which for lack of space cannot be mentioned here. and in others it had been lowered by adding distilled water. Of course. in 100 cubic centimeters of sea-water. If now in reality the mechanics of growth is the same for animals as for plants. is reached where the water begins to have a poisonous It was found. According to the laws of osmosis the cells of amount of water absorbed by the these Tubularians differed with the concentration of the sea-water. the values for the increase in growth on the axis of ordinates. 33) will give an idea of the dependence of growth upon the concentration of the sea-water in Tuhidaria. show that growth in animals is 1 The substance which I is acid. began at Naples some experiments on Tubularia mesemhryanthemum. but in more diluted sea-water. .

increase in size. is determined in which result by chemical processes within the a higher osmotic pressure. The abscissae represent the concentrations. formed than those from which Everyone knows that by exercise our muscles No satisfactory explanation of this fact has . Curve representing the influence of diluted sea-water. a conclusion seems to solid walls. This does not determine the peculiar character of growth. while the normal concentration is indicated by the vertical line — between 3 and 4. while this lie is not the case in most animal The solid cell-wall. in the fact that An obstacle to such many plant-cells have cells. The maximum growth is at a concentration between 2 and 3 per cent of salt. Both these qualities are inde- pendent of the cell should not agree in and I see no reason why the animal these two salient features with the plant-cell. which Pig. character cell. solid cell-wall. In order that the foregoing explanation of the mec hanism of growth in the animal cesses. chemical processes of such a character take place that substances of higher osmotic pressure are they originate. but does not allow dissolved in it all salts to do the same. the osmotic qualities of the outer layer of protoplasm. in case of growth. and. however. it cell might be based only upon knoTVTi prowas necessary to find out whether. by first. 33. the ordinates the corresponding growth in the unit of time.98 The Mechanistic Conception of Life determined by the same mechanical forces which determine growth in plants. allows water to pass through freely. secondly.

during normal activity. as soon as it ceases to be in size. for the gastroconemius that had been tetanized from twenty to forty minutes it varied from 1.^ This increase of osmotic pressure inside the muscle-cell leads. like gro^iih in whether segmentation. This increase in osmotic pressure is probably caused by the formation of Two years after the publication of this lecture I showed that the muscle The recent work of Pauli swells in an isomotic solution if this solution is acid. is influenced high concentration of sea-water it may be stopped entirely. begins to decrease Activity. by the amount of water contained in the If we decrease the amount of water in the egg of the cell. We fomid that while this concentration for the resting gastroconemius was about 0. The active. Cooke in my laboratory. The protein salt is more strongly dissociated than the protein base (1912).5 per cent. e.75 per cent to 0. I must expect that during activity substances are formed in the muscle. increase in volume. cell plaj^s still Therefore the amount of water contained in the another role in the process of organization and influences the process of cell-division. to a taking up of water from the blood and lymph. or rather a so-called Ringer's mixture. we were able to show directly that during activity is the osmotic pressure inside the cell-wall raised. In experiments which were carried on by Miss E. in which the gastroconemius of a frog neither lost nor took up water. loses Ranke had already sho\\Ti that the blood of a tetanized frog water and that this water is taken up by the muscles. plays the same role in the growth of a muscle that the temperature plays in the I tried to ascertain gro^\i:h of the seed.. which determine a higher osmotic pressure than those from which they originate.Physiological Morphology been given. were If 99 of g^o^^i:h my interpretation of the method correct.2 per cent to 1. therefore. . and Handovski indicates that the swelling is caused through a formation of salt between the acid and a weak base.g. and the consequence is an same muscle.85 per cent. This is exactly the case. and if we use a sufficiently general. sea-urchin segmentation is retarded. We deterof mined the concentration of a solution of NaCl. 1 acid. a protein.

the different regions the unsegmented egg correspond already to the different regions of the differentiated embryo. flat in the blastoderm of a chick. a of amniotic folds. very simply. board. It can be shown. In the blastoderm of a chick. . of from the According to him. sticking two tacks distance apart. 1 But this so-called is theory Another method of producing twins from one egg discussed in the last chapter of this book. the center of the blastoderm growing more rapidly than the periphery. parts of the blastoderm? Why is growth different in different But instead of trying to it answer it from the physiological standpoint he answered anatomical standpomt. His raised the question. The idea that the formation of the vertebrate embryo is a function of growth has been investigations of His. If we by plate of elastic rubber. His has sho^\^l how inequahty of growth determines the differentiation of organs. these processes of folding are due simpl}' to inequalities of growth. that such a process of unequal growth must. and the system all According to His.100 The Mechanistic Conception of Life THE artificial PRODUCTION OF DOUBLE AND MULTIPLE MONSTROSITIES IN SEA-URCHINS^ IV. In this faster than the periphery. we get the same system I of folds as that shown by the still embryo of a chick. so that the rubber in the stretched. indeed. and lay it on a drawinga short we can imitate the stronger growi:h in the center into the middle of the rubber. is a process of folding. and then pulling them in opposite directions. mention this way of demonstrating the effects of unequal growth as the ideas of His are doubted by some morphologists. for example. the first step in the origi- formation of the embryo nates a head fold. a medullary groove. the center growing If we then fix the tacks in the middle remains drawing-board. made the basis of the embryological In a masterly way. lead to the formation of exactly such a system of folds as we find take a thin. There tail fold. way we may imitate unequal growth.

" causal morphology can make no use an explanation. ficial would give But it has been said that the arti- isolation of one cleavage-cell causes a process of postDriesch. of heteromorphosis would show that. the heat of our blood inherited. adhesion. It may be changed by must be asked. Zoologie. we may are ovum somewhat chemically. Neverthe- not necessarily in opposition to the theory of growth Starting with the idea of His. it is embryo grow faster than others. what. .Physiological Morphology of 101 preformed germ-regions gives no answer to the question. in some animals at the arrangement of preformed germ-regions gravitation. however. If we answer of such ''heredity. etc. cells we should rise to expect that every one of the isolated one-fourth of an embryo. light. but although our parents had the same temperature. mode of the first cleavage by submitting the ovum LV. Thus the phenomena least. its later If germ-regions with modifications were true. determines the arrangement of the different germ-regions the egg. which differs only in size from the normal embryo. is not the result of certain chemical processes in our Still it may be possible that the molecular forces of the chemically different substances of the egg determine a separation of these substances and thereby give rise to the chief directions of the future embryo. from the standpoint in of causal morphology. pressure. 1 In this f. LIII. way the nuclei were brought into somewhat Zeitschrift wissensch. well imagine that the different regions of the different offered in the preceding chapter. and each one be the theory of preformed Driesch has shown^ that by shaking a sea-urchin's egg in the four-cell stage the four cells may capable of giving rise to a complete embryo. of the why some parts less. and that these chemical differences of the different germ-regions determine the differ- ences of growth in the blastoderm. Our is blood has the temperature of about 37°. moreover. changed the to one-sided generation or regeneration. but tissues. be separated.

which often. have made a series of experiments to the results of which these objections I shall describe these cannot be made. The protoplasm swells until the membrane m bursts and part of the protoplasm m addition of about 100 per cent distilled water. Fig. remained in connection with the protoplasm inside the membrane after the eggs were brought back into normal sea-water. 35 — Fertilized egg of a sea-urcliin (Arbacia) put into dilute sea-water. 35. and they then developed into perfectly normal single embryos. and not in the nucleus. The part of the protohad connected the two drops. each of the two droplets may develop into a blastula. as experiments somewhat I they have not yet been published. fully.102 The Mechanistic Conception of Life from those they would have held m the case of Still. 34 Fig. 34) the eggs took up so much water that the membrane burst and part of the protoplasm escaped in the form of a drop (6. 34). movement. Fig. 6 flows out. such an egg two larvae may arise. Since we caimot assume that in every case the by accident or rapid . the twins were separated. might object again that the preformation existed in the protoplasm. however. I brought eggs of a sea-urchin. within ten to twenty minutes after impregnation. Fig. as the part remaining inside the These eggs gave rise to adherent twins. as well normal and perfectly plasm. 34. it part where the twins remained gro\Mi together. formed the Of course. often happened that. as Indicated In this solution (m. which at first membrane. normal embryos resulted. so that from Fig. One of the germ-regions I different places normal segmentation. developing into a complete embryo. the ejected part h of the protoplasm. into sea-water that had been diluted by the Fig. though cannot enter into details at this place.

is now understood by V. but triplets all and quadruplets It is grown together. we must conclude that every part of the protoplasm may give rise to fully developed embryos without regard to preformed germ-regions. however. and I obtained not only double embryos. . protected from microbes and infusoria. But this fact alone does not explain why it is that we get I In the light of more recent experiments it is possible. but it would be something totally different from what ovum by the molecular qualities of these liquids. or with the formation of One of the steps that to show in every case the connecting link between the chemical processes and the formation of organs. I have tried to show that in a few cases at least this connecting link was to be sought in the changes of osmotic pressure determined by the chemical changes which an embryo or a single organ. might be called a preformation of germregions. physiological morphology has to take is take place in the growing organ. remarkable that the development of these monstrosities goes on nearly at the same rate as that of the normal embryo. that term. with the process of secretion. THEORETICAL REMARKS are determined All life phenomena by chemical processes. perfectly well possible that this prearrangement consists in a separation of different liquid substances in the Such a separation. that after all only such pieces can develop into a normal embryo which contain the different germ-regions (1912). cases each of the drops of the protoplasm In many In such may give rise to an embryo. provided they are equally well supplied with oxygen and equally The development in most eggs takes place in so regular and typical a manner that it seems as if there were a prearrangement of some kind.Physiological Morphology same part of the 103 protoplasm escapes. of course. It is. 1.^ eggs a repeated outflow of the protoplasm takes place. This is equally the case whether we have to do with the contrac- tion of a muscle.

That organs which is differ in shape are very often chemically different a well-known fact." as in their chemical constitution. this we must bear in mind that the must As necessarily be different for different organs. 'Hhat differences in the form of organs are accompanied by differences former from the latter. 1 For these phenomena the wave theory of Form der Pflanzenorgane. Stoff und Gesammelte Abhandlungen. or. In this way we are led through the mechanics of growth to a conclusion which forms the nucleus of Sachs's theory of organization. as for example in the formation of a root. if led to the idea that differences in the forms of different organs differences in their chemical constitu- the chemical constitutions be similar. Differmust be the outcome of differences in the chemical processes which determine growth. J. and the formation of a stem. we must avoid a mistake very often made even good theories. in the case of Huyghens explained by refraction. differences of growth can only be determined either by differences in the amount of energy set free in the growing organ or ences in the energy by differences in resistance. II. namely. The formation of urea in the liver and the synthesis of hippuric acid by the kidneys are the consequences of chemical differences. and that according to the principles of science we have to derive the According to Sachs there are at least many 2. by differences in resistance to growth.104 The Mechanistic Conception of Life In order to understand processes of growth differences in the forms of organs. namely. 1893. growth is a process in which energy is used up in overcoming the resistance to growth.^ In adopting the theory of Sachs and applying it to animal morphology. Therefore we are must be determined by tion. the endeavor to explain which special cases are complicated by unknown conditions. his theory of light the phenomena of but he could not and did not attempt to explain the sensations of color. ''spezifische Bildungsstoffe " in a plant as there are different organs. Sachs. .

For instance. They all of lie upon the walls of the blood-vessels as well as in the spaces between the capillaries (Fig. Its chemical But it from a chick's egg only a chick can would be a mistake to attempt at present an explanation of how the unknown chemical nature of the germ determines all the different organs and characters that belong to the species. 38). . but color sensations depend not only on the of the ether. succeeded experimentally in showing to be probable that some of the substances contained in the blood determine this reaction. 36). wave motion but also on the chemical and physical I structure of the retina. think it perfectly safe to say that every animal has specific germ substances. These substances. make. and that the germ substances of different animals differ chemically. the yolk sac of the Fundulus tiger-like coloration. and possibly changes in the and not a preformed arrangement of the germ. cells the pigment have crept upon the surface I of the neighboring bloodit vessels (Fig. In this way the chroma- tophores are arranged in stripes. the pigment cells first But the arrangement is days of development not such as to produce any definite markings. according to the laws of surface tension. The pigment chemically closely related to hemoglobin. if they diffuse from the blood-vessel and touch the chromatophore. qualities determine that arise. however. the protoplasm of the chroma- tophore flow toward and at last over the blood-vessel and form a sheath around it. Later on. of chromatophores. which later on give rise to the colored places of the yolk sac. but I found that this coloration originates in a manner much more simple.Physiological Morphology light 105 remains true. and so its formation may from during the the first be connected with the of formation of the blood corpuscles. The pigment cells are is formed irregularly on the surface of the yolk. embryo has a markings We might say that these may be due to a certain arrangement of molecules or complexes of molecules (determinants). while the gaps between the blood-vessels become empty surface tension.

37.106 The Mechanistic Conception of Life Fig. the same egg still a few days later. The black pigment cells have completely crept on the blood-vessels and formed a sheath around them. 37 Fig. Fig. and 38. Here we notice that some pigment ceUs show a tendency to creep on the blood-vessels. . 37. 36. 36 Fig. 38 Figs. This was the first observation pro\'ing that tropisms play a role in the arrangement of the organs in the body. Fig. The red chromatophores are omitted in this — drawing. 38. The black pigment cells show no definite orientation in regard to the blood-vessels. From nature. The origin of the pattern on the yolk sac of a fish embryo {Fundulus heteroclitus). 36 is a drawing of the bloodvessels at the surface of the yolk sac at an early stage of development. the same egg a few days later. Fig.

while polyps only would grow out in the opposite direction. cal root It is possible that the hj'potheti- substances of Sachs may have a greater specific gravity than the substances which form stems. hydroids. while the opposite took place in the case of elements containing polyp substances. but the task of deriving such a coloration in the adult from a similar arrangement of molecules in the germ plasm would prove too much even for a genius like Huyghens. The reasons why roots grow on the under side of the and stems on the upper side can only be and chemical conditions inside the stem of Antennularia have been worked out. what determines the polarization may suffice to suggest the possibility . 3. At present we stem of Antennularia given when the special physical can only think of possibiHties. I found. Since outgro\\i:h can take place only at the free surface of a stem or branch. But there are still other possibilities which we must omit here. and without the possibility of such a derivation the theory is of no use. sion of the question of like Cerianthus. more But we must wait with our attempts For the same reasons It I will at explanation and chemical conditions for the form are not go into a discusof ani- worked mals out. indeed. it In the case of Margelis and other solid bodies pro- might happen that contact with of surface in the duced an increase touched elements in case they contained specific root substances. and therefore take the lowest position in the cell. The conif sequence would be an increase in the surface of the roots they came into contact with solid bodies. in some forms at Naples that roots of hydroids which grew free in the water began to grow in these cases much faster and to branch off abundantly when brought into contact with until the physical solid bodies. roots can on this assumption grow only at the under side and stems only at the upper side of an element. 107 We do not know what processes determine the coloration of animals which owe their markings to interference colors.Physiological Morphology determine the marking.

1 1 stated . if both cavities communicate with one another. In my experiments. as there must be a minimum becomes impossible. but to other circumstances. as every sphere gives rise to a blastula. one single necessarily formed a whole as it the four-cell stage embryo after it had been isolated. in so far as these circumstances determine the I got double or number of blastulae. assumed the shape of a single sphere or ellipsoid. triple embryos when the egg formed two or cell of three droplets or spheres. under normal conditions. and only one embryo. which. is possible.^ Without the size for the cleavage-cells — that the minimal size is about one-eighth of the mass of the seaurchin's egg and I do not think that this is very far off the limit. The egg of a sea-urchin under normal conditions gives is rise to but one embryo. or nucleus. but the limit is not due to any preformation. by reason surface tension of the protoplasm. as a two separate segmentation cavities are formed. the number of embryos which come from one egg is not determined get twins. that which of the the geometrical shape of the protoplasm. while in cases of heteromorphosis migration or arrangement in every direction or in several directions 4. In Driesch's experiments. if we by the preformation of germ-regions in the protoplasm. When this happens. there must be a limit to the number of embryos that can arise from one egg. Whether the first or the second case will happen depends upon the molecular condition of the part of the protoplasm connecting the two drops. Therefore. When we make the protoplasm outside the egg membrane and remains within it assume spherical forms. This circumstance due simply to the eggs burst. rule. is that of a sphere. the chief one being that with too small an amount of protoplasm the formation of a blastula from merely geometrical reasons. Of course.— 108 The Mechanistic Conception of Life cells that in polarized animals the tissues or may have such a peculiar structure as to allow the specific formative substances to migrate or arrange themselves only in one direction. but by the geometrical shape of the egg and the molecular condition of the protoplasm.

But the aim analytical. inasmuch as their object has been to derive the laws of organization from the ena. or constructive. which are determined by the have chosen the name Physiological Morphology for these investigations. .Physiological Morphology formation of the blastula. just as the phj'sicist and chemist form new combinations from the elements of non-living nature. common source of cell. that to form new combinations from the elements of living nature. of course later stages I it is 109 not possible to get the blastula. is i.e. all life phenom- the chemical activity of the is In what way this to be done indicated in the chapter on the mechanics of growth. which is. It of Physiological Morphology is is not solely synthetical has another and higher aim..

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ON THE NATURE OF THE PROCESS OF FERTILIZATION .VI.

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1899. the so-called spermatozoa. He believed that the spermatozoa represented the future embryo. frogs 1 and by Spallanzani who put the males of during the act of cohabitation into trousers and convinced Lectures delivered at courtesy of Ginn Reprinted. from Biological & Co. the for the act of fertilization. volatile contained sufficient imaginary ''aura seminalis" was That contact between sperm and egg was really necessary for fertilization was at last proved experimentally by Jacobi (1764) who showed is that fish eggs can only be fertilized if the sperm brought into direct con- tact with the eggs. It was generally assumed that no direct contact between sperm subside until and egg was necessary and that something in the sperm. fertiliza- The idea that spermatozoa are not parasites did not it was proved about 160 years later that the spermatozoa originate from the cells of the testes. to Not until recently have biologists begun become conscious of the uncertainty of conclusions which are not tested and verified by adequate experiments. but that the spermatozoa and not the liquid constituents were the essential element in the sperm was a fact which could not be established except experimentally.. Leeuwenhook demonstrated in 1677 the existence of motile elements in the sperm. by 113 .VI ON THE NATURE OF THE PROCESS OF FERTILIZATION^ I Experimental biology is a very recent science. Boston. That sperm was needed to bring about fertilization of the egg was too obvious a fact to escape even those biologists who never made an experiment. Woods Hole. The majority of his contemporaries assumed that the spermatozoa were parasitic organisms which had nothing to do with tion.

but a chemical constituent. Prevost and Dumas famous chemist) to prove that the spermatozoa are the essential element in the sperm. In 1843 the entrance of the spermatozoon into the egg was directly observed by Barry and this fact has since been verified by an endless number of investigators for the egg of all kinds of animals. that causes Against this second view Miescher the development of the egg. They made the simple experiment of filtering the sperm and demonstrated that the lost its sperm whose spermatozoa had been retained by the filter had power of fertilizing the eggs. raised the objection that his investigations showed the same . offered. is responsible for the process of fertiliza- The second theory theory it is is a chemical one. the morphological structure of the spermatozoon which tion. It is probably no exaggeration to say that with the general recognition of the experimental method in biology it would probably have taken about as many years as it took centuries to establish the simple is fact that the spermatozoon the essential element in sperm. Nevertheless four theories or rather suggestions were The first theory of fertilization it is is a morphological one.114 The Mechanistic Conception of Life himself that under such conditions the eggs remained unferti- Hzed although the "aura seminalis" was not prevented from acting upon the eggs. But even this did not convince many of the descriptive biologists and nine years later K. According to this not a definite morphological or structural element of the spermatozoon. E. According to this theory. The mere observation of the fact that the spermatozoon must enter the egg in order to bring about fertilization did not lead to any understanding of the mechanism of the activation of the egg.'' It (the latter the was reserved to two experimenters. This ended the reign of the ''aura seminalis. von Baer still expressed the opinion that spermatozoa had nothing to do with fertilization.

of course. because it assumes exactly the same for the spermatozoon that Liebig Liebig thought that the enzymes their assumes for the enzymes. According to this conception the egg considered as a definite machine which its work in a certain direction. experiments he was not familiar with such Moreover. I do not beheve that this objection valid. owed power of producing fermentation to the motions of of atoms. however. stimulus which causes the egg to undergo its development. This is have an enormous effect upon shown among others by the work of Emil Fischer on the relation between the molecular configuration of sugars his and their fermentability. nothing but a disguised repetition of the chemical or physical theory of All these theories are so vague that we do not need to be to in biology. It should be said. A third theory was a physical theory is (Bischof). any further discovery. theory assumes that a peculiar condition of motion exists in the spermatozoon which its is transmitted to the egg and causes development. Miescher was not able state whether the This spermatozoa contain enzymes or not. When to Miescher made possibilities. that this idea not so very different from the chemical conception. and not from vague Among these observations the most important them has led make new discoveries . is certain atoms or groups The was do is fourth conception the stimulus conception. which is if once wound up will The spermatozoon is the originated by His. by the spermatozoon is of a physical or a chemical and in this way. surprised that none of If we want to from definite speculations. we must start facts and observations. We know that simple variations in the configuration of a molecule life phenomena. the stimulus conception is fertilization. It to be said in connection with this stimulus conception that is the main point at issue carried omitted as to whether the stimulus character.Nature of the Process of Fertilization compounds in the egg is 115 and the spermatozoa.

Die Zelle und die Gewebe. A beginning of parthenogenetic development had been observed in the case of a great many marine It develop outside of the female in sea-water. who discovered that the unfertilized eggs of bees develop and give rise to males. certain that a few crustaceans show parthenogenesis. . It was. the so-called teratomata. 1893. when put back after they had been in such sea-water for about two hours. 239. exist in wasps. p. The most impressive fact concerning the parthenogenesis of animals was contributed by Dzierzon. On the other hand. But all these were considered to be of a pathological character. Jena. admitted that we cannot utilize these phenomena It must be. II Eight years ago I observed that if the fertilized eggs of the sea-urchin were put into sea-water whose concentration raised was to segment. in ovaries of mammals now and then eggs were found that were cells. while the Similar conditions seem to fertilized eggs give rise to females. moreover. broke up into a large number of cells at once instead of dividing successively by the addition of 1 Hertwig. was not until very recently that such experiments were made. facts with any It degree of certainty for the theory of fertilization. however.116 The Mechanistic Conception of Life It are those on parthenogenesis. but that the some neutral salt they were not able same eggs. animals which was found that divide into such eggs when left long enough in sea-water may two or three cells. had been observed for a long time that the imfertihzed egg of the silkworm can develop parthenogenetically. moreover. suggest the possibility of at least partial parthenogenesis in the eggs of mammals. kno^\^l that plant lice can give rise to new generations without fertilization. O.. It is. ^ segmented into a small number of These facts and the occurrence of a certain class of tumors in the ovary. as in this case certainty can only be obtained bj^ the experiment. but no farther.

. of MgCl2 to 100 c. Morgan^ had used the same concentration of sea-water as Norman^ and I had used in our previous experiments." Biol.c. etc.Nature of the Process of Fertilization into two. by the As is Loeb." Journ.^ As soon as the egg is put back into normal sea-water it at once breaks up into as many cleavage-cells as nuclei or distinct chromatin masses had been preformed in the egg. went as far as about sixty cells.. A small number two or four cells. f. very small amount of KCl to sea-water he could force the unfertilized eggs of Chaetopterus to throw out their polar bodies. * Mead. III. "Experiments on Cleavage. was able to show the beginning of a segmentation when put back into normal sea-water. Morgan tried the same experiment on the unfertilized eggs of the sea-urchin.. little NaCl was for KCl did not have the While continuing my studies on the effects of upon life phenomena. 1896. of sea-water. 1899. W. sixteen cells. f.c. and Morgan used the same concentration. 1898. I had added about 2 grams of sodium chloride Norman used instead of this 3§ grams to 100 c. 3 Norman. four. in a few cases. and found that the unfertilized egg. Woods Hole A. 2 Morgan. E ntwickelungsmechanik. 117 neces- Of course it is sary for this experiment that the right increase in the concentration of the sea-water be selected. 1892. of sea-water." Arch. but no larvae ever developed from these eggs. eight. etc. "The Action of Salt Solutions. I led to the fact that the peculiar actions of protoplasm are influenced to a great extent ions contained in the solutions which surround the 1 cells. . Mead'' made an observation somewhat similar He found that by adding a to Morgan's upon Chaetopterus.. fact is as follows: The explanation of this The concentrated sea-water brings about a change in the condition of the nucleus which permits a division and a scattering of the chromosomes in the egg. H." Arch. and. of Morph.. of eggs divided into The same salts substitution of a effect.. W. if treated for several hours with concentrated sea-water. T. VIII. " The Rate of Cell-Division and the Fimction of the Centrosome. " Segmentation of the Nucleus without Segmentation of the Protoplasm. VII. Entwickelungs- mechanik.. J. Led.

some of these ions are able to combine with. Ill. like our heart . the It only necessary to increase number of sodium ions in the muscle or to reduce the num- ber of calcium ions or do both simultaneously. charged with a certain quantity of positive electricity) and CI ions (CI electricity). Joum.. J.g. and thus are able to impart properties to a tissue which it does not possess ordinarily. that by changing the amount of sodium and calcium ions contained in the muscles of the skeleton we can make them is contract rhythmically like the heart. which were in every to those produced way similar by the first fertilized egg. There seemed to be three ways in which accomplished.^ ions By changing the relative proportions of these we change the physiological properties of the protoplasm." Amer. 1900. it atoms charged with the same amount of negative When an egg is in sea-water. sodium chloride. I have found. At any are rate. e..of this On the basis and similar observations it I thought that by changing the constitution of the sea-water might be possible to cause the of eggs not only to show a beginning development but to develop into living larvae. . the proteins of the proto- plasm. the various ions enter determined by their osmotic pressure and the It is probable that in proportions permeability of the protoplasm.. 1 The second way was to increase the osmotic pressure Loeb. the physical qualities of the proteins of the protoplasm (their state of matter and power of binding water) determined by the relative proportions of the various ions present in the protoplasm or in combination with the proteins.. of Phys. for instance. this might be The way was a simple change in the con- stitution of the sea-water without increasing its osmotic pressure.It is due to the Ca ions of our blood that the muscles of our skeleton do not beat rhythmically. 118 The Mechanistic Conception of Life if we have a salt in solution. "On lon-Proteid Compounds and Their Role in the Mechanics of Life-Phenomena. but a certain number of NaCl molecules are split up into Na ions (Na atoms well kno^Mi. we have not only NaCl molecules in solution.

of sea-water for two hours the eggs began Such eggs to develop when put back into normal sea-water.c. the the egg developed into one blastula. of -^^-n MgClg soluinto a of the sea-urchin tion and 40 c. It became extremely irregular. In this case I had no better results than few cells.c. but I I next tried the effects of Morgan. however. increase in the sea-water by adding a certain amount of magnesium chloride. after by putting the unfertiHzed eggs solution of 60 c. The were in a certain way similar to those I had obtained when caused the fertilized eggs of sea-urchins to burst. a rare thing that the whole mass of rule. as a stage. The disconnection results I of the various cleavage-cells led. reached the blastula stage. See next chapter.^ All the various effect of causing artificial solutions I prepared had only the one the unfertilized egg to divide into a was not an able to produce a blastula. and the outlines of the egg showed. as a rule.Nature of the Process of Fertilization of the sea-water 119 by adding a certain amount of a certain salt. that found. The way did not lead to the result I desired. I then tried beyond develop the combination of both methods. but these did not the first stages of segmentation. In such cases a part of the proto- plasm flowed out from the egg but was able to develop. As these eggs had no membrane. These extraovates had no membrane. I do not think that anybody has a number ever seen before such blastulae as resulted from these unfertilized eggs.^ solution. positive results. the amoeboid motions of the cleavage-cells led very frequently to a disconnection of the various parts of one and the same egg. Very few eggs began to divide. The first third way was by combining both of these methods. is ordinary sea-water of a The osmotic pressure of roughly estimated to be the same as that I fn NaCl solution or a ^-n MgCl. moreover. The blastulae same outline as the egg had in the morula was. but the irregularity in this case 1 Later experiments gave. . of experiments. to the formation of more than one embryo from one egg. and of course showed some irregularity in their outlines.

the eggs. The outlines of each blastula were much more spherical than in the previous experiment. It was my next task to find a solution which would allow I the eggs to reach the pluteus stage. on the blastula stage. not only reached the blastula stage. Today it may seem strange that I had . Thus I had succeeded urchins to the same stage to which the fertifized eggs can be raised in the aquarium. The most absurd sources of error were suggested. but went into the gastrula and pluteus stages. and died. The blastulae that into originated from these eggs looked much healthier normal than those of the former solution with and more more MgCla- Of course as these unfertilized eggs had no membrane it happened but rarely that the whole mass of an egg developed into one single embryo. They developed more slowly than the normal eggs. very few biologists were willing to accept the correctness of my statements. as a rule. Though I do not wish to go into the technicalities of these experiments. but when my first papers on artificial parthenogenesis appeared. These eggs reached the pluteus stage on in raising the unfertilized eggs of sea- the second day (considerably later than the fertilized eggs do).^ 1 The reader who to is interested in this meet such objections. fertilized eggs in I have not yet succeeded in raising the my laboratory dishes beyond the pluteus stage. and twins were much more frequently produced than a single embryo. after they were put back normal sea-water. I second day. triplets. found that this can be done by reducing the amount increasing the lized eggs for -2^-n and amount of sea-water. But although had thus far satisfied my my desire to see the unfertilized eggs of the sea-urchin reach the was not able to keep these eggs alive long enough to see them grow into the pluteus stage. Quadruplets. By putting the unfertiabout two hours into a mixture of equal parts of of magnesium chloride MgCl^ and sea-water.120 The Mechanistic Conception of Life far less was than that observed in the unfertilized eggs of I recent experiments. I must mention a few of the precautions that I took in order to guard against the possible presence of spermatozoa in the sea-water.

Part of the unfertilized eggs were put into the for the eggs that did in same normal sea-water that was used None of these eggs that remained normal sea-water about twenty- formed a membrane or showed any development. 1900. made another set of control experiments by putting a lot of eggs of the same female into a solution w^hich differed less from the normal sea-water than the one which caused the formation of blastulae or plutei from the unfertilized eggs.." Amer. artificially had a membrane. In the earlier experiments in which no fertilization membrane was developed. of Anat. The method used in these experiments was primitive inasmuch as no fertilization membrane was formed. except that a few of them were divided I into two cells after four hours. which is described in the next paper. publication in the American Journal Here wish only to mention the following points: These experiments were made after the spa^\Tiing season was practically over. nevertheless a change in the cortical layer of the egg was brought about by the combined action of the hydroxyl-ions of the solution. develop. 3. 6. The spermatozoa contained if in the sea-water lose. . egg is fertilized or not in the production of a membrane. J. Gemmill.^ their fertilizing power within five hours 4.water. "The Normal Larvae from the UnfertiJoum. distributed in large quantities of sea. tell We have a criterion by which we can whether the The membrane the unfertilized has no and egg fertilized egg forms a None of the unfertilized eggs that develdistinct membrane. and Phys.^ With each experiment a number of control experiments were made." Joum. 2. III. or instruments with spermatozoa. accord- ing to the investigation of Gemmill. Bacteriological precautions were taken against the possi- bility of contamination of the hands. the Ova and Spermatozoa of Certain Animals. oped 5. 1 Loeb. A few years later I found a method for the artificial production of a fertilization membrane.. dishes..Nature of the Process of Fertilization technical side of the experiments will find in all 121 the necessary data of Physiology} my I 1. and the 3 increased osmotic pressure. of Phys. "On the Artificial Production of Vitality of lized 2 Egg of the Sea-Urchin.

yet not In this case one single egg formed a membrane or reached the blastula stage. changes of these physical qualities we know specific The one enzymes plasmase. qualities are It seems probable that changes of these two mainly responsible for muscular contraction and perhaps amoeboid motions. we may assume that by treating the eggs for solution of equal parts of a ^ n MgCl. The third agency is temperature.). will have to be in the following direction: The bulk of our protoplasm consists of colloidal substances. Among is the agencies that cause of three that are (trypsine. solution Ill two hours with a and sea-water we can cause them to develop parthenogenetically into plutei.122 The Mechanistic Conception of Life it was shown.^ Hence. . The concentration varies for various ions. I think If we we should confine ourselves to the following question: What it alterations can be produced in an egg by treating for two hours with a solution of equal parts of ^^n MgClg and of sea-water ? Even in this regard we can only give a very indefinite answer which. and although they were injured less than the ones which developed. that although these eggs received the same sea-water as the ones which developed. as in nine different series of experiments these results were confirmed. it do not consider advisable to enter into theoretical discussions beyond these 1 Through other control experiments I convinced myself that a treatment of eggs or spermatozoa with equal parts of a -g'^n MgClg solution and sea-water diminishes the impregnability of the eggs and annihilates the fertilizing power of spermatozoa in a very short time. If the sea-water had contained any spermatozoa these eggs should have reached the blastula stage. material easily changes its This of state of matter and its power binding water. In our experiments it is obvious that only the I second possibility can have been active. however. etc. What conclusions may we draw from these results ? wish to avoid wild and sterile speculations. The second is ions in definite concentration. especially powerful.

While we are able to produce the process of fertilization by a treatment of the unfertilized egg with certain salts in certain concentrations.^ 1 Loeb. I consider the chief value of the experiments on artificial parthenogenesis to be the fact that they transfer the problem of fertilization from the realm of morphology into the realm of physical chemistry . like the three agencies In the last volume of these lectures I pointed out that impossible to derive all the various elements that constitute heredity from one and the same condition of the egg.Nature of the Process of Fertilization statements. the inference must be that the transmis- sion of the hereditary qualities of the male causes the process of fertilization the and the agency that are not necessarily one and same thing. Boston.. . especially KgPO^. An account of this work is given in the two following papers. In the following years the methods of artificial parthenogenesis were improved and this led to the unraveling of the mechanism by which the spermatozoon causes the egg to develop. J. 2 This paper was written immediately after I had succeeded in producing larvae from the unfertilized egg. it is qualities of the colloids.^ Our recent experiments suggest the possibility that different constituents of the egg are responsible for the process of fertiliza- tion and for the transmission of the hereditary qualities of the male.." Woods Hole Biol. whether the spermatozoa act in the same way. Led. 1899. " The Heredity of the Markmg in Fish Embryos. but it It is true the spermatozoon contains a considerable proportion of may contain enzj-mes or effects it may con- tain substances which have similar upon the physical mentioned above. we cannot hope to bring about the transmission of the hereditary qualities of the male by any such treatment. Hence. 123 The next question that should be raised would be that salts.

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VII. ON THE NATURE OF FORMATIVE STIMU- LATION (ARTIFICIAL PARTHENOGENESIS) .

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By formative means those which give rise to nuclear and cellular : He considers as the classic example for formative stimulation the fertilization of the egg and the parallel drawn by him between If this process is and the causation of a pathological process of growth so characteristic that we may quote it in full we admit the identity between the pathological and the embry- onic neoformation. nutritive. 1909. paper on ''Stimulation and stimulation stimuli he division. that the spermatozoon carries chemical substances is into the egg.: VII ON THE NATURE OF FORMATIVE STIMULATION (ARTIFICIAL PARTHENOGENESIS)! PREFACE The XIV. entirely dissolved in the egg was not his view. the egg will have to be considered as the analogue of the pathological mother cell and the act of impregnation as the is analogue of pathological stimulation. 1858) in which he discriminates between three forms of functional. perfectly correct. 127 . title of this paper was chosen in reference to Virchow's Irritability" {Virchow's Archiv. which serve as specific stimuli. and formative. I therefore 1 Address delivered at the International jSIedical Congress at Budapest. The supposition prevalent spermatozoon but is at Virchow's time. into the egg. and like\\ise the analogy between the causation of the development of the egg by a spermatozoon and the causation of a pathological growth seems correct. since there is This view not essentially altered through the discovery of the entrance of the spermatozoon no reason to consider the spermatozoon as the direct morphological starting-point for the development of definite parts of the egg. as seems to be the case. by calling forth new chemical and morphological arrangements of the atoms. 1. that the correct. If. they carry into only certain chemical substances. which form the stimulus for its development. Each specific contagium offers the same possibilities. the spermatozoa are it dissolved in the egg.

which changes does the egg undergo in this formative stimulation by a spermatozoon ? Or in other words. in the cell in the process of formative which change occurs stimulation. however. since Virchow. If no sperma- tozoon enters. and second.128 The Mechanistic Conception of Life it believe that may be of interest to the medical profession to follow me in a brief survey of my experiments on artificial parthenogenesis and the causation of the development of the egg by a spermatozoon. and pathologist may therefore interest the and the physician in general to become familiar with the essential features of the data thus obtained. as a rule no segmentation of the egg takes place and it perishes after a comparatively short period of time. or what changes a it cell undergoes in such a This task has. been to It is the task of modern biology ascertain first what is the nature of these stimuli. It is known that aside from a few exceptions the animal egg if can only develop a spermatozoon enters into it. tumor is. Pathologists have not yet succeeded in determining what the physico-chemical nature of the formative stimulus in the case of a process. and second. fertilization of the Virchow already emphasized the egg is fact that the the model of all phenomena of forma- tive stimulation and that the spermatozoon may be considered as the formative stimulus in this case. I tried The questions which to solve were the following: B}^ which physico-chemical agencies does the spermatozoon cause the egg to divide and to develop into an embryo. is what is the mechanism by ? which the unfertilized egg caused to segment and to develop . I Cellular physiology has shown that cells tissues and organs into develop only from cells through nuclear and cellular division. to divide The conditions which cause new normal or pathological called formative stimuli. been accomplished to a high degree in the animal egg. and to develop tissues have.

II We will begin with a description of the method of in artificial parthenogenesis since here this It the egg of the Cahfornian sea-urchin. in the eggs of may be mentioned that many animals the itself effect of the entrance of the spermatozoon manifests of fertilization. easier to get the dark as to the chemical character of the active substances. Figs. i. in almost instantly by a characteristic change. 1 . cells The second way which could lead to a decision of the ques- tion concerning the nature of formative stimulation lay in the direction of artificial parthenogenesis. method has been worked out most completely. not by extracts of sperm but directly by physico-chemical agencies. but met at first with only negative results for the reason that I used at first only extracts from the sperm taken. have spent a good deal of time in trying to succeed in this task. where- by the former is it lifted separated from up from the protoplasm of the egg and by a more or less wide. the nature of the agencies insight into the of This method of Since in this case it we know an mechanism by which they cause the development the egg.. of the causation of the development of the animal egg. procedure has a special advantage. the formation Briefly stated this of of the so-called membrane process may possibly consist the entrance sea-water between the surface film and the protoplasm of the egg. namely. while if we work with extracts of sperm we are in is we employ. of the same if it species of animals I from which the eggs were is Only recently have is found that the extract of sperm effective only taken from a foreign species. clear space. We it shall return to this curious fact later on and show that has a to the bearing upon the problem of the immunity of our lysins of our body.e.Nature of Formative Stimulation Two ways first 129 were open to find an answer to this question: to try to cause the development of the unfertiUzed egg I with extracts from sperm.

a second influence as needed. how we know that the e. the eggs are transferred into normal the eggs form a perfect fertilization membrane. The answer that if we apply the acid but prevent the changes leading to a cell memdo not brane formation. the artificial production of a membrane in the egg by fatty acid only calls forth a nuclear division. of an n/10 lower monobasic fatty acid. if..g. sea-water all however. and possibly a cell- however. this process of The experiments showed that tion is membrane forma- the essential condition which causes the egg to develop. If the temperature is very low not only cell-divisions it begin but the egg may develop into a swimming larva.g.130 The Mechanistic Conception of Life 7) and 2 (page 1 show these changes m the sea-urchin egg. reaches the so-called blastula stage. another question has to be settled. butyric.. we shall see later on that we can . At room temperature. but that at ordinary temperature the latter does not go far. is the formative stimulus in this case. we shall see later. 2 shows the same egg after the entrance of the spermatozoon. propionic. Before we describe this second influence. e.c. to cause a complete development. Fig. On the other hand. membrane a catalytic. represents the unfertilized egg.c. in putting the eggs for about membrane of fertilizaThis method consists (at a two minutes temperature of 15°) into a mixture of 50 c. We by a therefore see that the artificial membrane formation In order is fatty acid induces the developmental process. In all these eggs in the course of the next hours after the memcell- brane formation those changes begin which lead to a division. of sea-water-f3 c. after this the egg slowly begins to disintegrate. Fig. In 1905 I succeeded in finding a method by which it is possible to call forth the formation of a tion without apparent injury to the egg. valerianic acid. is. or In this mixture no membrane formation takes place. formation and not any other action of the acid. namely. acetic. divisions of the nucleus and of the occur.

After the eggs are transferred brane formation consists in putting these eggs for three hours in sea-water free trace of KCN has is. sugar solution).water they develop often but not always. therefore. e. however. if the larvae are not fed. Delage has. not quite as reliable as the other method artificial mentioned previously. If after this time the eggs are transferred into normal sea-water they develop at room temperature in a wa}' similar to the eggs which are fertilized by sperm. The one which never fails consists in putting the eggs about twenty minutes after the artificial membrane any other rendered formation into hypertonic sea-water (or any other hj-pertonic solution. but sickly development is abnormal and the egg inhibit this sickliness and perishes the more rapidly the higher the temperature. these means The causation this of the membrane formation by in is a fatty acid starts.g.^ Nature of Formative Stimulation call forth 131 the membrane formation not alone by all fatty acids but by a number of different agencies and that act as formative stimuli. arises. into normal sea.e. raised two such larvae until they were sexually mature. 1 The larvae originating from eggs fertilized by sperm live no longer than those originating from eggs which develop parthenogenetically. that the formative stimulus in activation of the egg of the sea-urchin consists of the two phases. therefore. the development the sea-urchin egg. into sea-water or solution the osmotic pressure of which has been 50 per cent higher than that of the sea-water. ... The question how can we and grant a normal development to the egg? I found that two different means are at our disposal for this purpose. This method therefore. at The second method of causing the eggs to develop normally room temperature after the artificial causation of the memfrom oxygen or into sea-water to which a been added. i. We see. The feeding of these larvae is a tedious process and for this reason I have not undertaken the task. In this solution the eggs remain from twenty to sixty minutes —according to the temperature and the concentration of hydroxj'1-ions in the solution.

We will now try to gain some insight into the mechanism of these two agencies. therefore. Asterias found for and I for The experiments on annelids and star-fish. It was noticed that the agencies which cause cytolysis also cause membrane formation. the eggs of annelids made on and of star-fish (Asterina). of Then ensues a pause this sometimes several minutes and after pause a sudden . for We may add that these observations do not hold good Similar observations were (Polynoe) the sea-urchin egg only. especially with saponin. it probably counteracts a secondary detrimental effect connected with the Ill membrane formation.132 The Mechanistic Conception of Life first namely. digitalin. In Polynoe and star-fish the artificial often sufficient to allow the eggs to membrane production is develop into larvae. solanin. result. the artificial causation of the membrane formation of the egg with a and second the subsequent short treatment hypertonic solution. the eggs are treated subsequently Lillie with one of the above-mentioned methods. yielded a curious If the unfertilized eggs are put into a weak solution of first effect saponin in sea-water we notice as the on the eggs the formation of a fertilization membrane. that the calling forth of the membrane is the essential feature in formative stimulation and that the subsequent treatment of the eggs with a hypertonic solution (or an isotonic solution free from oxygen) has merely a corrective effect. solanin. confirm the fact. as R. How can the fatty acid cause the formation to get an answer to this question In order of a membrane? we must find out whether act like there are other agencies which all fatty acids. and soaps. Experiments with these agencies. But the number of eggs is which reach the larval stage and the type if of segmentation improved Polynoe. bile salts. namely. and digitalin. (or a longer treatment with an isotonic solution free from oxygen or containing KCN). first the specifically cytolytic agencies like saponin.

42. 39 Fig. however. 39-43. 41 Fig. 40. Fig. Fig. 40 Fig.e. 41.Nature of Formative Stimulation cytolysis of the whole egg follows. 39. it will develop into a larva. unfertilized egg at the beginning of the experiment. after the membrane formed. urchin egg in a weak solution of saponin in sea-water. and 43. 41. they behave as the . — cytolysis of the egg occurs) out of the sea-water containing saponin and free them from ing all traces of saponin by washif them repeatedly with sea-water. If the egg is taken out of the saponin-sea-water in this stage. but before the d^--V-\-- FiG. The stages represented in Figs. it is left in the saponin solution it undergoes the rapid cytolysis represented in Figs. 42 Fig. The following figvires show the changes it underwent in this solution. If. membrane formation under the influence of saponin. Camera drawings from In this nature. Fig. after it is put back into normal sea-water. condition the egg was put into sea-water containing a small amount of saponin.. washed and put into a hypertonic solution for about one half-hour. cytolysis began at G. 42 and 43 were reached a few minutes later. 43 Membrane formation and subsequent cytolysis of the seaFigs. eight minutes later. five minutes after the membrane formation. this If is 133 we take the eggs during pause (i. In the above drawing of the egg.

e. . calls had alreadv observed that chloroform forth the membrane formation and Herbst had seen the same effect brought about by benzol and toluol. and for this reason these authors could not notice that the membrane formation was followed by the that the development of the egg. i.134 The Mechanistic Conception of Life called forth membrane formation had been nuclear division at by a fatty acid. hypertonic sea-water they developed A further group of cytolytic agencies is ether or alcohols. If. Cytolysis of the eggs by these agencies If is also preceded by a membrane saved from formation. if was the proper cause for the membrane The one membrane formation solution . is The second group specific fat solving of cytolytic agencies like formed by the Hert^vig hydrocarbons amylen. vince myself that for if I have. larvae with skeletons.. but their rather slow and depends on the presence of free ox>'gen. been able to con- amylen or benzol are allowed to act only if one moment and them the eggs are then quickly transferred into normal sea-water a membrane formation can be produced in of some ^vithout subsequent c}i:olysis. Bases can also action is call forth membrane formation. and in a much lesser degree chloroform. however. 44-47). the eggs are taken out from such solutions immediately after cji^ol^-tic membrane formation they can be destruction (Figs. but do not go beyond the room temperature. however. benzol. first Such eggs begin to develop. ^^'ith If such eggs were afterward treated into larvae. if One gains the impression as the alkali acted in this case only as an accelerator of oxidations and as if a product of oxidation formation. toluol. etc. the eggs are treated for half an hour Tvdth hypertonic sea-water they can develop to normal plutei. But these substances act so violently membrane formation is followed almost immediately by a cytolysis of the egg. usually becomes manifest only treats the eggs afterward for a short time with a hypertonic such a treatment causing them to develop into larvae.

45. The cytolyzed Fig. egg has in this case an entirely different appearance from that of an egg cytolyzed in saponin. 45 Fig. 47 Figs. 44 Fig. Fig. Camera drawings. Membrane formation and subsequent cytolysis of the egg under the influence of the addition of a minute quantity of salicylaldehyde to sea-water. 135 increase in temperature can also produce a cytolytic I centrotus purpuratus have observed that at 34° or 35° the eggs of Strongyloform often but not always a membrane Fig. 46 Fig. unfertilized egg at the beginning of the experiment. Fig. — of fertilization. instantly Such a temperature kills these eggs almost and consequently they are no longer able to develop after this treatment. membrane formation in the salicylaldehyde-sea-water. beginning of the cytolysis. 44-47. 44.Nature of Formative Stimulation An effect. Fig. not killed so rapidly after the membrane . 47. The eggs of the star-fish Asterias for- however. 46. cytolysis completed. hesii are.

membrane formation is called forth Von Knaffl has shown that if a into ^'ghosts. I succeeded . When I discovered this fact there was no analogue known which allowed us to draw an inference concerning the mode of action of a hypertonic solution. The destruction of this hj^^othetical substance can be brought about in two ways: first. but that the egg formation. was able to show that such eggs can develop into larvae the by by raising the temperature. This superficial cortical layer of the egg is very thin. while the agencies which do not call forth cytolysis do not cause a membrane formation. that call the agencies which cause c>i:olysis also forth the membrane formation. it still is as a rule sickly after this membrane To fix our ideas provisionally we assume that membrane formation a substance is formed which must be abolished or destroyed before the egg is able to develop If we permit the egg to begin its development while contains this hypothetical detrimental substance in a sufficient quantity it is sickly and dies prematurely.136 The Mechanistic Conception of Life Lillie if formation. therefore. and R. IV We have already mentioned that the lies c5i:olysis which under- the membrane formation causes the development of the egg." all high temperature acts for some time on these eggs they perish c}i:olysis and are transformed We have been able to convince ourselves. we draw the inference that the membrane formaupon the c\i:ol3"sis of the surface layer of the egg. through the normally. by treating the egg for a short time wdth a hypertonic solution. We shall see later on that we must discriminate between a cortical layer and the core of the unfertilized egg. The essential tion depends From feature of the developmental stimulus consists in the cj^tolysis of this cortical layer of the egg and this cytolysis is caused by the spermatozoon. We find in addition that the cytolytic power of these agencies runs parallel with their power of causing this membrane formation.

I succeeded two effects of the we wish to bring about a separation of these two agencies is in the spermatozoon we cannot use the spermatozoa of the same species of sea-urchins from which the egg taken.^ V Thus far we have dealt only with artificial parthenogenesis. for in this 1 case the spermatozoon penetrates at once into the this chapter is A further discussion of the facts in contained in the next paper Fertilization. putting The second method of saving the life it after the membrane formation is of the egg consists in for about three hours into sea-water which practically free from oxygen. it. it the hypertonic effect. We are now about to take up the causation of development by a spermatozoon. inasmuch as it causes the cytolysis of the thin cortical layer of the egg. deprived of oxygen inefficient if remains without any It remains also a trace of KCN is added to it is Since KCN inhibits the oxidations in the cell obvious that the hypertonic solution only acts by a modification of the process of oxidation." Nature of Formative Stimulation in 137 artificial showing that such a solution if it is only effective in If parthenogenesis solution is contains free oxygen. eggs are transferred after this time into normal sea- water containing free oxygen they are often able to develop normally. show that the spermatozoon also calls forth the normal development of the egg by at least two substances and It is possible to that one of these substances acts like butyric acid or saponin in artificial parthenogenesis. or contains a trace of If these KCN whereby the oxidations in the egg are suppressed. while the second substance has an effect similar to that of the hypertonic solution. on " The Prevention of the Death of the Egg through the Act of . Is the formative stimulation of the spermatozoon genesis? of the egg by same character as that in artificial partheno- This question can be answered in the affirmative. correctness for this view in separating these If is The proved by the fact that spermatozoon.

Elder. It takes. way almost simultaneously both membrane substances of the spermatozoon become causes the effective. which the formation. found that a spermatozoon . If we treat them. the cytolytic one. These eggs show at room are dealing with we temperature only the beginning of nuclear division and then disintegrate. all develop at room temperature The other eggs develop without any subsequent treatment with a hypertonic solution normal larvae. The experiments. however. we soon two groups of eggs. to the egg of Under ordinary conditions sperm of the star-fish it cannot cause the egg of the sea-urchin to develop. however. however. star-fish. while after the addition of sea-urchin sperm this result is obtained in one minute.c.water all the eggs of the sea-urchin form fertilization membranes in such a mixture if only a trace of living sperm of a star-fish {Asterias ochracea) is is more n/10 alkaline through the addition of some NaHO. however. The one group acts as if only one of the two agencies. in sea. of sea. mostly from ten to fifty minutes.c.g. some time.. the cytolytic substance. effective. A My Mr. after the membrane minutes formation by star-fish sperm for from thirty to fifty with a hypertonic solution they mostly into normal larvae. If 0. until the living star-fish sperm brings about this effect. If the sea-urchin eggs.138 The Mechanistic Conception of Life it protoplasm of the egg as soon as In this comes in contact with it.6 NaHO added. point. while at a lower temperature they little may develop a farther. becomes little c. and the ''corrective" substance. result differently e. are put if back into normal sea-water and we watch their further fate. all of which have formed membranes upon the addition notice that of living star-fish sperm.water which has been rendered a added to 50 c. had taken effect. when we add spermatozoa of a foreign species. namely. into What of eggs ? causes this difference in the behavior of both groups histological examination of these eggs decides this assistant. the sea-urchin.

since in the formation of the membrane the spermatozoon is thro\\'n out. only if A normal development will result the spermatozoon enters the egg since in this case only both agencies. This substance is probably situated at the only calls spermatozoon. enough of the lysin sticking to the surface of the spermatozoon can be dissolved to cause the cytolysis of the surface film of the egg which gives rise to the membrane formation. one a so-called lysin. Such eggs receive from the spermatoas if zoon only the and they act therefore only the mem- brane formation had been called forth in them by the treatment with but>Tic acid. surface of the of these agencies is a cytolytic substance. the cytolytic and the corrective. a spermatozoon pene- trates partially through the surface of the egg without entirely penetrating into the protoplasm. lysin. while the eggs which behave of star-fish as if only an artificial membrane formation had taken place do not contain any spermatozoon. This lysin forth the the membrane formation and it acts like the but>Tic acid in method of artificial parthenogenesis. This fact is portance to disprove the possibilit^'that the membrane formation .Nature of Formative Stimulation 139 had entered into those eggs which develop after the addition sperm without subsequent treatment \\'ith a hypertonic solution into normal larvae. get into the egg. In the eggs of Strongylocentrotus purpuratus the memin the of im- brane formation can in general only be called forth by living star-fish sperm while the extract of dead star-fish sperm same concentration remains without effect. The second agency it seems to be more in the interior of the spermatozoon and exercises an influence similar to the short treatment of the egg with a hypertonic solution. We have already mentioned that foreign spermatozoa If penetrate only slowly into the egg. This behavior of the eggs under the influence of foreign sperm is comprehensible under the assumption that the spermatozoon also causes the development of the egg through two agencies.

In this case only the lysin entered the egg but not the It was. they are treated afterward with a hypertonic solution they develop into normal plutei. We know cells of that blood contains lysins species. was able to show three years ago that the blood of certain . however. sensitive to lysins. or Such eggs can be caused to develop into plutei by after the treat- them membrane formation for a short time with hypertonic sea-water. Such eggs act as membrane formation with the aid of butyric acid had been caused. while If which destroy the blood corpuscles of foreign it does not destroy the the same species. therefore. I have called forth only the membrane formation of sharks or in the sea-urchin egg with the living if sperm even roosters. necessary to treat such eggs spermatozoon. VI The idea that a lysin contained in the spermatozoon is the formative stimulus which causes the egg to develop can be tested experimentally.140 The Mechanistic Conception of Life star-fish was caused by added with the sperm. ing of star-fish dilute watery extract of the it which was killed by heating to 60° C. If. At room temperature they begin to develop but they are sickly and soon perish. the idea is correct that the spermatozoon acts lysin upon the egg through a which calls forth the possible to call forth the lized I membrane formation it should be membrane formation in the unferti- egg of the sea-urchin by foreign blood and such is the case. That it is possible to separate a in these experiments blood which was lysin from the sperm can be proved for the eggs of another species of sea-urchins. the eggs of which are very it is In these eggs possible to call forth the membrane formation with a very sperm more. Strongylocentrotus franciscanus. subsequently with hypertonic sea-water in order to cause them to undergo normal development at room temperature. In the place of star-fish sperm the sperm the of other foreign species can be used. namely.

The lysin of the blood is like that of the spermatozoon lose its relatively resistant to heat. therefore. They began to develop. and rabbits. CXXIV. . and such eggs which had been treated with the living sperm of roosters or with butyric acid. If. I succeeded in causing membrane formation in the behaved like the eggs sea-urchin egg (purpuratus) with the blood serum of cattle.Nature of Formative Stimulation worms. sheep.^ curious that SrCl. development. 37. and Babl2 increase the membrane-forming power of the blood. namely SipuncuUdes . I think the difference differences in the permeability of the eggs for lysins. 141 can call forth the membrane formation in the sea-urchin egg even times or more with sea-water. but they became sickly at room temperature and soon disintegrated. however. Not only blood but also the extracts of the organs of foreign 1 Since this was written the blood and the extracts of organs of a number of invertebrates were used successfully to produce the membrane formation and development of the egg of the sea-urchin. contains but not the second substance necessary for the It is. necessary to substitute for the action of the latter the treatment with a hypertonic solution if we wish to call forth a normal development of the egg treated with serum. Pfingers Archit. therefore. and the degree of permeability seems to vary slightly for the eggs of different females. the lysin. 2 The substance which causes membrane formation can be precipitated with acetone (Loeb. if it is diluted a hundred is This effect not produced caused by in the eggs of every female sea-urchin but of only about 20 is per cent of the females. 1908). pigs. they were treated after the membrane formation for a short time with a hypertonic solution they developed at room temperature. full The blood. The blood does not to call forth membrane formation by heating it It is for power some time to 60° or 65° C. Instead of wasting time on an examination of the effects of the blood of invertebrates^ I examined the effects of the blood serum of warm-blooded animals.

therefore. but not against those of foreign species. an immunity of the eggs as well as of the the lysins of their rest of the cells against own blood or tissues. The extract from the dead sperm of the sea-urchin is ineffective for the eggs of the sea-urchin solely for the reason that it cannot diffuse into the sea-urchin egg. e. or the cells contain antibodies against the lysins of their own body. causes membrane formation in the eggs of franciscanus. our cells it upon the nature of this If the lysins contained in our blood do not injure can only be due to one of two facts: The lysins of either not diffuse into our cells. is For if the sea- urchin lysin carried by the living sea-urchin spermatozoon (which acts as a motor for the lysin) into the sea-urchin egg. sharks.142 The Mechanistic Conception of Life membrane formation in the sea-urchin egg. . 1 If eggs were sensitized with SrCl. Experiments with the extract of dead sperm of their owti species on the egg oi franciscanus or purpuratus fail. certain worms.g. fowl. though this was trueonlj. species call forth coecum of the star-fish was very effective. however. the lysin is very active and probably more active than the If lysin of foreign species...^ What causes this difference in the action of the lysins from their that the lysins of own and a foreign species? our own blood do not hurt our There We know cells while they hurt the cells of foreign species. of star-fish. light Our experiments throw a immunity.exceptionally. the sea-urchin egg contained an antibody against the lysin of the sea-urchin sperm. the seaurchin sperm should not be able to it call forth membrane formation when enters the sea-urchin egg. As far as the lysins of the blood are possibilities. exists. while they cells of foreign our own blood can can diffuse into the forms. certain mollusks. concerned we cannot decide between the two We can. and the same is true for extracts from the tissues of extract of the An these species. they could be caused to develop by extracts from the coecvun of the sea-urchin. reach a decision for the lysins of the sperma- tozoa. We have already mentioned the fact that the extract of dead sperm of foreign species.

Nature of Formative Stimulation

143

We now understand the paradoxical fact, that by foreign sperm we can cause membrane formation and development of the sea-urchin egg in two different ways: namely, first by the living sperm and second by the extract from the dead sperm;
while the sperm of the same species can only cause the eggs to

develop when

it is

alive.

We now

understand the fact alluded

to at the beginning of this chapter that

my

first

experiments

to cause the development of the egg with extract of sperm did

not succeed, since

I

took

it

for

granted that

it

was necessary

to use the extract of the sperm of the same species from which

the eggs were taken.
diffuse into the egg.

The

lysins in this case

were not able to

The
of the

further unraveling of the nature of the

egg-cell against the dissolved lysins of the blood

immunity of the and the tissues
of the fact

same

species depends

upon the explanation

that the lysins of a species cannot diffuse into the egg of the

same

species.

formed

for the

It would be of interest if the same principle immunity of the egg-cell would hold also for the

immunity

of the body-cells against the lysins in the blood of

their o%vn species.

which the power is a lysin and we may express the suspicion that the lysins which we have thus far kno\\TL
therefore, say that the substance to

We

may,

sperm owes

its fertilizing

only as protective agencies against bacteria play a great physiological role in the

mechanism

of

life

phenomena.

We may

call

our theory of the developmental action of the spermatozoon
the lysin theory;

thereby designating that the impulse for the
is

development of the egg
spermatozoon.
In

given by a lysin contained in the
parthenogenesis

artificial

we

substitute for

the natural lysin a cytolytic substance.

Aside from the lysin
(but not

action the normal development demands, as a rule

always), a second corrective influence which in artificial par-

thenogenesis

may be

given by a hypertonic solution.

;

144

The Mechanistic Conception of Life
VII

The experiments on the artificial parthenogenesis of other forms of animals show that the eggs of different animals possess
a varying tendency for parthenogenetic development.

There

are eggs which can easily be induced to develop, so easily in
fact,

that the experimenter cannot always be sure whether he

has caused the development by a substance applied by him or

whether some accidental condition of the experiment was
responsible.
of certain

The eggs of the silkworm, of the star-fish, and worms belong to this class. In working with starfish eggs we can observe that occasionally a few of them develop
normal sea-water, apparently without any demonstrable

in

cause, into

swimming
slightest

larvae.

The eggs

of the

Californian

sea-urchin Strongylocentrotus purpuratus,

on the other hand,

show not the
causes

tendency to segment parthenogenetically

only the above-mentioned very specific and quantitative method

them

to develop.

For

this reason I selected these eggs

for the investigation of the nature of the experimental stimulus,

since I could always be sure that the

same

results;

while, e.g., in the star-fish eggs

perfectly certain that

some

internal

same stimulus gave the we can never be condition in the egg or some
Although eggs with

overlooked unimportant secondary condition in the experiment

may

not have caused the development.

such a strong tendency for spontaneous development as the
star-fish eggs are

not the best material for the study of the

nature of the developmental stimulus yet
the question

we have

to answer

how

it

happens that some eggs have a greater

tendency for parthenogenetic development than others. Mathews has observed that by gently shaking the star-fish
eggs the

number
I

of eggs

which develop "spontaneously" can
In the eggs of the sea-urchin nobody
I

be increased.

made

a similar observation in the eggs of

Amphitrite, an annelid.

has ever been able to obtain such a result.
believe that
if

am

inclined to

a sea-urchin should be found, the eggs of which

Nature of Formative Stimulation
possess a greater tendency to develop spontaneously,
also be found that the
it

145

might

number

of eggs developing spontaneously

might be increased by agitation.
I tried

whether

it is

possible to cause the eggs to cytolyze
exercise only a slight pressure with

also mechanically.

If

we

a finger upon the ovary of a star-fish

we

find that

many

of

the eggs which afterward leave the ovary are cytolyzed.
cytolysis
is

This

not caused by a bursting of the egg membrane; on
is,

the contrary, in this case the cytolysis of the egg

as usual,

preceded by the formation of a membrane of fertilization and
this

membrane remains
it is

intact in the star-fish egg which

is

caused to cytolyze
egg, however,

by mechanical

pressure.

In the sea-urchin

impossible to produce cytolysis

by a

slight

pressure.

The eggs
fish

of the star-fish

form a membrane.
eggs only
if

which develop spontaneously first Shaking causes a development of the starfirst

the-shaking

leads to a

membrane formation.

The
is,

greater tendency of the star-fish to develop spontaneously

therefore,

due to the greater ease with which cytolysis can
call forth

be produced in this egg.

How

can mere agitation or pressure
It

membrane
is

formation or cytolysis ?
easily understood
Biitschli, that

seems to

me

that this fact
first

most

under the assumption,
is

suggested by
It

the cytoplasm

an emulsion.

would then

follow that the

membrane formation

as well as the cytolysis

depends upon the destruction of this emulsion.

We know that

The different emulsions have a different degree of durability. have an eggs which upon gentle pressure undergo cytolysis
emulsion with a lesser degree of durability than the eggs in which
pressure has no such effect.

Let us assume that membrane

formation as well as cytolysis depends upon the destruction
of

an emulsion;

in this case the

membrane formation depends

upon the destruction of the emulsion in the cortical layer of the egg only. The lysin of the egg destroys only the emulsion in

146

The Mechanistic Conception of Life

the cortical layer of the egg and thereby causes development.

The

greater tendency of the eggs of certain animals for spon-

taneous parthenogenetic development thus depends upon the
relatively small degree of

durability of the emulsion which

constitutes the cortical layer of the egg.

But

it

should be

stated that this hypothesis

is

not essential for the lysin theory

of the activation of the egg.

VIII

The assumption that the membrane formation
layer of the egg
Biitschli

is

only a

superficial cytolysis of the egg presupposes that the cortical
is different from the rest of the cytoplasm. had already reached such a conclusion on the basis I am inclined to accept this view histological observations.

of

on the basis

of

my

observations on the action of cytolytic

agencies on the unfertilized egg.

The

action of these agencies

on the unfertilized egg always occurs in two stages which are often separated from each other by a considerable interval of
time.

The

first

stage
is

is

the cytolysis of the superficial layer;

the second stage
is

the cytolysis of the rest of the egg.

This

most obvious

in

experiments with weak solutions of saponin

or solanin in sea-water.

In this case

first

a

membrane formaand

tion occurs, then a pause ensues, often of several minutes,

then cytolysis of the whole egg follows.
benzol
is

If instead of saponin

used a pause can also be observed between membrane
is

formation and cytolysis of the whole egg but this pause

short,

often only a fraction of a second, or at the best a few seconds.
It

can also be shown directly that there
the

is

a qualitative

difference between the cortical layer of the protoplasm and the
rest. If for
artificial

membrane formation
is

the lower fatty

acids,

from the formic to the capronic
observed,

acid, are used, cytolysis

of the cortical layer only

i.e.,

membrane formation
If,

follows but

no cytolysis

of the

whole egg.

however, the

higher fatty acids of the same series

from the heptylic acid on

" We can accept this with a slight modification which refers to the nature of the emulsion. membrane formation and development but not a cytolj'sis of the whole egg. general cytolytic agencies destroy the cortical layer as well as the rest of the egg.Nature of Formative Stimulation 147 and upward are applied the membrane formation is always followed after a short pause by a cytolysis of the whole egg. IX How to a can the cytolysis of the cortical layer of the egg lead membrane formation? Von Knaffl has expressed the is following view on this point: it ''Protoplasm rich in lipoids. is probably mainly an emulsion of these and of proteins. The droplets of the emulsion are surrounded by a thin layer of a substance which lessens the surface tension between the . The protein of the is egg can only swell or be dissolved altered if the state of the lipoids by chemical or physical means. The third substance serves the purpose of making the emulsion more durable (Lord Rayleigh's theory). Every physical and chemical agency which is able to liquefy lipoids calls forth a cytolysis of the egg. from that of the There are certain cytolytic agencies which while all destroy only the cortical layer. The lysins contained in the blood and the spermatozoon act according to my present experience only upon the cortical We get a layer of the cytoplasm but not on the rest of the egg. If we go back to the idea of Butschli that protoplasm has the structure of an emulsion we are led to the view that the differs emulsion of the cortical layer of the egg rest of the egg. An emulsion requires not only in addition two substances or phases as von Knaffl assumes but a third substance. The mechanism of cytolysis consists in the liquefaction of the lipoids and the subsequent swelling or liquefaction of proteins by absorption of water This confirms Loeb's view that is membrane formation caused by the liquefaction of lipoids.

We I is imfertilized egg it. The existence of these two phases has been established by Hardy. although this does not belong to our problem. To fix our ideas provisionally we may assume that these phases are first protein with little water and second water with little protein. According to Overton and Koeppe the surface film of consists of lipoids. The emulsion cholesterin. . this is absolutely insoluble in any lipoid solvent. since the surface film which is lifted the fertilization protein. I assume that only this stabilizing substance consists of lipoids.^ We wish to add a few remarks concerning the nature of this surface film." perfectly intact when the egg transformed X Since we can cause the formation of a membrane of fertili- zation in the star-fish egg 1 by gentle agitation or mere pressure. especially The two other phases which constitute the emulsion need not be lipoids. This view off in not the form of This is membrane does not consist of a lipoid but of suggested by the fact that this membrane is Moreover.148 droplet The Mechanistic Conception of Life and the second phase of the emulsion. membrane remains into a ''ghost. according to this view. It is immaterial for the problem discussed in this paper which view we adopt temporarily. it membrane is a membrane tion formed through the reaction of a constituent of the liquefied of precipitacortical layer with a constituent of the sea-water (Ca?). If we use saponin destroyed by the precipitation of cholesterin tion of water leads to the lifting by saponin. If the sea- urchin egg is treated with a lipoid solvent like benzol the is stabilizing film of cholesterin let dissolved and the protein dropthe film is can absorb water. of a system of protein droplets poor in water surrounded by a stabilizing film of a lipoid (cholesterin or lecithin). As have assumed here that the fertilization membrane is preformed in the and lifted up in consequence of the cytolysis of the layer beneath stated in my book on Die chemische Entwicklungserregung des tierischen also possible that the fertilization Eies. The absorpup of the surface film which surrounds the egg. cells and according to Koeppe cytolysis is deteris mined by the solution or tearing of this film. at the surface of the egg consists. tenable.

concertain quantity taining eggs with a fertilization membrane. That merely a change in the nature of the surface film prevents the entrance of a spermatozoon into the egg after the membrane formation can be proved by the fact that the surface film.Nature of Formative Stimulation this 149 membrane if is apparently preformed in the unfertilized egg. has different qualities or a different structure when it is in close contact with the cytoplasm than when it is lifted off from the cytoplasm by a layer of sea-water. therefore. The correctness of this view can be proved. if it is lized egg. . even if we tear the membrane mechanically This proves that already preformed in the unferti- a spermatozoon can penetrate into the egg. In this process the surface film undergoes a change. The membrane is. and is thereby dissolved. membrane The reason that almost the liquid which existed between the mem- brane and the cytoplasm diffused into the surrounding seawater. We have assumed that the membrane formation is determined by the action of a lysin or cytolytic agency upon the cortical layer of the egg. a of dissolved white of egg. we add to the sea-water. impermeable for the colloids dissolved in sea-water. whereby a protein in this layer absorbs sea-water. (free If the eggs are brought back into normal sea-water it from protein) diffuses again into the space between the membrane and its the cytoplasm. or blood serum. and this be true the process of membrane formation must consist in the lifting up of a preformed film from the underlying cytoplasm through the entrance of sea-water between this film and the cytoplasm. and the fertilization mem- brane resumes its former distance from the cji^oplasm and round shape. the collapses is and closes tight all around the cytoplasm. This assumption leads to two consequences: first it must be possible to show that the fertilization membrane is permeable for sea-water and crystalloid substances but impermeable for If colloids. since the spermatozoon can enter into the egg before but not after the membrane formation. tannin.

membrane is permeable for salts. This may be due to the fact that the degree of swelling of the colloid of the cortical layer varies under different conditions. of that layer is able to absorb water and swell. Through a lysin or some other cytolytic agency a certain substance of the cortical layer. but not for colloids. to the egg in the case of ''formative stimulation. it We see. and that the lifting up of the fertilization membrane is determined by the swelling and subsequent liquefaction of a colloid. This dissolved colloid exercises an osmotic or colloidal pressure and sea-water must diffuse from the outside imder the fertilization membrane of the egg until the tension of this membrane equals the osmotic or colloidal pressure of the dissolved colloid. . Others expressed the opinion that the entrance of the sperm nucleus or lipoid. however. XI We now possess is a pretty complete picture of what happens i. of a centrosome was responsible for the development. presumably a whereby a protein substance Formerly it development the caused was thought that the spermatozoon of the egg by carrying a ferment or enzyme into it and that this ferment set the mechanism of development into action. We now can understand why not in is all cases of fertilization a distinct fertilization membrane formed. in order to observe after two or three hours the formation is of normal astrospheres or egg.150 The Mechanistic Conception of Life If salts are added to the sea-water or if it is diluted by the addition of distilled water the tension and the diameter This proves that the of the membrane do not change. explains also This why it is that the fertilization membrane as a rule assumes a spherical shape." when it caused to develop.^ spindles.e. that suffices to call forth the artificial membrane formation in the unfertilized sea-urchin egg. is dissolved or precipitated.. This disproves the suggestion that the fusion of egg essential for the is and sperm nucleus 1 development of the The fusion of the nuclei of course of importance for the transmission of paternal qualities.

all cytolytic substances are lipoid soluble (or destroy it is probable. The spermatozoon as well as the blood and the tissues contain a substance (lysin) which causes only cytolysis of the cortical layer. or fertilization parthenogenesis are superposed in two spermatozoa by sperm and the same egg. But if this is not the case. cause only the cytolysis of the layer. The lower fatty acids.^ tion The further development will be connected with the queshow can the cytolysis of the cortical layer of the egg lead development? I to its may mention the possibility that the cytolysis of the cortical layer facilitates the diffusion of oxygen or of HO ions (bases) or other substances. . were correct the velocity of developif ment should be accelerated artificial not doubled if enter the egg instead of one. no matter what is the nature of the spermatozoon. into the egg. Any agency which causes formative stimulation of the egg. Cytolysis of the cortical layer leads often but not always to the formation of the Since lipoids) 1 membrane of fertilization. In neither case is a shortening of the time which elapses between two successive periods of segmentation observed. cortical from formic to capronic. at least this cytolysis (without causing the cytolysis of the rest of the egg) induces development. Since most cytolytic agencies cause a cytolysis of the if whole egg they can be used only the eggs are withdrawn is from their influence after the cortical layer destroyed but before the rest of the egg has undergone destruction.Nature of Formative Stimulation The ferment theory tozoon is 151 of the activation of the egg If it by the sperma- also wrong. XII Let us summarize our results concerning the activation or For the normal development two agencies are required: the one is the cytolysis of the thin cortical layer of the egg. necessary for the development. that the formative Another reason is that the velocity of segmentation is purely determined by the egg. but not proved.

a short treatment of the egg with a hypertonic solution containing oxygen or a longer inhibition of the development of the egg in normal sea-water which is free from oxygen. according is Lord Rayleigh. The cytolysis of the cortical layer of the egg causes its is development. which acts similarly to the hypertonic solution in our method of artificial parthenogenesis . the mode of action of which is not yet so clearly understood as that of the cytolytic agency. 152 The Mechanistic Conception of Life stimulus in the activation of the egg consists in a liquefaction or precipitation or cortical layer of the some other modification of the lipoids of the egg which results in an imbibition or soluIf tion of a colloidal substance of the cortical layer.. The spermatozoon carries in addition to the lysin a second substance into the egg. namely. but this development often abnormal and comes prematurely to a standstill. necessary for the durability of the emulsion. In order to induce a more normal is type of development a second agency often required. the cyto- plasm has the structure of an emulsion it is possible that lipoids form the to stabilizing envelope for the droplets which.

VIII. THE PREVENTION OF THE DEATH OF THE EGG THROUGH THE ACT OF FERTILIZATION .

.

VIII THE PREVENTION OF THE DEATH OF THE EGG THROUGH THE ACT OF FERTILIZATION^ I The unfertilized egg dies in a comparatively short time. fertilized. that is to say. 155 . I found that the death of the egg can be prevented by withdrawing the oxygen. a life-saving act for the egg. If they remain. of fertilization is. theoretically at least. I Ten years ago made the following observations. in which life way can the spermatozoon save the of the egg If ? the ovaries of a star-fish are put into sea-water the eggs they cannot be and in this condition by spermatozoa or by chemical means. either They so-called polar bodies. the by sperm or by physico-chemical agencies. 1 Reprinted from the Harvey Lectures. Lippincott & Co. The death of the egg manifests itself morphologically in a darkening and blackening eggs are not caused to develop of the otherwise clear egg. all or a number of them gradually become mature. The question arises. At summer temperature they may die in from four to six hours. If is immediately after the extrusion of the polar bodies sperm added. by courtesy of Messrs. or by diminishing the rate of oxidations in the egg through the addition of a trace of potassium cyanide. The act therefore. the eggs develop. their nuclear mass is diminished by the extrusion of the two are shed. The life-saving action of lack of J. 1911. while the act of fertilization gives rise to a series of generations which. at that period likewise be caused to develop They can by certain chemical If and physical agencies. is of infinite duration. for some time in sea-water. are generally immature. however. they perish very rapidly. B.

by putting them into a narrow glass tube which is sealed at the bottom.156 The Mechanistic Conception of Life in various ways. or tion does not take place. however. P. since large masses of the oxygen which diffuses from the air through the sea-water is consumed by the uppermost layer of the eggs. If oxygen can be shown egg itself The maturation the ox. the process of matura- Maturation of is. This opposition was based on the fact that potassium cyanide was used in part of the experiments. in sea-water in contact with It is not even necessary to drive out the air the life of the unfertilized eggs can also be preserved by hydrogen. death does not occur. If the oxygen is withheld from the eggs immediately after they become mature their life is also saved. A. The eggs sink to the bottom of the tube. hence by withholding ox^^gen from the immature eggs their maturation and death are prevented. and those which are lying near the bottom receive no oxygen. The eggs of the second group remained alive.y^gen is of the depends upon oxidations. unripe. overlooked the fact that lack of oxygen acts in exactly the same w^ay as the addition of potassium cyanide. the oxidations in the immature eggs are inhibited by potassium cyanide. The objection was raised that the potassium cyanide in these experiments acted only by preventing the development of bacteria. were divided into The eggs a female. which were two groups: the one group remained oxygen. the other was put into sea-water whose oxygen had been removed by a current of hydrogen. . also a function of oxidations. This proves that the death of the mature but unfertilized egg determined by oxidations. therefore. who raised this objection. On account of this lack of ox^^gen the eggs at the bottom of the tube do not mature and do not perish. If these oxidations are inhibited first When these experiments were published they caused opposition. if withheld from the immature eggs. the eggs of the first group perished in a few hours. Mathews is has repeated this experiment and obtained the same results. The authors.

however. certain that the death of the star-fish eggs fertilized is which are not egg. of the heap receive oxygen. It is. Warburg found for the eggs of the sea- urchin at Naples that fertilization raises the velocity of the process of oxidations to six times their original value. but by the If process of oxidation in the egg. a spermatozoon enters the egg.Prevention of Death by Fertilization and that it is 157 is entirely immaterial is how lack of oxygen pro- duced. The sterilization of the eggs of the first group was complete. the latter remains alive in spite of the fact that the entrance of the spermatozoon causes an acceleration of the oxidations in the egg. The following experiment The eggs of a star-fish were separated into three parts: one part was put the aseptically into a series of flasks with sterilized sea. the third part was put into sea-water to which a large quantity of a putrid culture of bacteria had been added that had star-fish. It was found that mature eggs died within the same period of developed on the dead eggs of the in all three cases the time. while the dead eggs in the normal sea-water were completely destroyed in a few days by the action of the bacteria. as was shown by the fact that the eggs although dead preserved their form for two months.water. second part was put into ordinary sea. while Wasteneys and I found that fertilization caused an increase to three in the velocity of oxidations of Arbacia in Woods Hole or four times the rate found in the unfertilized eggs. if no spermatozoon enters the the egg is not caused to develop by chemical treatment If. hydrogen or whether the eggs are put together whereby only those lying on the surface sufficient It is. or it not caused by bacteria.water without asepsis. perishes very rapidly. incorrect. The eggs was of the star-fish can put into sterilized sea-water without bacterial infectried. therefore. easy to show directly that the aboveis mentioned objection easily be tion. whether the oxygen driven out by carefully purified in a large heap. however. .

differences in regard to the above-mentioned conditions. w^e j&nd great fatal effects of the oxidations. the substances which determine the heredity of paternal characters and the substances which cause the egg to develop. In this paper we are concerned only with the second group of substances. analysis of the process of fertilization by the spermato- zoon shows that we must discriminate between two kinds of the hereditary effect and the activating or developeffect. mental The experiments on artificial parthenogenesis make it very probable that the two groups of substances. or if some faulty combina- tion of conditions which. The problem as to what determines this difference has not yet been investigated. namely. The analysis of the causation of development of the egg by a spermatozoon has shown that the latter acts by carrying at least two substances or groups of substances into the egg.: 158 The Mechanistic Conception of Life How The can we explain the fact that fertiHzation saves the life of the egg ? Let us make the following preliminary assumption unfertilized egg contains a poison. those which cause the development of the egg. causes the death of the egg. In the unfertilized but mature egg oxidations carries The spermatozoon into the egg among other substances something which protects the egg against the and allows them even to carry on We might say that the mature but unfertilized egg is comparable to an anaerobic being for which oxidations are fatal. while the eggs of the sea-urchin remain alive for a longer period of time after they have become mature. If we compare the eggs of different animals. oxidations take place. oxidations at an increased rate without suffering. . II The effects. if The they eggs of certain annelids (Polynoe) also perish rapidly become mature without being caused to develop. take place. and that the spermatozoon transforms the egg into an aerobic organism. are entirely different.

in a have shown number of papers that the essential feature in the causation of the development of the egg is a Fig. it begins to develop. If we cause membrane formation in an unfertilized sea-urchin egg by artificial means. Disintegration of the sea-urchin egg after the membrane formation with butjTic acid or with foreign serum or with the extract of sperm or of foreign cells.Prevention of Death by Fertilization The first 159 of these substances causes the formation of a mem- brane. — modification of its surface. if the eggs are not treated after the membrane formation with a hypertonic solution or a suppression of oxidations. A indicates the area where the disintegration begins. 48 Fig. I was able to show that . the second serves the purpose of rendermg the egg immune I against the fatal action of oxidations. much more rapidly than if it is not exposed to any treatment. 50 Figs. 48-50. which in many cases leads to the formation of a membrane. but very soon perishes. 49 Fig.

causes the rapid death of the egg through an acceleration that the of oxidations. gastrula. c. they develop after being transferred to normal sea-water in the same way as if a spermatozoon had entered them. for a short time with a hypertonic solution. The membrane this rapid formation. to a second treatment. artificial Warburg has membrane formation in the recently shown unfertilized sea- urchin egg causes the same increase in the rapidity of oxidations as the entrance of a spermatozoon. therefore. can be prevented either formation. but that these oxidations lead to the rapid decay of the eggs at room temperature for the reason that the egg contains a toxic substance. or a toxic complex of conditions. which in the presence of oxidations leads to the rapid death of The second treatment serves the purpose of rendering the egg immune against the toxic effects of the oxidations. no longer the hypothesis that the membrane formation causes a rapid increase of the oxidations in the egg and in this causes it way to develop.c. and if we afterward treat the eggs the egg. develop into a blastula. pluteus. they are put perish. I ventured artificial hypertonic solution of a certain osmotic pressure (for instance. n/2j NaCl) after this time. of sea-water +8 c. and the larvae produced from eggs fertilized by a spermatozoon. If we first cause the artificial membrane formation in the unfertilized egg by any of the various means which I have described in former papers. we have to subject them about 50 If.c. after artificial membrane by withdrawing the oxygen from the egg or by inhibiting the oxidations in the egg by the addition of a trace of potassium cyanide.water they but develop into normal larvae. If we wish to cause the unfertilized eggs to develop to the pluteus stage after the membrane formation. . live as long as They reach the and successive larval stages. back into normal sea. fifteen This may consist in putting them minutes after the membrane formation into a for one-half to one hour.160 The Mechanistic Conception of Life death of the sea-urchin egg.

namely. however. fertilize its For reason: this purpose we must the egg of the sea-urchin with a sperm different from own. we it fertilize the sea-urchin egg with the sperm fifty of star-fish. of the egg by two different substances. the deprivation of the egg for three hours from oxidations. or the treat- ment with lack larvae. takes from ten to minutes to cause the membrane formation star-fish in the eggs. however. after the treatment with the hypertonic solution for half an hour. by adding a trace of potassium cyanide to the solution. the reason bemg that the sperm can penetrate only very slowly into the egg of as a rule. It is. second treatment consists in abolishing the faulty condition which makes oxidations fatal to the egg. not possible to fertilize the egg of the sea- the sea-urchin. urchin by star-fish sperm in normal sea-water. for this treatment another treatment. the eggs are put back into normal sea-water they can develop into normal We can show that the spermatozoon also causes the development of the egg by two different agencies comparable in their action to the agencies used in the methods of chemical fertilization which we have just described. either by removing the oxygen from the solution or If. and for the following causes the The spermatozoon it is of the sea-urchin enters so rapidly it into the egg that impossible to show that development If.Prevention of Death by Fertilization Hence the physico-chemical activation one is 161 of the unfertiUzed egg of the sea-urchin consists of two kinds of treatment. We can substitute. But I found . change causes the acceleration of oxidations which in is my opinion The the essential feature of the process of fertilization. The a change in the surface of the egg which may or may This not result in the so-called formation of the membrane. of oxygen for about three hours. This second treat- ment may or a little consist in exposing the eggs for about half an hour more to a hypertonic solution.

the best results were obtained when 0.c.6 c. of sea-water.c. the one kind of eggs behave as if they had been fertilized with sperm of their own kind. If we examine the eggs of a sea-urchin which have been treated in an alkaline medium with the sperm of the star-fish. in about fertilization fifty minutes the eggs form the typical If membrane. Through the act of . is one of the substances of the spermatozoon dissolved in the superficial layer of the egg and causes the membrane formation. we watch by the further development of sea-urchin eggs sperm we notice very soon that there are two different kinds of eggs present. star-fish penetrates am inclined to draw the following of the conclusion from these observations: The spermatozoon some time very slowly through the surface film of the sea-urchin egg. however. of n/10 NaOH were added to 50 c. and this percentage seems to be identical with the percentage of the eggs which develop into larvae. with the sperm of Asterias. I no sperm nucleus can be found inside of them. That is to say. which form only a disintegrate. For the fertilization of the Californian sea-urchin. however.162 The Mechanistic Conception of Life if eight years ago that alkaline than fertilized it we make the is sea. When it lingers for partially imbedded in the surface film. but at ture they slowly perish room tempera- by cytolysis. with active sperm. Strongylocentrotus purpuratus. these eggs begin to segment. If. these eggs are treated for half an hour with a hypertonic solution they develop into larvae.water slightly more naturally the eggs of the sea-urchin can be by the sperm of the star-fish. The other kind of eggs. they segment regularly and develop into swimming blastulae and gastrulae. As far as the other membrane and then eggs are concerned. all In this case. we find that only a certain percentage of these eggs contain the sperm nucleus. act as if they had been treated with one of the agencies which cause the membrane formation in the unfertilized seafertilized star-fish urchin egg.

immmie against Ill Seven years ago the fertilized egg I fomid that a number of agencies destroy rapidly than the unfertilized egg. for instance. line solutions of If we use slightly alkaof the sodium chloride the greater resistance . while in a pure sodium chloride solution the unfertilized egg of the Californian sea-urchin may be kept alive for several days. While the membrane-forming substance of the spermatozoon may be situated at its surface.Prevention of Death by Fertilization 163 membrane formation the further entrance of the spermatozoon mto the egg is prevented. The latter. much more Thus. a by the spermatozoon is by two different substances. These a data support the assumption action of the spermatozoon is made above that the life-saving it due to the fact that carries substance into the egg which renders the latter the toxic action of oxidations. the second substance which transforms the egg from an anaerobe into an aerobe must be situated it in the interior of the sperma- tozoon. the fertilized egg is destroyed in such a solu- tion in less than twenty-four hours. since can only act if the spermatozoon penetrates into the egg. one of which causes the membrane formation. contains a toxic substance. This membrane formation leads to an increase in the rate of oxidations and the beginning of the de- velopment of the egg. concerning the fertilization of the sea-urchin egg by the star-fish sperm. while the second renders the egg proof that the activation of the egg also caused immune against the toxic action of the oxidations. The spermatozoon carries a second substance into the egg which renders it immune against the fatal actions of the oxidations. since the fertilization membrane is impermeable to sperm. or superficially at least. however. or a faulty complex of conditions which has to be abolished. before the oxidations necessary for development can take place T\dthout the egg being destroyed by them. We see in these observations.

Or of phenylurethane. . suppression of the oxidation saves the 1 This observation does not agree very well with the assumption that the narcotic action of these substances is due to a retardation of oxidation. cytolyzed in a neutral unfertilized egg sodium chloride solution in a few hours. or of a solution of chloral hydrate. or a solution of sodium + calcium. When we the and unfertilized eggs into hypertonic solutions. have been able to show for a number of toxic solutions that their effect can be either completely annihilated or at least diminished if we take the oxygen away from the solution. is greater than that While this is probably to some extent true. is The egg of the Atlantic form of sea-urchin. or at least partially. while the may put live for fertilized a considerably longer period of time. for instance. can be kept alive for a considerable period of time in we either carefully remove the oxygen from the solutions. while a of the egg. or if we diminish the rate of the oxidations in the eggs by adding a trace of sodium cyanide. Arbacia. In this case we have the direct proof that solutions which are fatal for the egg when the same solutions the oxidations are allowed to go on are rendered completely.+ 164 The Mechanistic Conception of Life is unfertilized egg perhaps still more striking. What causes this difference It sensitiveness between of the fertilized and unfertilized eggs? is possible that the permeabihty of the fertilized eggs unfertilized. we find also that the fertilized eggs suffer much more than of is unfertilized. Not only the toxic action of salt solutions upon the fertilized egg could be inhibited by the suppression of the oxidations in the egg. harmless if we stop the oxidations in the egg.^ These observations prove directly that of in the presence certain toxic substances or mixtures of substances its the oxidations in the egg lead to rapid life destruction. but also the toxic action of sugar solutions. yet it not the whole explanation of the difference in the I behavior of the two kinds of eggs. Thus. or a solution if of sodium barium. or of solutions of alcohol in the sea-water. fertilized eggs of the sea-urchin which perish very rapidly in pure salt solutions.

in addition to the membrane-forming substance. or group of substances. caused by the and that the life- saving action of the spermatozoon consists in the fact that the latter. carries a second substance.Prevention of Death by Fertilization 165 We therefore believe that we may conclude is that the rapid death of the unfertilized egg of certain species oxidations which take place in these eggs. into the egg which renders it immune against the harmful effect or consequences of oxidations. .

.

IX. THE ROLE OF SALTS IN THE PRESERVATION OF LIFE .

.

1911. Biologists have long been aware of the fact that the ocean has an incomparably richer fauna than fresh-water lakes or streams and it is often assumed that life on our planet originated in the ocean. This proportion identical with the proportion in which these salts are contained in sea-water. fats. It is salts is not the same in both about three times as high in the sea-water as in our blood serum. Ca. but the concentration of the three cases. 5 molecules of CaCl. The fact that the salts of Na.2 molecules KCl. 653-65. namely.S. as a kind of inheritance. namely. October 19. 100 molecules NaCl.IX THE ROLE OF SALTS IN THE PRESERVATION OF I LIFE^ Less is known of the role of the salts in the than of the cerned. N. we diluted sea-water in our blood. and that. and proteins. 1 still carry Statements of this kind have Carpenter lecture delivered at the Academy of Medicine of New York. by courtesy of Professor James McKeen Cattell. although no one can say why We have a point of attack for the investigation of the role of the salts in the fact that the cells of our body live longest in a liquid which contains the three salts. and CaCU is in a definite proportion. role of the three other carbohydrates. animal body main food-stuffs. fact that The Yet be it seems to be a indefinitely.^. neutral salts. November 17. 1911. Reprinted from Science.. 881. no animal can live on an ash-free diet this should so. are not oxidizable. 169 . and 1 . KCl. NaCl. As far as the latter are con- we know at least that through oxidation they are capable of furnishing heat and other forms of energy. XXXIV. 2. however. No. and K exist in about the same proportion in our blood serum as in the ocean has led some authors to the conclusion that our ancestors were marine animals.

but they serve to emphasize the two facts. animals are put into cane-sugar solution. the osmotic pressure of which is equal to that of sea-water. If these animals are suddenly thrown into (at a distilled water. the animals die just in distilled water. that the three salts. intend to put before you a series of experiments which light seem to throw some on the mechanism by which the life. KCl.. NaCl. diluted with an equal quantity of distilled the sea-water is water in one case. allowed to elapse before bringing no recuperation is possible. solutions surrounding living cells influence their duration of II In order to give a picture of the extent to which the life of many animals depends upon the cooperation of the three salts I may mention experiments made on a small marine crustacean. they can recuperate. of the Bay of San Francisco.170 The Mechanistic Conception of Life mainly a metaphorical value. higher or lower than therefore. about as rapidly as The same is is. one would expect that the deadly if effect of distilled water would be inhibited distilled enough cane sugar were however. exist in our blood in the same relative proportion as in the ocean and that they seem to play an important I role in the maintenance of life. Since in this case death is caused obviously through the entrance of distilled water into the tissues of the animals. their respiration stops If temperature of 20° C.water.) in about half an hour. is true if the osmotic pressure of the sugar solution that of the sea-water. the solution equal to that of the sea-water. the added to the water to make the osmotic pressure of If. they are put back immediately after the cessation of respiration into If ten minutes or more are sea. them back into the sea-water. The sugar solution about as toxic for these animals as the distilled water. Gammarus. and of isotonic cane-sugar solution in the . although in the latter case water enters into the tissues of the animal. while it in the former case If does not. and CaCl.

. of the same osmotic pressure about half an hour.. for their nutrition. Ill What It is is the role of the salts in these cases ? The botanists have always considered nitrates salt solutions as nutritive solutions. All this leads us to the con- clusion. The little. salts.Role of Salts in Preservation of Life life is 171 other. both CaClo and alone to the If KCl are added to the sodium chloride solution. salts of We find also that the proportion in which the three sodium. calcium. the animals die as quickly or more so than in the pure cane-sugar solution. addition of KCl NaCl KCl and CaCl. sodium chloride. however. in both cases the duration of cally the If shortened by practi- same amount. calcium and potassium chloride in the proportion in which . for The relative tolerance of various cells is. other salts be substituted for the three salts the animals The only substitution possible is that of SrCl. definite concentra- tion or osmotic pressure but that this osmotic pressure must be by definite salts. for CaCl.. the solution must not only have a furnished chloride. these three salts exist in the sea-water (and in the blood) fact could also be this demonstrated for many other marine animals. e.. a well-kno^\^l fact that plants require definite and potassium arises salts. . If. and animals abnormal salt solutions however. that in order to preserve the life of the crustacean Gammarus. and potassium have to exist in the solution cannot be altered to any extent. namely. a point which we shall discuss later on.g. not the same. If to this solution a little calcium chloride be added in the proportion in which it is contained in the seawater the animals die as rapidly as without it. and CaCl. tion now whether the three salts and the quesXaCl. they also die in the crustaceans are brought into a pure solution of NaCl. prolongs their are life but added to a cane-sugar solution isotonic with sea-water. If die. KCl. the animals can live for several days. as the sea-water.

. proportion. very dilute and does not exceed the concentration of m/8. who found that there is a certain antagonism between K and Ca in their action on the heart. in distilled and fresh water. be added the to the solution in the right indefinitely. and certainly in a very dilute solution of sodium chloride. Gammarus and the tissues but they differ from Gammarus and the fish majority of marine animals inasmuch as the can live long. The discovery of antagonistic salt action was made by Ringer. but as a -protective solution. prove that a mixture NaCl + KCl +CaCl2 does not act as a nutritive solution. newly hatched fish are is not the case. KCl and if its CaCl. action . Fundulus. marine fish. with a case of antagonistic salt an antagonism between NaCl on the one hand and KCl and CaClo on the other. succeeded in showing that as long as the sodium chloride solution required. behave in this respect like of the higher animals. that they only serve to render NaCl harmless I is if the concentration of the latter salt is too high. therefore. solution of If put into a pure sodium chloride of the concentration in which die. therefore. From this fact I drew the conclusion that KCl and CaClg do not act as nutritive substances for these animals.^ are only necessary in order to prevent the harmful effects which it is NaCl produces is if alone in solution and concentration too high.172 The Mechanistic Conception of Life which are needed for the preservation of animal life. animals can live These fish. proved beyond question that this the young. this If. indefinitely. is not NaCl has a concentration KCl and CaCl. We are dealing. the animals very soon however. KCl and CaCl. play the Experiments which I made on a small role of nutritive salts. in other words. salt is contained in sea-water. of The experiments on Fitndi^ZiiS. the addition of above m/8 Only when the solution of does it become harmful and require the addition of KCl and CaCl. and in some cases. When he put the heart of a frog into a mixture of NaCl + KCl he found that the contractions of the heart were not normal.

and other bivalent metals. they usuall}' die without forming an embryo. eggs are put immediately after solution which is fertilization into a pure sodium chloride isotonic with If. Cu. Ni. the toxicity of the diminished or even salts abolished. at can live long enough to form to render the pure solution of least to the extent that the eggs an embryo. It had generally been assumed that the antagonistic action of two salts was based on the fact that each salt. calcium salt. Since these experiments were antagonistic action of salts have made many examples of the become kno^^Ti. namely. IV What is the mechanism of antagonistic salt action? I believe that an answer to this question lies in the following If these observations on the eggs of Fundidus. JVIn.. however. of Ba. only a trace of a or of the exception of any other salt with a bivalent metal (with Hg. similar to that which Schmiedeberg had found between e. We shall see that in certain cases of antagonistic salt action at least this view is not supported bj' fact. when applied singly. line salt Biedermann had found that alkasolutions cause twitchings in the muscle and Ringer found that the addition of Ca inhibited these twitchings. A of mixture of NaCl+CaCl. Ringer drew the conclusion that there existed an different heart poisons. The fact that a substance as poisonous as Zn or lead can render harmless a substance as indifferent as sodium chloride seems so paradoxical that it demanded an explanation.g. . are able sodium chloride harmless. acted in the opposite way from that of its antagonist. also caused abnormal contractions of the heart. Pb. or Ag) is added to the m/2 NaCl solution is solution. Zn. Ko. sea-water. but these were rendered normal by the addition KCl. atropin and muscarin. Even salts which are very poisonous. antagonism between potassium and calcium.Role of Salts in Preservation of Life b}' 173 but they were rendered normal the addition of a Uttle CaClo.

has found that in the larvae of Arenicola a slight antagonism between NaCl and ZnS04 can be proved. For the antagonistic action of a salt of lead or zinc against the toxic action of sodium chloride can only consist in the lead action possible? salt protecting the embryo against this the toxic action of the NaCl. R. into a pure m/2 solution of chloride the animals die very quickly. while of CaCl. but that they live indefinitely in the CaCl. the so-called micropyle. it is How happen that is for the long as in the egg shell. more rapidly by an m/2 solution of by the same solution one or two days One can imagine that the micropyle conwhich is wad Lillie of a colloidal substance if hardened gradually to a leathery consistency 1 the egg remains in the sea-water. This egg membrane possesses a small opening. is separated from the surrounding solution by the egg membrane. alone is the fish out of the shell the addition no longer it is sufficient and the addition if of KCl also life becomes necessary ? of the fish after Moreover. as long as in the egg. sodium chloride solution does it if we add both embryo. immesodium diately after hatching. .. as and KCl. I have gained the impression that killed this micropyle is not closed as tightly immediately after fertilization as later on. This shows that the general laws of antagonism between two salts differ in degree but not in principle in the living organism and the dead envelop of the fish egg.. But how is protective We have mentioned that if we put the young fish. since the newly fertilized egg is NaCl than tains a it is killed after fertihzation. to the solu- tion we find that the fish die even much more w^ithout the addition. we try to preserve the taken out of the egg in an m/2 sodium quickly than chloride solution by adding ZnSO^. if the addition of CaCl.^ If is we look for the cause of this difference our attention it is called to the fact that the fish.174 The Mechanistic Conception of Life this explanation casts light and on the nature of the protective or antagonistic action of salts. through which the spermatozoon enters into the egg. to the NaCl solution suffices. or lead acetate.

beginning If we add varjdng amounts a certain of NaCl we NaCl. a large percentage of the eggs develop^ often as many as 100 per cent. of distilled we add^ quantity of ZnSO^ If all required to render the NaCl solution harmless. therefore. Ko. and Xi.) If tion into distilled w^ater. find that. Pb. Further data are furnished by experiments which I made together with Professor Gies. however. It is also possible that similar changes take place in the whole membrane. Zn.: Role of Salts With the process impermeable is in Preservation of Life it 175 of hardening. This process of hardening if brought about apparently very rapidly we add m/2 NaCl solution a trace of a salt of a bivalent metal like Ca. also on the eggs of Fundulus. but that the reverse is also the case. Mn. table This can be illustrated by the foUo^Wng . Gies and I were able to show that not only are the bivalent metals able to render the sodium chloride solution harmless. Sr. to 100 is c. is the eggs are killed rapidly and not a single one able to form an embryo. for instance ZnSO^. many of the bivalent metals. becomes more to the for the NaCl solution. Ba.c. that NaCl is required to render the solu- tions of less. or tanning. rendering the fish. and the larvae and embryos water that formed which in the distilled water are able to hatch. etc. m/2 Na solution harmless for The process of the embryo of the depends apparently upon the fact that the addiof the egg tion of the bivalent metals renders the micropyle or perhaps- the whole membrane more impermeable to NaCl than was the case before. concentration of inhibits the toxic effects of ZnSO^ and many eggs are able to form an embryo. But these are only one part of the facts which throw a light upon the protective or antagonistic action of salts. namely. this salt with. harm- (That the SO^ ion has nothing to do with the result wasthe eggs of Fundulus are put immediately after fertiliza- shown before by experiments with Na^SO^.

m/32 ZnSOi 3 8 44 38 37 34 29 '. cooperative action of both salts this on the egg membrane. c. c. in this case at least.c. c. is concentra- able to render the ZnSO^ If in the solution comparatively harmless. c. m/32 ZnS04 m/32 ZnSOi m/32 ZnSO. c.c.c. c.c.c. that this antagonism depends. As long salts acted.c. is this we must its draw the further conclusion that the salts.c.c.c.c.c. as if we assumed that each of the two antagonistic applied singly. c. c. m/8. distilled c. And from toxic. m/32 ZnSO^ m/32 ZnSO^ m/32 ZnSO. water+8 100 100 100 100 c. so that comes into direct contact with the protoplasm of the germ.c. c. c. c.176 The Mechanistic Conception of Life TABLE Nature of the Solution I Percentage of the Eggs Forming an Embryo 100 100 c. c.c. We therefore arrive at a new conception of the mutual antagonism of two salts. it was impossible to understand these experiments . is due to comit paratively rapid diffusion through the membrane. NaCl+S NaCl+8 NaCl+8 NaCl+8 m/4 NaCl+8 3/8 NaCl+8 m/2 NaCl+8 5/8 NaCl+8 6/8 NaCl+8 7/8 NaCl+8 m NaCl+8 m/64 m/32 m/16 m/8 c. in the opposite way from its antagonist. namely.c. c.c. 100 100 100 100 c. that NaCl renders the egg membrane comparatively impermeable for ZnSO^.c.c.c. m/32 ZnS04 m/32 ZnSO.c. c. m/32 ZnS04 m/32 ZnSOi m/32 ZnSO. if it is fact that each of these alone in the solution.c. c.c.c. namely. c. 8 6 1 if its This table shows that the addition of NaCl. or by which action membrane becomes completely com- paratively impermeable for both salts. distilled water 49 c. c. we now assume that ZnSO^ renders the 5/8 m NaCl solution harmless by rendering the egg membrane comparatively impermeable for NaCl we must also draw the opposite conclusion. 100 100 100 c. c.c. upon a common.c.c. tion exceeds a certain limit. namely.

We may. But if we NaCl alone is toxic because it is not able to render egg membrane impermeable. the example. NaCl or some other salt of sufficiently high osmotic decreases the volume of the contents of the loss of cell through water and the protoplasm retracts. since they leave of Fundulus are theoretically in this case at no doubt that least the "antagonistic" action of salts consists in a modification of the egg the membrane by a combined action of two salts. solution toxic for the These experiments on the egg of importance. not only able to render NaCl harmless. but some authors. and vice versa. are put into a solution of pressure.Role of Salts or find an analogue for realize that in Preservation of Life 177 the them in colloid chemistry. are at all permeable for Nobody denies that salts diffuse much especially more slowly into the cells than water. and that ZnSO^ if alone in is same reason. CaCl2. urea. V It is not easy to find examples of experiments in the litera- ture which are equally unequivocal in regard to the character of antagonistic salt action . is from corners of the Overton maintains that this plasmolysis permanent. whereby membrane becomes less permeable for both salts. still mention that salts alone have such antagonistic effects. while both combined are harmless (since for the 'banning" of the membrane the action of the two salts is required) these experiments become clear. and On the other hand. cells has long been a question whether or not salts. especially rigid cellulose walls. but also LiCl. ZnSO^ was alcohol have no such action. those of Spirogyra. and others. Overton's view is based partly If on experiments on plasmolysis cells of plants. for in the cells of plants. deny categorically that they can diffuse at all into the cells. and concludes from this . NH^Cl. for the sake of completeness. glycerin. but I think that some recent experi- ments by Osterhout It satisfy this demand. Overton and Hoeber.

This stage of If reversion of plasmolysis had been overlooked by Overton. which superficially resembles plasmolysis. A .2 NaCl solution has approximately CaClo.2m CaClo solution.. of the cell the restitute the this does water should back into the and normal volume of the According to Overton not happen. but which in reality has nothing to do with plasmolysis. Osterhout explains this difference in the concentration of the two I salts required for plasmolysis by the assumption that NaCl CaCl. In one experiment which Osterhout describes. since otherwise salt should gradually diffuse from the solution osmotic pressure cell into the cell. a portion of a Spirogyra filament was plasmolyzed in . m m m the same osmotic pressure as a . it." as Osterhout calls has nothing to do with the hyper- tonic character of the solution was proved by the fact that hypotonic solutions of toxic substances may produce the same phenomenon. CaCl2. a shrinking of the contents of the sodium chloride cell takes place again. Osterhout's experiments also show that the antagonism of NaCl and CaClg depends partly on the facts that the two salts . remains permanently in the h^^jertonic solution. But on placing 29 another portion of the same Spirogyra filament in a tion the expected plasmolysis does not occur m NaCl solu- plasmolyze the cells until and it is impossible to they are placed in 4 m NaCl. however. the cell. a conclusion diffuses more rapidly into the cell than which reached also on the basis of my earlier experiments on animals. Osterhout has recently shown that Overton's observations were incomplete in a very essential point and that in reality the plasmolysis. which occurs in this case when the cell is put into the h^q^ertonic solution. . disappears again in a time which varies wdth the nature of the salt in solution. and through this increase in the finally diffuse cell.29 but not in 195 .. but is a phenomenon of death.178 The Mechanistic Conception of Life that only water but no salt can diffuse through the cell-wall. . That this second ''false plas- molysis.

n/10 butyTic acid to 100 c. After a longer the mixture of period of time the plasmolyzed cells can expand again in a mixture of NaCl and CaCl. If we add 0.4 c. By it that . or less acid they can live for a of acids week or more. . perfectly balanced solutions. 179 from diffusing into the and this conclu- sion is based among others upon the following experiment. The explanation for this observation lies in the fact that in NaCl and CaCl. the two salts render their diffusion into the cell mutually more difficult. of butyric acid to 100 c. In solutions which contain 0. 195 m CaClo and placing other portions of the same filament in it.5 c.38 m XaCl. the diffusion was inhibited or considerably retarded.c. On mixing 100 c. of solutions of NaCl of various concentration. .2 m CaCL and in . yet they do not prove that these salts diffuse into the ditions.Role of Salts inhibit each other in Preservation of Life cells.c. we find that above a certain limit . These experiments are the analogue of the observation on the embryo of the eggs of Fundulus in which a pure solution of ZnSO^ diffused rapidly through the membrane or micropyle. but that occurs much later than if they are in the pure NaCl solution.c. if both salts were present. of distilled water these fish die in 2| hours or less. While the observations of Osterhout show that Overton was not justified in using the experiments on plasmolj^sis to prove that the neutral salts cannot diffuse into the cells. cell under normal conare in strongly In Osterhout 's experiments the cells h^T^ertonic solutions and it does not follow that such solutions act like isotonic. prompt and very marked plasmolysis occurred. VI Wasteneys and I have recently sho^Mi that the toxic action upon Fundulus can be annihilated by salts.375 m NaCl.195 m CaCla nor in .375 m XaCl with 10 c.c. dividing a Spirogyra filament into several portions it was found was plasmolyzed in .c. while. If we add. but neither in . 5 c.c.c. however..

c. It is needless to say that the NaCl used in these experiments was strictly neutral and that the amount of acid present in the mixture of acid and salt was measured. m/2 m/2 m/2 m/2 m/2 m/2 +0.c.c.c. c.0 8.0 + 4. of the Solution 2 hours 4 hours 1 day . c.c.2O+ 4 H.c.c.c.c. c.c.5 NaCl+0.c. c. HoO H. 180 the The Mechanistic Conception of Life NaCl can render the acid harmless. c.5 NaCl+0. c. Six fish were put into 500 c. 2) 3) c.c.c.c.c. c. c.c.c.0 12.5 NaCl-f-0.c. 6. The following experiment may serve as an example.c.c. c.c.5 n/10 butyric acid n/10 butyric acid n/10 butyric acid After certain intervals the ascertained. c. 2 days. 4 days.5 NaCl+0.2O+ 6 c. number of surviving fish II.5 NaCl+0.0 in 10. . HoO+10 HoO+12 H2O+I5 c. 94 92 90 88 85 4) 5) 6) 7) H0O+ 8 c.0 m/2 NaCl 100 c. c. of each of the following seven mixtures. NaCH-0.5 c. was The result is given in Table TABLE Number After II of Surviving Fish in 0. n/10 Butyric Acid 15.c. 1) 100 96 c. c. c.c.c.0 c. namely. 3 days.. n/10 n/10 n/10 n/10 butyric acid butyric acid butyric acid butyric acid c.5 c.

however. probably not the direct cause of the death.. It is well ^ It is possible that in the killed in the following experiments with acid the fish is way.^ Role of Salts Wasteneys and of acid I in Preservation of Life 181 could show that the rate of the absorption salt. the combined action of acid and salt causes a dehydration of the surface film 1 salts It seems that the first. whereby this surface layer becomes able for the acid. the fish into a weak acid solution in distilled water just strong enough to (e.0 c. can state a If little more kill definitely the cause of death by we put c. little later is the epidermis falls off in pieces and This. we notice that the acid very soon makes the normally transparent epidermis of the fish opaque. whereby the If salts are present in the right concentration. cells and of the more permeare killed.c. p.O+2. 308. the fish in from one to two hours n/10 HCl). Thus the internal friction of certain protein solutions is increased by acids while the addition of neutral salts inhibits this effect (Pauli). LXXV. known that the action of acids upon proteins can be inhibited by neutral salts. by the fish is the same in solutions with and without This proves that the action of the salts consisted in this case not in preventing the diffusion or absorption of the acid. mentioned . The well as action of the acid upon the epidermis of the body as upon the gills is prevented through the addition of neutral salts.c.g. 1899. but in modifying the deleterious effect of the absorbed acid. experiments on the antagonism between acids and were published by the author in PflUgers Archiv. and a shreds. Vol. 500 H. 2 The beautiful osmometric experiments of R. The swelling of gelatin caused by acid is inhibited by salts (Procter). The acid can now diffuse into the epithelial cells and act on the protoplasm. The acid causes certain proteins in the surface layer of the epithelial cells of the gills skin to swell. We acid. but I am inclined to assume that the fish die from suffocation caused by a similar action of the acid upon the gills. Lillie should also be in this connection.

into 500 c. in wide limits. it must be possible to show that not only KCl. independent of the osmotic pressure of the surrounding solution since the concentration of the low. is —the antagonism which NaCl toxic if it is alone in solution. not essentially diminished by the presence of is but while tanning in the presence of salts the acid consumed or is in the action of the surface layer of the layer. but that the reverse true This can be proved in the case of KCl. KCl in sea-water is very The experiments were carried out by Mr. and KCl and CaCl. on the one hand. after washing them twice with distilled water. is of the same nature as that between NaCl and ZnSO^ in the case of the eggs of Fundulus.182 The Mechanistic Conception of Life it of these cells. is We choose as an example the antagonism between NaCl. absorbed in this no salt is present part of the acid diffuses into the epithelial cells and kills the latter. on the other. and that salts. . is electrolytes a little The case of the antagonism between three more complicated. it. life In this saved. The method consisted in putting six fish. To demonstrate we have again to experiment on organisms which are. if cells. and CaCl. it is rendered is harmless by the two other also. most important in life phenomena. Wasteneys and myself on Fundulus. as does in the experiments on gelatin or as in the cases of tanning of hides by the combined action of acids and gills salt solutions.c. If the mechanism of the antagonism between NaCl. VII We have thus far considered the cases of antagonism between two electrolytes only. way the gills are kept intact and thus destroying and the of the fish is As long as the amount is of acid is small the amount absorbed salts . This combined dehydrating or "tanning" action of acid and salts on the surface of the epithelial cells of the diminishes the permeability of this layer for the acids and cells prevents them from diffusing into the the protoplasm..

which only 1/50 of that of the sea-water. the fish were put into pure solutions of is KCl of the concentration in which this salt contained in the sea-water m/2 KCl in 100 c. This is is not due to the low concentration of the KCl solution. if not completely The following table may serve as an example TABLE After Days III . KCl is considerably diminished.: Role of Salts of the solution.c. we add to the toxic quantities of KCl increasing quantities we find that as soon as the solution contains 17 or more molecules of NaCl to one molecule of KCl. If as that in which the NaCl solution of the KCl exists in the sea- of NaCl.2 c.c. in Preservation of Life 183 It was ascertained from day to day how many When (2. since the fish can live indefinitely in a pure same concentration water. of the solution) they died mostly in less than two days. fish survived. the toxic action of comiteracted.

m NaCl. but it becomes more incomplete the this reason it is higher the concentration of KCl. but it more powerfully. to if ratio ? If we add too little only 1 to 10 molecules NaCl 17 1 molecule of KCl.c. 7/32 m NaCl. KCl rendered KCl rendered KCl rendered KCl rendered harmless in 100 harmless in 100 harmless in 100 harmless in 100 c..c. NaCl .c. the This indicates that the antagonistic salts. but that If in addition is needed. of the solution exceeds 2.c.c.c. 1/17 1/19 1/17 1/16 5/32 . 100 c. .0 c.c. For of antagonization is still to some extent possible. m/2 1. m/2 2. effect requires This furnishes the reason why an m/2 CaClg solution of NaCl can. .75 c.c.c. 1/17 What happens if we vary this NaCl to the KCl solution. . namely.c. 6/32 m NaCl 100 c.c. the solution becomes more harmful than is KCl alone in solution. m/2 0..c.c.g.9 c. If the amount of KCl in 100 c. 1/18 4/64 NaCl. . . as that of several hundred or a thousand times as great as the following tables shows. m/2 KCl. we add to 100 c. 1/17 5/64 5/64-6/64 1/16-1/19 . .c. . c. m/2 1 1 . . KCl/NaCl.c. KCl/CaCla.c.65 c. solution 50 molecules to one molecule of KCl. m/2 2 2 c. possible to render not an m/2 solution NaCl harmless by the as mere addition acts of KCl. as a rule.. not be rendered completely harmless by the mere addition of KCl. in in in m NaCl. m/2 0.e. m/2 0.c. m/2 NaCl c.7 c. .c. e. c. if we add considerably more than molecules NaCl. . m/2 NaCl enough KCl to make the ratio KCl: NaCl = 1/17 we find that the antagonization of KCl: NaCl becomes incomplete.1 184 The Mechanistic Conception of Life TABLE IV Coefficient of Antagonization. m m m . 1/16 3/64 NaCl. m/2 3 3 c. .c. is the coefficient of antagonization. becomes toxic again. m KCl rendered harmless KCl rendered harmless KCl rendered harmless KCl rendered harmless KCl rendered harmless in 100 c. c. CaClg acts upon KCl i.2 c. .6 c. . 9/32 m NaCl 6/64 . . and the more so the higher the a rather definite ratio of the two concentration of NaCl.c. . . in 100 c.c.. similarly does NaCl. 100 c.

c. antagonizes KCl also powerfully. c. of the solution.5 .550 CaCL.c.3 m/100 1. .6 m/100 CaCl^. CaCL. that CaCIg produces its protective effect through the formation of a comparatively insoluble compound (in this case on the gills or the rest of the surface of the animal) while NaCl acts through the formation of a compound which is more soluble.75 3. but. Why is less CaCl.65 c.. H^O 100 c. 165 . for instance. . very remarkable that CaCl. m/2 KCl in 100 c.c. is due to the fact that the minimal value of CaCl^ at which renders the KCl harmless cannot be determined as sharply as the limit for NaCl.3 m/2 m/2 m/2 m/2 m/2 KCl KCl KCl KCl KCl in 100 c. CaCU. .c.6 c. but not this limit. This view is corroborated by the observation which we made. does not permit us to render harmless more than 6.137.c. in the case of the sulphates.366 . renders harmless any given concentration of KCl below 6. probably.c.c.. if salts. required than NaCl ? We can only answer with a sug- by T. but that Mg much less efficient. . .0 100 c.c.2 2. than the calcium salts are often while the magnesium incomparably more soluble. .O 100 c. in in in H. .6 above c. in consequence of the fact that the substances formed at the surface of the animal or the gills. m/2 KCl in 100 c. 2. H. H^O This it require 0. that Sr is gestion first offered just as effective to antagonize is KCl as CaCl.c. c.1 require 0. B. 1 . not more insoluble. c. Robertson.O H. 103 The coefficients are not as regular as in the case of antagoni- zation of KCl by NaCl.0 m/100 1. diffuse slowly into the cells. .c. of the is solution. in 100 c. . .. This limit exactly the same which we found in the case of antagonization of KCl by NaCl.c. c.. Even the combination of NaCl and CaCl.5 require require require m/lOO m/100 0.c.c. This would correspond with the wellknown fact that many strontium salts are just as insoluble. BaCl. CaCL.c.1 Role of Salts in Preservation of Life 185 TABLE V Coefficient of Antagonization KCl/CaCL 1 . the fish do not remain alive as long if Ba is used as if the more harmless It is Ca and Sr are used. namely.

KCl and NaCl. a film which surrounds the embryo and keeps the such eggs are brought back into solution containing calcium is the film restored and the cells come into close contact again. can antagonize even a 6/8 tion of NaCl. It is therefore not impossible that the mechanism of the is antagonism between KCl and NaCl similar to that found between NaCl and ZnSO^. 1.5 c. Such an opportunity is offered us through the . who found that in sea-water deprived of calcium the segmentation cells of a sea-urchin embryo If fall apart through the disintegration or liquefaction of cells together. KCl Experiments with pure CaClg solutions give the result that is this substance harmless in a solution of that concentration in is which this salt contained in the sea-water. but that we must be able to see directly in which way abnormal salt solutions cause the death action of salts that of the cell. not so unequivocal as the previously mentioned cases of antagonism between only two VIII It is necessary for our understanding of the life-preserving we do not depend merely on conclusions drawn from experiments. we ceases to find that CaCl. live indefinitely in a solution of m/2 CaClg Botanists have also found that weak solutions of CaCl^ are comparatively little toxic. centration of the in addition to is It seems only due to the high con- NaCl in the sea-water and in the blood that. CaClj is needed. while if m and a 7/8 m solushow any antagonistic effect the NaCl solution exceeds m/2 or 5/8 m. But the case electrolytes. This gives us the impression that the effect upon the surface film of protoplasm produced by CaCla plasm.c. is especially important for the protection of the proto- This conclusion receives an indirect support by the well-known experiments of Herbst.c.186 If The Mechanistic Conception of Life we try to render NaCl harmless by KCl and CaCL. Fundulus can in 100 c.

however. If is we put the egg into an abnormal solution this surface film is modified and changed. around the protoplasm of each is kept together.Role of Salts in Preservation of Life If 187 ferti- observation of the eggs of the sea-urchin. so is that if one become familiar with these pictures. off of small granules or This process gradually continues from the periphery until toward the center different. begins on the surface of the protoplasm. suggest the possibility that the combination of the three salts in their definite proportion and concentration has the function structure or cell. The previously mentioned observation shows the important role of Herbst again calcium in this process. the whole egg is disintegrated. is For little different salt solutions the picture of the disintegration a but sufficiently characteristic for a given solution. . sodium and potassium chloride. if This process of disintegration can be observed are put into a pure solution of the eggs sodium chloride. as a rule. therefore. a gradually takes place. one able to diagnose to some extent the nature of the solution from the in way which the cell disintegrates. of the egg remains if One gains the impression as the protoplasm of the egg were held together by a continuous surface film of a definite texture. and consists very often in the formation and falling droplets. or in a mixture of sodium chloride and calcium chloride chloride. These observations on the sea-urchin egg. by which the protoplasm protected against and separated from the of surrounding media. and the change of the surface film often followed by a gradual process of disintegration of the rest of the cell. all three salts are used in the proportion in which they occur in the sea-water no disintegration takes place and the surface perfectly smooth and normal. we put The the Hzed eggs of the sea-urchin into an abnormal destruction of the cell salt solution. destruction. or in a mixture of If. of forming a surface film of a definite texture.

c. was found that these animals in a solu- a mixture of tion of NaCl. The solutions were all 3/32 m. Ostwald worked with fresh-water crustaceans which he put into mixtures of various live longer in salts.c. mm. NaCl+2. In NaCl the Spirogyra died in 18 hours. 3/32 3/32 3/32 CaClg 2 to 4 weeks.+2. Osterhout was able to show that the spores of a certain variety of Vaucheria die in a pure 3/32 m solution of NaCl in 10 to 20 minutes. The reac3/32 tion of the solution was strictly neutral and the NaCl the purest The results remained the same after the NaCl obtainable.0 3/25 CaCL NaCl+2. for which ^'adaptation" and CaCl. 740 59 3/25 NaCl 3/25 3/25 c. and CaCl. Nature of the Solution Total Length of Roots after 40 Days H2O 100 100 100 c.c. KCl. in NaCl+KCl+CaCl. prove that the life-preserving combinanot due NaCl + KCl +CaCL in definite proportions is . which are said to be '^ adapted" to a mixture of NaCl. Experiments on fresh-water organisms. or NaCl+CaCl. in NaCl+KCl in two m days. These tion of cases.2 c. CaCl. and in 100 c. of the same osmotic pressure. to a mixture of NaCl. KCl.. Osterhout caused wheat grains to develop in such solutions and measured the total length of the roots formed. 3/32 m KCl 6 to 8 weeks.188 The Mechanistic Conception of Life IX The objection might be raised that the beneficial action of the three salts could only be proved on marine animals or on tissues of higher animals. in these definite propor- show that It this objection is not valid.c. in definite proportions. tions cannot be claimed.c. m NaCl + 1 c. m NaCl + 1 c.0 3/25 CaCl2+2. had been recrystallized six times. they lived 65 days. mm.c.c. c. Experiments with Spirogyra gave a similar result. while they live in 100 c. to many other similar observations might effect of the be added. mm.2 3/25 which m KCl 254 324 mm.c. or NaCl + KCl +CaClo than NaCl+KCl.

While a marine of Gammarus dies in half an hour in an isotonic solution (of NaCl or cane sugar. The One is surrounded by a cortical the formation and this layer destroyed or modified in the process of is fertilization. even a pure solution of NaCl. But the reader resist- has noticed that there are considerable differences in the ance of various organisms to abnormal solutions . firmed and amplified by Harvey. make it doubtful whether the egg is salts which are not soluble I believe in fats can enter the fertilized is egg at all.Role of Salts in Preservation of Life 189 to the fact that organisms are ''adapted" to this mixture but to a specific protective effect of the combination of the three salts upon the cells. than in any other osmotic solution. that the explanation of the difference unfertilized egg is much more layer simple. result of this modification of the fertilization membrane. or of NaCl for several days at a temperature of 15°. therefore. while the fertilized eggs of the same female are killed in a pure neutral solution of CaClo in a few hours. which were conegg. for which I have been able to . red blood corpuscles or even the muscle of a frog can be kept for a day or longer in such a solution the muscle of a frog lives longer addition if course even the NaCl solution contains in KCl or CaClg). What causes this difference? Six years ago I found that the unfertilized eggs of the sea- urchin (Strongylocentrotus purpuratus) can keep alive and remain apparently intact in a pure neutral solution of CaCl. The same is difference is found What causes this difference? it Several authors have suggested that fertilized due to the fact that the more permeable to salts than the unfertilized But recent experiments by Warburg. in the proportion in which these salts exist in the blood and in the ocean. for other salts also. X It seems. to be a general fact that wherever tissues of a comparatively high osmotic or animals require a medium pressure — like our tissues — their life lasts much longer in a mixture of NaCl+KCl+CaCl.

the layer destroyed by fertilization the surrounding directly in contact with the protoplasm it salt solution if comes is and the solution abnormal can cause the disintegration of the surface layer of the protoplasm.190 The Mechanistic Conception of Life it is show that readily permeable for salts. against the But tions aside from differences of this kind there are other condiof resistance of cells to various fertilized which influence the degree I solutions. however. Microorganisms which solutions of a high can live in strong organic acids or salt concentration probably possess a surface layer which shuts off their protoplasm from contact with the solution. to believe that during or previously to am inclined cell-division. effective wall For the protoplasm of muscle the rather tough sarcolemma forms not an absolute but nevertheless an surrounding solution. have dra^\Tl the conclusion that in a pure NaCl solution the rate of oxidations of the egg of Strongylocentfotus is increased kills and that the eggs.. lack of oxygen prolongs the fertilized egg just as well in solutions of NaCl-f-CaClg or of for NaCl+BaCl. prevents the surrounding salt solution from coming in contact with the proto- plasm or at cortical least is it retards this process. I am inclined to believe that differences in the resisting power of various cells or organisms to abnormal salt solutions are primarily due to differences in the constitution of the protective envelopes of the animals or the cells. which salts these authors do not claim that I they can raise the rate of oxidations of the egg. is As long it as the cortical layer of the unfertilized egg intact. besides . If. either the oxida- by the withdrawal of oxygen Warburg and Meyerhof or the addition of KCN or NaCN. moreover. it is this increase in the rate of oxidations this increase of oxidations which But cannot be observed of the in the eggs of Arbacia when they are put into a pure NaCl solulife tion and. have foimd that the eggs of the seaif urchin will live longer in abnormal salt solutions tions in the egg are stopped.

This assumption is not compatible with two cell. whereby these bility films acquire those physical qualities of duracell and comparative impermeability. The experiments mentioned effect in this paper indicate that the role of salts in the preservation of life consists in the ^'tanning" which they have upon the surface films of the cells. The above-mentioned facts of the antagonism between acids and salts suggest the idea that the surface film of cells consists exclusively or essentially of certain proteins. that water diffuses very rapidly into the and second. namely. cannot On this assumption we can understand that neutral life salts should be necessary for the preservation of although they do not furnish energy.Role of Salts phenomena film of the in Preservation of Life 191 of streaming inside the cell. in which the work can if only go on in the proper way. As far as the dynamical effects of salts are concerned it is . the nature of the surface layer of the Overton and others assume that facts. changes in the surface is protoplasm occur. without which the exist. that life depends upon an exchange of watersoluble and not of fat-soluble substances between the cells and the surrounding liquid. XI If we now raise the question as to why cell salts are necessary for the preservation of the life of the we can clear. point to a number of cases in which this answer seems Each cell may be considered a chemical factory. the diffusion of substances through the cell-wall is restricted. more easily injured by the If we suppress the oxidations we suppress also the processes leading to cell-division and thereby retard the deleterious action of the abnormal salt solution upon the surface layer of the protoplasm of the egg. This diffusion depends on cell. whereby this film salts. this layer consists of a continuous membrane of fat or lipoids.

The fact that the addition of of the calcium to an NaCl solution prevents the twitchings muscle.192 The Mechanistic Conception of Life not impossible that some of them belong also to the type of those mentioned in this paper. the apparent dependence of of contractility of the muscle upon the presence NaCl.. merely prevents or retards the the sarcolemma. .g. do not find their explanation in the facts discussed here. suggests the diffu- CaCl. sion of salts. NaCl through But other role of effects of e. or the PO^. which occur in the pure possibility that the NaCl solution.

EXPERIMENTAL STUDY OF THE INFLUENCE OF ENVIRONMENT ON ANIMALS .X.

.

INTRODUCTORY REMARKS calls What animal is the biologist the natural environment of an rigid from a physical point of view a rather It is combina- tion of definite forces. 195 . chemical and physical forces upon living Owing to limitation of space this sketch it is necessarily very incomplete. of the University of Cambridge. C. The manner which a germ cell can force upon the adult certain characters will not be understood until we succeed in varying and controlling hereditary characteristics. obvious that by a purposeful and systematic variation of these and by the application of other forces in the laboratory.X EXPERIMENTAL STUDY OF THE INFLUENCE OF ENVIRONMENT ON ANIMALS^ I. England. results must be obtainable which do This is not appear in the natural environment. and must not be inferred that studies which are 1 Reprinted from Darwin and Modern Science (1909). and of bringing about phenomena which cannot be expected in nature. by courtesy of Professor A. Seward. The systematic physico-chemical analysis of the effect of is outside forces upon the form and reactions of animals also our only means of unraveling the mechanism of heredity beyond the results which can be obtained by a mere cytological in investigation. and this can only be accomplished on the basis of a systematic effects of study of the matter. It was perhaps not the least important of Darwin's services to science that the boldness of his conceptions gave to the experimental biologist courage to at will the effects life enter of upon the attempt of controlling animals. the reasoning underlying the modern development of the study of the effect of environment upon animal life.

for the the result of numer- spermatozoa are carried to the eggs by chance and are not attracted by the rule latter. — It was held until recently that hybridization species is not possible except between closely related successful hybridization and that even among these a It is. and gravita- We shall also treat separately the effect of these forces upon form and II. The tion. cannot always be counted upon. recently been showni has by experiment that in fresh sea-water the sperm may live and retain its fertilizing power for several days.196 The Mechanistic Conception of Life not mentioned here were considered to be of minor importance. and beUef. well known that the majority of marine animals lay their unfertilized eggs in the ocean and that the males shed their sperm also into the sea-water. instinctive reactions. the influence of which we shall discuss. which indicate the nature and extent of our control over life phenomena and which Darwin. the effects of chemical agencies a) Heterogeneous hybridization. It is thus unavoidable that at certain periods more than one kind of spermatozoa is suspended in the sea-water and it is a matter of surprise that the most heterogeneous hybridizations do not . are in succession chemical agencies. and when sea-water constitutes a veritable suspension of sperm. The numerical excess of the spermatozoa over the ova in the sea- water is the only guaranty that the eggs are fertilized. All the writer could hope to do was to bring together a few instances of the experimental analysis of the effect of environ- ment. This statement is is ous experiments by various authors. contrary to common As a all or the majority of individuals of a species in this occurs the It a given region spawn on the same day. light. temperature. for instance. forces. also have some relation to the work is of In the selection of these instances preference given to those problems which are not too technical for the general reader. This view was well supported by experience.

star-fish. the is eggs of Strongylocentrotus purpuratus (a sea-urchin which found on the coast of California) can be quantities stars. hydroxide or some other fertilized the eggs of the sea-urchin with the sperm of widely different groups of it animals. It seemed to the writer that the cause of this limitation of hybridization might be of another kind and that in the constitution of the sea-water it by a change might be possible to bring about heterogeneous hybridizations. sperm When this is is.. but not by the sperm of other groups of echinoderms. though in smaller numbers. California. or. The eggs of a sea-urchin can be fertilized by sperm of their o^\^l species. 197 The reason for this becomes obvious when done no egg as a we bring together mature eggs and equally mature and active of different families. by the sperm of other species of sea-urchins. while in normal sea-water or with single egg of the sodium hydroxide not a fertilized same female could forms be with the star-fish sperm which proved effective in the hyperalkaline sea-water. and still less by the sperm distant groups of animals. and about 10~^ n Woods Hole. which in normal sea. This assumption proved correct. n/10 sodium hydroxide to 50 of sea-water. holothurians. In 1903 was sho\\Ti that if we add from about 0.c. rule. or crinoids. the so-called micropyle. c.g. (in 1 Sea-water has a faintly alkaline reaction cal chemist its concentration of hydroxy ions terms of the physiis about 10~^ n at at Pacific Grove.Influence of Environment on Animals constantly occur. e.8 a small but definite quantity of sodium alkali. fertilized. Massachusetts). If we it slightly raise the alkalinity of the sea- water by adding to can be to 0. less fertilized in large by the sperm of various kinds of star-fish. brittle-stars. brittle- and holothurians. and that the micropyle allowed only the spermatozoa of the same or of a closely related species to enter the egg.5 c. The sperm of the various . to be that the spermatozoon The consensus of must enter the egg through a more opinion seemed of narrow opening or canal.water are impossible.c.

between sea-urchin and mollusks. capitata gave the best results. . are the hybrids between sea-urchin and star-fish. Recently. {Antedon Kupelwieser afterward obtained results which seemed to indicate the possibility of fertilizing the eggs of Strongylocentrotus with the sperm of a mollusk (Mytilus) .water). We thus see that possible to by increasing the alkalinity of the sea-water it is effect heterogeneous hybridizations which are at present impos- sible in the natural environment of these animals.8 n/10 sodium hydroxide to 50 c.c. conceivable that in former periods of the that in solutions like sea.water the degree of alkalinof carbon dioxide in the earth's history such heterogeneous hybridizations were possible.— 198 The Mechanistic Conception of Life was not equally effective for these hybridizations. however.c. If it be true. capable of development. Godlewski used the same method for the hybridization of the sea-urchin eggs with the sperm of a crinoid rosacea) . have resulted in an increase of the alkalinity of the sea-water. while the sperm of Pycnopodia and Asterina fertilized only 10 or 2 per cent respectively of the same eggs. or better still. This be obtained in sea-water the alkalinity of c. as Arrhenius assumes. of star-fish the sperm of Asterias ochracea and A. of sea. the writer succeeded in fertilizing the eggs of Strongylocentrotus franciscanus with the sperm of a mollusk result could only Chlorostoma. result of such an increase must have been to render which in the possible heterogeneous hybridizations in the ocean present state of alkalinity are practically excluded. It is. since it was possible to fertilize from 50 per cent to 100 per cent of the sea-urchin eggs. such a diminution must also that the Ice age was caused or preceded by a diminution in the amount and one of carbon dioxide in the air. It is known ity must increase when the amount is atmosphere diminished. But granted that such hybridizations were possible. which had been increased (through the addition of 0. ? would they have influenced the character of the fauna In other words.

But these objections are refuted in which he showed definitely that if the egg of the sea-urchin is fertilized with the sperm of a crinoid the fusion of the egg nucleus and sperm nucleus takes by Godlewski's experiments. By way have purely maternal characteristics and of illustration it may be said that the larvae of the sea-urchin reach on the third day or earlier (according to species and temperature) the so-called pluteus stage. p. h) Artificial parthenogenesis. interest to find out It was. in the case of might be argued that zation the sperm nucleus does not fuse with the egg nucleus. a matter of some whether or not the larvae produced by the egg with the sperm of star-fish fertilization of the sea-urchin or mollusk would form the normal and typical pluteus skeleton.Influence of Environment on Animals and if 199 so. 11). These hybrid larvae heterogeneous hybridi- were exclusively maternal It in character. This was invariably the case in the experiments of Godlewski. corresponding stage. all the experiments on heterogeneous hybridization of the egg of the sea-urchin with the sperm of star-fish. Kupelwieser. and moUusks have led to the same result. that. and the writer. therefore. namely. crinoids. in which they possess a typical skeleton (Fig. of the star-fish nor those of the while neither the larvae molluskform a skeleton at the therefore. So far as the question of heredity is concerned. place in the normal way. Hagedoorn. 10. The number is of experiments and this statement therefore not yet final. the and spermatozoon cannot transmit its hereditary substances to the larvae. brittle-stars. —Possibly life in no other field of biology has our ability to control conditions been proved to phenomena by outside such an extent as in the domain of . that the larvae differ in no way from the pure breed of the form from which the egg is taken. what is their character ? In all cases of heterogeneous hybridization the vitality of the egg or the embryo seems is still weakened and limited it is still doubtful whether any heterogeneous hybrid can reach maturity.

may have This suggests also the possi- bility that at that period outside forces may have supplied the conditions for the development of the egg which at present the spermatozoon has to supply. formic.200 The Mechanistic Conception of Life The reader knows that the eggs of the over- fertilization. and during the last ten years living larvae have been produced by chemical agencies from the unfertilized eggs of sea-urchins. indeed. and a number of annelids and mollusks. different The facts of artificial parthenogenesis. The writer found that brane in the unfertilized membrane of fertilization (Figs. a mental image of the process of evolution with the possibility that parthenogenetic propagation preceded sexual reproduction. what In each form the method of procedure is someand a long series of experiments is often required before the successful method is found. have. the immediate change of the surface of the egg.. holothurians. brief consideration of the For this. as the chemical fertilization or activation of the egg is called. monobasic acids of the fatty series. of animals cannot develop miless a sperma- In this case a living agency arises is the cause of possible to development and the problem whether it is accomplish the same result through the application of well- knowTi physico-chemical agencies. This is. the formation of the so-called 1 and 2). if for no other reason. acetic. a means of artificial parthenogenesis may be of interest to the student of evolution. we can produce this memegg by certain acids. namely. in fact this holds true in regard to the eggs of practically all forms of animals with which such experiments have been tried long enough. It seemed necessary in these experiments to imitate as completely as possible by chemical agencies the effects of the spermatozoon upon the egg. propionic. If we wish to form we have to reckon some bearing on the problem of evolution. whelming majority tozoon enters them. especially the e. effect is a characteristic When a spermatozoon enters the egg of a sea-urchin or certain star-fish or annelids.g. perhaps. true. . star-fish.

efficient 201 in Carbon dioxide was also is also very efficient this It found that the higher acids are more it is than the lower ones. fatty acids etc..) formation. the eggs will segment and develop If care normally.g.g. that unfertilized egg. and alkali. The nature of this injury became when it was discovered that all the agencies which cause i. digitalin. or after and the eggs perish the first nuclear division. direction. namely oleic acid or one of or esters. It thus happens that the . also in unfertilized eggs.Influence of Environment on Animals butyric. in In all cases in which the unfertilized egg has been treated as to cause it such a way is to form a membrane it begins to develop. It is possible that the sea-urchin injured in the process of clear membrane formation. ^vith a butyric acid.. The physico-chemical process which underlies the formation of the membrane seems to be the cause of the development of the egg. when transferred back to normal sea-water. or saponin. all taken. or ether. the majority of which may be perfectly normal and fertilized may live as long as larvae produced from eggs egg is with sperm. and possible that the sperma- tozoon induces membrane formation by carrying into the egg a its salts higher fatty acid. But if we treat the same eggs after membrane (at 15° C.e. is all For the eggs of certain animals membrane formation required to induce a complete development of the e. Thus the unfertilized eggs of the sea-urchin Strongylocentrotus purpuratus of the Californian coast begin to develop when membrane formation has been induced by treatment fatty acid. solanin. the destruction of the red blood corpuscles. e.. but the development soon ceases in the early stages of segmentation. or chloroform. e.g. hemolysis.. for from thirty-five to fifty-five minutes with sea-water the concentration (osmotic pressure) of a definite of which has been raised through the addition of amount is some salt or sugar.. in the star-fish and certain annelids. etc. For the eggs of other animals a second treatment is necessary. cause membrane formation alcohols bile salts. practically the eggs can be caused to develop into plutei.

formation. all we the eggs form the typical membrane washing in of fertilization. such eggs are after- ward treated for a short period with hj^Dertonic sea-water they . cytolysis is due to a liquefaction compounds. a few minutes after membrane cytolysis. into larvae. therefore. caused by a is superficial or It is possible that the subsequent treatpartly needed to ment layer. The writer found last year that the blood of mammals. If the eggs are then taken out of the traces of saponin saponin solution. after a few minutes.. undergo cytolysis into the blood of an animal which belongs to a if injected different family. however. while if the cytol>i:ic agencies have time to act on the whole egg the latter is completely destroyed.g. as a rule. of the egg with hypertonic sea-water overcome the destructive effects of this cytolysis of the cortical Many pathologists assume that hemolysis or of certain fatty or fat-hke cell. that the blood corpuscles. It is well knoAATi possibly. e. The difference between cytolysis (or hemolysis) and fertilization seems to be this. they develop If. is. and cattle.202 The Mechanistic Conception of Life phenomena of artificial parthenogenesis are linked together with the phenomena of hemolysis which at present play so important a role in the study of immunity. pig. in the If this view is would be necessary to ascribe the fertilization of the egg to the same process. correct. the disintegration knoTvn in pathology as Membrane formation incomplete cytolysis. that the latter is caused by a superficial cytolysis of the egg. If we put unfertilized eggs of a sea-urchin into sea-water which contains a trace of saponin notice that. freed from all by repeated sea-Avater for normal sea-water. it the so-called lipoids. they are left in the sea-water con- taining the saponin they undergo. The analogy between hemolysis and fertilization throws. some light on a curious observation. and transferred to the hypertonic from thirty-five to fifty-five minutes. causes the egg of Strongylocentrotus to If form a typical fertilization membrane. the rabbit.

which do not form membranes. Presumably some blood enters the eggs and causes the cj'tolytic effects in a less degree than is necessary for membrane formation. The slightness of the cytolytic effect allows the egg to develop without treat- ment with hypertonic of cytolysis sea-water. The question may be raised whether the larvae produced parthenogenesis can reach the mature stage. We can also cause the development of the sea-urchin egg without membrane formation. however. of fertilization It is is probable that in this case the mechanism as in the case where the essentially the is membrane formation membrane blood. develop directly into normal larvae without any treatment with hypertonic sea-water. by artificial This question may be answered in the affirmative. carries another substance into the egg which counteracts the deleterious effects underh'ing or following membrane formation. This inference corroborated by observations on the fertilization of the sea-urchin egg with ox very frequently happens that not in this process. but perish tonic sea-water. that the cytolytic effect when no is fertilization formed. but in a sufficient degree to cause their development. since Delage has succeeded in raising several parthenogenetic sea-urchin larvae . Since the entrance of the spermatozoon causes that degree which leads to membrane formation. with this difference less only. all of the eggs form membranes Those eggs which form membranes if begin to develop. presumably. it is probable that. It is brought about. 203 Some substance contained in the blood causes. provided they are exposed to the blood for only a few minutes. writer were done in this The early experiments of the way and many same is experimenters still use such methods. they are not treated ^vith h^'per- Some of the other eggs. a superficial cytolysis of the egg and thus starts its development.Influence of Environment on Animals develop into normal larvae (plutei). in addition to the cytolytic or membrane-forming subit stance (presumably a higher fatty acid).

There is. from lime have a tendency The writer has recently found that twins can be produced not only by the absence of lime. The procedure for the production of twins in the sea-urchin this: egg consists simply in solution of the eggs are fertiUzed as usual in in a neutral normal sea-water and then. namely that the solution used for the production of twins must have a neutral or at least not an alkaline reaction. —The reader probably familiar with the fact that there exist two different t>T)es of human twins. This separation can be brought about by a change in the chemical constitution of the sea-water. the same egg. but also through the absence of sodium or of potassium through the absence of . in other words. fall the cells into which the egg divides have a tendency to free apart. In the second type the twins have invariably the same sex and resemble each other most closely. however. Driesch afterward noticed that eggs of the sea-urchin is treated \\dth sea-water which to give rise to twins. In the one type the twins differ as same parents born at different much as two periods they may or may not have the same sex. namely. a second condition. while twins of the former type are produced from two different eggs.204 The Mechanistic Conception of Life beyond the metamorphosis into the adult stage and since in all the experiments made by the writer the parthenogenetic plutei lived as long as the plutei produced from fertilized eggs. that the two cells into which the egg divides after fertilization become separated from each other. The experiments first of Driesch and others have taught us that twins originate from one egg in this manner. T^\'ins of the latter type are produced from children of the . Herbst observed that if the fertilized eggs of the is sea-urchin are put into sea-water which freed from calcium. c) On the production of twins from one egg through a charige is in the chemical constitution of the sea-water. after repeated washing sodium chloride (of the concentration of the sea- . one or two of the three important metals in the sea-water.

It is its career as a also possible to control the tendency of such twins to grow together sea-water. If. each of these two cells develops into an independent embryo. and potassium chlorides the propor- tion in which these salts exist in the sea-water) the tendency of the twins to grow together is much more pronounced than if we use simply a mixture of sodium chloride and magnesium chloride. way which 55 to 57). revealed by observation This of the first segmentation of the egg in these solutions. or heal together so completely that only slight or even no this imperfections indicate that the individual started pair of twins. sodium. are placed in a neutral mixture of potassium chloride and calcium chloride. contains since it Since normal sea-water all three metals. as the result of alteris ing the composition of the sea-water. The mechanism of the origin of twins. cell-division is modified in a cells (see Figs. or of sodium chloride and calcium sodium chloride and magnesium chloride.water and allowed to develop. or of chloride. we perceive the reason why twins are not normally produced from one egg. for some reason. These twins may remain separate or grow partially together and form double monsters. or of sodium chloride and potassium chloride. leads to a separation If of the first two the egg is afterward transferred back into normal sea-water. and has besides an alkaline reaction. of sodium. into They are then transferred normal sea. and potassium. These experiments suggest the possibility of a chemical cause for the origin of twins from one egg or of double monstrosities in mam- mals. From 50 to 90 per cent of the eggs of Strongylocentrotus purpuratus treated in manner may develop into twins. If by a change in the constitution of the we use as a twin-producing solution a mixture (in magnesium. calcium. The eggs must remain in this solution until half an hour or an hour after they have reached the two-cell stage. the liquids which surround the first cell-division human egg a short time before and after the .Influence of Environment on Animals 205 water).

— In conclusion it may be pointed out that the reverse result. 57 Fig. 57) are separated from each other. 55 Fig. is the fertilization membrane. The two daughter-cells imtil its long axis touches the fertilization membrane. 58 Cell-division in the egg of Strongylocentrotus purpuratus Figs.206 The Mechanistic Conception of Life and at the are slightly acid. 54 Cell-division in a sea-urchin egg. 55-58. or NaCl is lacking. instead of remaining connected If about one hour later the eggs are put as in the normal cell-division (Fig. — M the first two cells and the formation of identical twins are provided. formed (Fig. which leads to the formation of twins. Fig. back into normal sea-water each of the two ceUs develops into an embryo (Fig. the fusion of normally double organs. 52 Fig. Stockard succeeded in causing . or CaClj. P a layer of colloidal embryo from an egg. This type of cell-division leads to the formation of one tus. namely. 51 Fig. 58). can also be brought about experimentally through a change in the chemical constitution of the sea-water. 51-54. 51) is dissolved. in normal sea-water. 53). 56 Fig. 53 Fig. Strongylocentrotus purpuraFigs. same time lacking in one of the three important metals. This cell-division can be observed if the egg is put after fertilization into a neutral mixture of salts in which either KCl. which seems to serve the purpose of keeping all the cells of an egg substance together. and the egg thus gives rise to two instead of to one embryo. the conditions for the separation of Fig. In such a neutral solution the substance which forms the elastic layer (PM. During the segmentation the protoplasm of the egg spreads Fig.

^ We have confined ourselves to a discussion of rather simple change in the constitution of the sea-water upon It is effects of the development. which in all cases gives more or less to result When evidence of its double nature.c. about 50 per cent of the eggs gave one-eyed embryos. the embryos were studied the one-eyed condition was found from the union or fusion of the ''Anlagen" of the two eyes. it appeared as though the optic vesicles were formed too far forward and ventral. however. as yet impossible to connect in a way the effects produced in this and similar cases with the cause which produced them. Entwicklungsmechanik. he added about 6 grams of magnesium rise to chloride to 100 of sea-water and placed the fertilized eggs in the mixture. 1907. 249. define in a simple III. and constitution of the sea-water.Influence of Environment on Animals the two eyes of a fish 207 embryo {Fundulus heteroclitus) to fuse into a single cyclopean eye through the addition of chloride to the sea-water. a priori obvious. Archiv f. so that their antero-ventro-median surfaces fused. rational is. Cases were observed which showed various degrees in this fusion. XXIII. and it is also impossible to way the character of the change produced. as a rule. It has often been noticed a) — by at explorers who have had a chance to compare the faunas in different climates that in the polar seas such species as thrive all in those regions occur. magnesium When c. This produces one large optic cup. . THE influence OF TEMPERATURE The influence of temperature upon the density of pelagic organisms and the duration of life. that an unlimited variations number of pathological might be produced by a variation in the concentration and experience confirms this statement. As an example we in the may mention the abnormalities observed by Herbst It development of sea-urchins through the addition of lithium to sea-water. however. This refers to those members of the fauna which 1 live at or near Stockard. in much greater density than they do in the moderate or warmer regions of the ocean.

the temperature of find an astonishingly rich "In the icy water below 0° C. and the same has been found tempera- ture on the velocity of physiological processes. that the quantity of plankton material exceeds that of the temperate and warm seas. which is condition ^\^th which we are familiar in the Arctic seas is repeated here. Robertson found that the rate of heart beat in the tortoise and in Daphnia is reduced to about one-half if the temperature is lowered 10° C. The same effect of temperature upon the rate of development holds for the egg of the frog. Keith Lucas. he states: fish. . Thus Snyder and T. Peter observed that the rate of development in a sea-urchin's egg is reduced to less than one-half is if the temperature (within certain limits) reduced by 10 degrees. calls especial In his account of the Valdivia expedition. case of the latter van't Hoff In the in had shown that a decrease for the influence of temperature by 10 degrees reduces their velocity to one-half or less. Hertwig. p. 1903. as Cohen and Peter calculated from the experiments of 0. conclusion that life Recent investigations have led to the are affected phenomena by temperature the same sense as the velocity of chemical reactions. Chun^ attention to this quantitative difference in the surface fauna and flora of different regions. Aus den Tie/en des Weltmeeres.208 The Mechanistic Conception of Life the surface. namely. Kerguelen ''The ocean is alive with transparent jelly Ctenophores (Bolina and Callianira) and of Siphonophore of this general observation lies colonies of the genus Agalma. in regard to the pelagic fauna in the region of the Islands. and Snyder found the same influence of temperature for the rate with which an impulse travels in the nerve. 1 Chun. and Maxwell. 225. B.'' The paradoxical character in the fact that a low temperature retards development. since they alone lend themselves to a statistical comparison." And again. and effect hence should be expected to have the opposite from that in mentioned by Chun. we The same animal and plant life. of the Antarctic. Jena.

therefore prolongs the life of the organism 2^^. namely. must be denser The excessive of increase of the duration of at the poles will necessitate the simultaneous existence of more successive generations the same species in these regions than in the temperate or tropical regions. temperature of the duration of life may possibly only life of organisms.Influence of Environment on Animals The of 209 writer found the same temperature coefficient for the rate maturation of the egg of a mollusk (Lottia). exactly than 10° in regions the reverse of what we should expect when authors more temperate state that the density of organisms at or near the surfac e of the ocean in polar regions is greater than in regions. i. therefore. 1 The high coeflEicient of be found near the upper temperature limit for the suflScient for our theory. of about one Since this prolongation of life is far in excess the retardation of development through a it is lowering of temperature. But this is . fertilized The experiments were made on the unfertilized eggs of the sea-urchin. It is. the cause of natural death. the duration of life was about doubled. of obvious that.e. and temperature of which is about 20° C. animal life development than in in there temperate or tropical life seas. its The in writer believes that this paradox finds explanation experiments which he has recently made on the influence of temperature on the duration of life of cold-blooded marine and and yielded the result that for the lowering of temperature by 1° C. in spite of the retardation Arctic seas. over a thouit sand times. Lowering the temperature by 10 degrees animals. development of where the temperature is near the freezing point of water. and a lowering by 20 degrees prolongs million times. must be from two to three times smaller animal life All these facts prove that the velocity of in Arctic regions. where the temperature of the ocean is about four to nine times smaller than in seas the from C.^ The writer is inclined to believe that these results have role some bearing upon a problem which plays an important in theories of evolution..

210 It The Mechanistic Conception of Life has been stated that the processes of differentiation and of the individual. and. Later he generalized this idea and stated that other cells may be considered immortal in the same sense in which AVeismann claimed this for the unicellular organisms. that the process of senile decay is essentially different from that of growth and development. chemical processes which determine death are certainly not identical with the processes which underlie their development. Robertson has also arrived at the conclusion. for quite different reasons. for a change of 20° altered this more than a hundred thousand times the much. velocity must be reduced by the same amount if the temperature is lowered to the same extent. The temperature coefficient for the duration of life of cold-blooded organisms seems. while he assiimed that all the other organisms with the exception of their germ-plasm are mortal. One can indeed well imagine that the same piece of skin might be transplanted through an indefinite series of generations of mice and that such a transplanted piece might outlive an indefinite number of generations of mice in exactly the same way as a cancer cell does. means of identification is the temperature One When two chemical processes are identical. he pointed out that this proved that the principle of immortality must also be granted to cancer-cells (1901). their coefficient. If respiratory motions and circulation could be maintained indefinitely even the metazoa might be foimd to be immortal.^ development lead also to the natural death If we express this in chemical terms it means that the chemical processes which underlie development also determine natural death. For a difference altered five in temperature of 10° C. m . death of the cells through lack of oxygen. to differ enormously from the temperature coefficient for their rate of development. C. 1 Weismann showed that infusorians or Tinicellular organisms general are immortal. the duration of life is hundred times as much as as the rate of development. at least for the sea-urchin eggs embryo. and since it is the originally transplanted cancer-cell and the cells derived from it by multiplication that survive. it is From and natural we may conclude that. Physical chemistry has taught us to identify two chemi- cal processes even if only certain of their features are knoA\Ti. B. T. howof these ever. The natiiral death of the metazoa is perhaps a secondary phenomenon due This leads to the to the cessation of respiratory motions or of the heart beat. Leo Loeb first called attention to the fact that the transplantation of a cancer can be repeated to an milimited series of generations.

Influence of Environment on Animals
h)

211

Changes in
of

the color of butterflies

produced through the

influence

temperature.

—The

experiments of Dorfmeister,

Weismann, Merrifield, Standfuss, and Fischer on seasonal dimorphism and the aberration of color in butterflies have so often been discussed in biological literature that a short reference to them will
suffice.

the fact that species
in a

By seasonal dimorphism is meant may appear at different seasons of the year
form or
color.

somewhat

different

Vanessa prorsa

is

the

summer form, Vanessa
species.

levana the winter form of the

same

By

keeping the pupae of Vanessa prorsa several
0° to 1°

weeks at a temperature of from
in

Weismann succeeded

obtaining from the

summer

chrysalids specimens which

resembled the winter variety, Vanessa levana.
If

we wish

to get a clear understanding of the causes of

variation in the color and pattern of butterflies,

we must

direct

our attention to the experiments of Fischer,

who worked with
and foimd
This

more extreme temperatures than

his predecessors,

that almost identical aberrations of color could be produced

by both extremely high and extremely low temperatures.
observations.

can be seen clearly from the following tabulated results of his

At the head

of each

to which Fischer submitted the pupae
vertical

column the temperature is given, and in the
varieties that

column below are found the
In the vertical column

were pro-

duced.

A

are given the normal forms:

0°to-20°C.

212

The Mechanistic Conception of Life
The reader
will notice that the aberrations

produced at a
are absolutely

very low temperature (from 0° to
identical with the aberrations

—20°

C.)

produced by exposing the pupae

to extremely high temperatures (from 42° to 46° C.)

Moreover

the aberrations produced by a moderately low temperature

(from 0° to 10° C.) are dentical with the aberrations produced

by a moderately high temperature (from

36° to 41°

C).
it is

From

these observations Fischer concludes that

errone-

ous to speak of a specific effect of high and of low temperatures,

but that there must be a

common

cause for the aberration found

at the high as well as at the low temperature limits.

This cause

he seems to find in the inhibiting

effects of

extreme temperatures

upon development.
If

chemical point of view,

we try to analyze such results as Fischer's from a physicowe must realize that what we call life
which are connected way; inasmuch as one reaction or group of reacoxidations).

consists of a series of chemical reactions,
in a catenary

tions (a) (e.g., hydrolyses) causes or furnishes the material for

a second reaction or group of reactions,

(6) (e.g.,

We know
it is

that the temperature coefficient for physiological

processes varies slightly at various parts of the scale; as a rule

higher near 0° and lower near 30°.

But we know

also that

the temperature coefficients do not vary equally for the various
physiological processes.
It
is,

therefore, to be expected that

the temperature coefficients for the group of reactions of the

type

(a) will

not be identical through the whole scale with the
(6).
If,

temperature coefficients for the reactions of the type
therefore, a certain substance
is

formed at the normal tempera-

ture of the animal in such quantities as are needed for the

catenary reaction
perfect

(6), it is

not to be expected that this same

balance will be maintained for extremely high or

extremely low temperatures
of reactions will

more probable that one group exceed the other and thus produce aberrant
;

it is

Influence of Environment on Animals
chemical
effects,

213

which

may

miderlie the color aberrations

observed by Fischer and other experimenters.
It
is

important to notice that Fischer was also able to produce

aberrations through the application of narcotics.

Wolfgang

Ostwald has produced experimentally, through variation of
temperature, dimorphism of form in Daphnia.
IV.

THE EFFECTS OF LIGHT
seriously questions the statement
is

At the present day nobody
chemical
character.

that the action of light upon organisms

primarily one of a
action
is

While
for

this

chemical

of

the

utmost importance

organisms,

the nutrition
it

of

which
of less

depends upon the action of chlorophyll,

becomes

importance for organisms devoid of chlorophyll.

Nevertheless,

we

find animals in
is

which the formation

of organs

by regeneralight.

tion

not possible unless they are exposed to

An
be

observation

made by

the writer on the regeneration of polyps
at

in a hydroid,

Eudendrium racemosum,
of this.
If
is

Woods

Hole,

may

mentioned as an instance
which
is

the stem of this hydroid,

usually covered with polyps,
fall
off.

put into an aquarium the

polyps soon

If

the stems are kept in an aquarium
of

where

light strikes

them during the day, a regeneration
few days.
If,

numerous

pol^'ps takes place in a

however, the

stems of Eudendrium are kept permanently in the dark, no
polyps are formed even after an interval of some weeks; but

they are formed in a few days after the same stems have been
transferred from the dark to the light.
suffices for this effect.

Diffused dayhght
these experi-

Goldfarb,

who repeated

ments, states that an exposure of comparatively short duration
is

sufficient to

produce this

effect.

It is possible that the light

favors the formation of substances which are a prerequisite
for the origin of polyps

and

their growth.
this observation are the

Of much greater significance than

214
facts

The Mechanistic Conception of Life
which show that a large number
extent, the color of the
of animals assume, to

some

ground on which they are placed.
fish
is

Pouchet found through experiments upon crustaceans and
produced through the medium of the eyes.
If

that this influence of the ground on the color of animals

the eyes are

removed or the animals made blind in another way these phenomena cease. The second general fact found by Pouchet was that the variation in the color of the animal is brought about through an action of the nerves on the pigment cells of
the skin; the nerve action being induced through the agency
of the eye.

tion were

The mechanism and the conditions for the change in coloramade clear through the beautiful investigations of
cells can, as

Keeble and Gamble, on the color change in crustaceans.
According to these authors the pigment
a rule, be

considered as consisting of a central bodj^ from which a system
of

more or
all

less

complicated ramifications or processes spreads

out in

directions.

As a

rule,

the center of the

cell

contains

one or more different pigments w^hich under the influence of
nerves can spread out separately or together into the ramifications.

These phenomena

of spreading

and retraction

of the
cells

pigments into or from the ramifications of the pigment

form on the whole the basis
influence of environment.

for the color changes

under the

that Macromysis flexuosa

Thus Keeble and Gamble observed appears transparent and colorless

or gray on sandy ground.

On

a dark ground their color
in their

becomes darker.

These animals have two pigments

chromatophores, a brown pigment and a whitish or yellow

pigment; the former

is

much more
cells,

plentiful
all

than the

latter.
is

When

the animal appears transparent

the pigment

con-

tained in the center of the

while the ramifications are free

from pigment.

When the

animal appears brown both pigments
In the condition of maximal

are spread out into ramifications.

spreading the animals appear black.

Influence of Environment on Animals
This
is

215

a comparatively simple case.

cated conditions were found
crustaceans,
e.g., in

Much more compliby Keeble and Gamble in other

Hippolyte cranchii, but the influence of the

surroundings upon the coloration of this form was also satisfactorily analyzed

by these authors.

While

many

animals show transitory changes in color under

the influence of their surroundings, in a few cases permanent

changes can be produced.

The

best examples of this are those

which were observed by Poulton in the chrysalids of various
especially the small tortoise-shell. These experiments are so well known that a short reference to them will suflfice. Poulton^ found that in gilt or white surroundings the
butterflies,

pupae became light colored and there was often an immense development of the golden spots, ''so that in many cases the
whole surface of the pupae glittered with an apparent metallic
luster.

So remarkable was the appearance that a physicist,

chrysalids, suggested that I had played a and had covered them with goldleaf." When black surroundings were used, "the pupae were as a rule extremely dark, with only the smallest trace, and often no trace at all, of the golden spots which are so conspicuous in the lighter form."

to

whom I showed the

trick

The
ings

susceptibility of the animal to this influence of its surround-

was found to be greatest during a definite period when the caterpillar undergoes the metamorphosis into the chrysalis stage. As far as the writer is aware, no physico-chemical explanation, except possibly Wieners' suggestion of color photography by
mechanical color adaptation, has ever been offered for the
results of the type of those observed
V.

by Poulton.

EFFECTS OF GRAVITATION
the egg of the frog.

a)

Experiments on
life

— Gravitation can only

indirectly affect
cell
1

two

different non-miscible liquids (or a liquid
Colours of Animals
p. 121.

phenomena; namely, when we have in a and a solid)
("International Scientific Series"),

Poulton, E. B.,

London, 1890,

216 The Mechanistic Conception of Life change in the position of This of different specific gravity. that the tendency of the black part of the egg to rotate upward along the surface of the egg leads to a separation of its first cells. possible to prevent or retard this rotation of the highly viscous protoplasm. thus confirming an older observation of When. however. compression experiments by Such may lead to rather interesting results. The dark one occupies normally the upper position in the egg and may therefore be assumed to possess a smaller specific gravity than the white substance. in this case. Pfliiger as 0. Obviously in this case the tendency . Morgan made an half interesting additional observation. through compression between two glass plates for about twenty time of its first division hours. which remained alive gave only one-half of an embryo. the frog's egg turned upside do\Mi at the and kept in this abnormal position. had already shown is that the first plane of division in a fertilized frog's egg vertical is and Roux established the Schultze found that if fact that the first plane of division identical with the plane of symmetry is of the later embryo. a small number of eggs may give rise to twins. nicely illustrated very very by the frog's egg. first He destroyed one-half of the egg after the segmentation rise to and found that the Roux. It is possible. the egg is turned with the white pole upward a tendency of the white protoplasm to flow itself. H. however. Schultze first pointed out. such a separation leading to the formation of twins. Morgan put the egg upside do^\^l after the destruction of one of the first two cells. and compressed the eggs between two glass plates. the surviving half of the egg gave rise to a perfect embryo of half-size (and not to a half-embryo of normal size as before). T. is which has two layers of viscous protoplasm one of which black and one white. dowTi again manifests When It is. so that a the cell or the organ may give results which can be traced to a is change in the position of the two substances. compressing the eggs between horizontal glass plates.

Influence of Environment on Animals of the 217 protoplasm to flow back to its normal position was partially successful of the living and led to a partial or complete separation from the dead half. —A striking influence of gravita- tion can be observed in a hydroid. from the Bay of Naples. change in the hereditary arrange- or the transformation of organs by external forces heteromorphosis. It is occasionally also from the short lateral branches. called the The writer had ment of organs this subject. whereby the former was enabled to form a whole embryo. however. b) Experiments on hydroids. to roots. while the stem had remained in lateral branches. We cannot now go any farther into in rela- which should.e. prove of interest tion to the problem of heredity. of course. straight. i. main stem which grows vertically upward and like lateral branches. possessed only half the size of an embryo originating from a whole egg. new stems grow possible to force mostly directly from the original stem. and that wherever . which has at regular intervals very fine and short bristleon the upper side of which the polyps grow. its apex upward when we put it obliquely roots grow vertically downward. which. one might conclude that the cells of substances of different specific gravity. namely. into the The writer observed that when the stem is put horizontally continues to grow vertically into the water the short lateral branches on the lower side give rise to an altogether different kind of organ. and these roots grow indefinitely in length if and attach its themselves to solid bodies. The main stem is negatively geotropic. This hydroid consists of a long.. thus upon this hydroid an arrangement of organs which is altogether different from the hereditary arrangement. Antennularia antennina. normal position no further growth would have occurred in the From the upper side of the horizontal stem out. while the aquarium. If it is correct to apply inferences drawn from the observaAntennularia also contain non-miscible tion on the frog's egg to the behavior of Antennularia.

they are expected to lead to any results. to diffused daylight from a window. a discussion of experiments on the physico-chemical character of the instinctive reactions of animals should not be entirely omitted from this sketch. they move toward the source of light in as straight a and peculiarities of their line as the imperfections locomotor apparatus will permit. (after When (when we expose winged aphides plant).g.218 The Mechanistic Conception of Life the specifically lighter substance comes in contact with the sea-water (or gets near the surface of the cell) the growth of a stem is favored. The ful simplest type of instincts represented by the purposee. as it may preferably be termed. It is also obvious that we are here dealing with a forced reaction in which the animals have no more choice in the direction of their motion than have the iron . or if the animals are thev are rendered sensitive through the agencies which we shall mention later. motions of animals to or from a source of energy. the experimental control of animal instincts Experiments on the a) mechanism of heliotropic reactions in animals. — Since the instinctive reactions of animals are as hereditary as their morphological character. light. If animals to move toward the source of naturallv sensitive. the reactions of which are not complicated by associative memory is or. will favor the formation of roots. while contact with the sea-water of the specifically heavier of the substances.. or they have flown away from the young caterpillars of Porthesia chrysorrhoea they are aroused from water copepods and falling in their winter sleep). and if the light is strong enough. we notice a tendency among these light. it is and with some of these that we intend to deal here. and it is also obvious that only such animals must be selected for this purpose. VI. associative hysteresis. or marine or fresh- many other animals. It is obvious that such experiments must begin with if the simplest type of instincts.

Under such conditions the animals actually died from starvation. the light preventing them from turning to the food. a peculiar photosensitiveness of the retina (or skin). the symmetrical muscles work with equal energy struck by stronger light than the as long as the photochemical processes in both eyes are identical. since it if can be showTi that they go toward the source of light in so doing they even move from places of a higher to places of a lower degree of illumination. If. are attracted by the light. This can be proved very nicely in the case of starving caterpillars of Porthesia. the symmetrical muscles will and in positively heliotropic animals those muscles will work with greater energy which brings the plane of symmetry back into the direction of the rays of light and the head toward the source of light. Animals of the type of those mentioned are automatically oriented by the light in such a way that symmetrical elements of their retina the rays of light at the same angle. and second a peculiar nervous connection between the retina and the muscular apparatus.Influence of Environment on Animals filings in their 219 arrangement in a magnetic field. The writer put such caterpillars into a glass tube the of the axis of which was at right angles to the plane window: the caterpillars went to the window side of the tube and remained there. is (or skin) are struck by In this case the intensity the same for both retinae or symmetrical parts of the This automatic orientation first is determined by two factors. light allowed them to do so. one eye is other. In symmetrically built heliotropic animals in Avhich the symmetrical muscles participate equally in locomotion. The writer has advanced the following theory of these instinctive reactions. of light skin. even if leaves of their food plant were put into the tube directly behind them. which we call positively heliotropic. however. As soon as both eyes are struck by the rays of light unequall}' work . which they eagerly ate when the One cannot say that these animals.

The former worked on a butterfly. Additional proof for the correctness of this theory was furnished through the experiments of G. of positive heliotropism hy acids The production means and the periodic depth migrations of pelagic animals. All the animals were in a marked that if degree positively heliotropic. therefore. — When we observe a dense mass of copepods collected from a some have a tendency to go to the light while others go in the opposite direction and many. that it is fresh- water crustacean. the latter on other arthropods. howas a rule necessary to lower the temperature of . as h) if The animal behaves. Parker and S. H. notice that we to make the negatively heliotropic or the indifferent copepods almost instantly positively heliotropic by adding a small but definite amount of carbon dioxide in the form of carbonated If water to the w^ater in which the animals are contained. Similar results may be obtained with any other The same experiments may be made with another ever. the carbon dioxide has again diffused into the air). animals are contained in 50 3 to 6 cally c.c.220 at the The Mechanistic Conception of Life same angle. namely Daphnia.c. These authors found it one cornea is blackened in such an animal. c. Vanessa antiope. moves light. there is no more reason it for the animal to deviate from this direction and All this holds will move in a straight line. are indifferent to light. Holmes. good on the supposition that the animals are exposed to only one source of light and are very sensitive to light. make the copepods energetilasts positively This heliotropism all half an hour (probably until acid. It is an easy matter if not the majority. with this difference. is continually in a circle when it is exposed to a source of is and in these motions the eye which not covered with paint directed toward the center of the circle. and other the darkened eye were in the shade. the of water all it suffices to add from about of carbonated water to heliotropic. fresh-water pond. J.

is The tendency to migrate the outcome of the fact that periodically varjdng external conditions induce a periodic change in the . These data have a bearing upon the depth migrations of pelagic animals. 221 If the water containing the Daphniae is cooled same time carbon dioxide added. Groom kno\Mi that and the writer. the tension must rise and this must have the effect of inducing positive heliotropism or increasing its intensity. At the same time the temperature of the water near the surface is lowered and this also increases the positive heliotropism in the organisms. tively heliotropic IMarine copepods can be of the made posi- by the lowering temperature alone. mitted in this case is not the tendency to migrate periodically upward and do\\Tiward. It is well many animals living near the surface of the ocean or fresh-water lakes. The faint light in from the sky is sufficient to cause animals which are vertically a high degree positively heliotropic to light. morning. These experiments have also a bearing upon the problem The character which is transof the inheritance of instincts. the animals which were before indifferent to light now become most strikingly and at the positively heliotropic. have a tendency to migrate upward toward evening and do'^Tiward in the morning and during the day.Influence of Environment on Animals the water also. the animals and slowly sink doAMi or become negatively heliotropic and migrate actively do^^•nward.g. When. the absorption of carbon dioxide by the green algae begins again and the temperature of the water lose their positive heliotropism. if These periodic motions not exclusively.. copepods. T. as move in the upward toward the experiments with such pelagic animals. dioxide Since the consumption of carbon of this gas in the water by the green plants ceases toward evening. by the are determined to a large extent. have shoAMi. but the positive heliotropism. or by a sudden increase in the concentration of the sea-water. as was pointed out years ago by Theo. heliotropism of these animals. e. rises.

when they are first aroused from This heliotropic sensitiveness lasts only as If long as they are not fed. Davenport and Cannon found that Daphniae. D. these animals take be inferred that as soon as up food. The caterpillars of Porthesia chrysorrhoea are very strongly positively heliotropic their winter sleep. react much more quickly after they have been made to go to the light a few times. carbon dioxide. Waller in the case of the response of a nerve to stimuli. is of course immaterial for the result. the formation of a substance or substances in their bodies takes place. We chemical notice also the reverse phenomenon. The writer is inclined to attribute this result to the effect of acids. A similar effect of the acids was shown by A. that its changes produced in the animal destroy heli- otropism.222 The Mechanistic Conception of Life It sense and intensity of the heliotropism of these animals. they are kept permanently without until food they remain permanently positively heliotropic It is to they die from starvation.. of the experiment react sluggishly to which at the beginning light. Since the workers show no heliotropism V. The writer observed many years ago that winged male and female ants are positively heliotropic and that their heliotropic sensitiveness increases and reaches its maximum toward the period of nuptial it flight. whether the carbon dioxide or side or any other acid diffuse into the animal from the outtissue-cells of whether they are produced inside in the the animals. Kellogg has looks as if an internal secretion from the sexual glands were the cause of their heliotropic sensitiveness. the bees are prevented from leaving the hive through the exit at the lower end. in fact so much so that by letting light fall into the observation hive from above. .g. diminishing or annihilating their heliotropic sensitiveness. e. produced in the animals themselves in consequence of their motion. namely. observed that bees also become intensely positively heliotropic at the period of their wedding flight.

and the latter at embryo itself. it is surprising that certain tissue-cells also show reactions which tissue-cells belong to the class of tropisms. repeated with the same result in Volvox. young fish entirely. sex-cells of such instincts as are based upon heliitself This problem reduces simply to that of the method whereby the gametes transmit heliotropism to the larvae The writer has expressed the idea that all or to the adult. For the transmission of this the gametes need not contain anything more than a catalyzer or ferment for the synthesis of the photosensitive substance in the body of the animal. briefly discuss the question of the transmission. to geotropism and stereotropism. 106). What if has been said in regard to animal heliotropism might. p. are of special interest These reactions of of their bearing by reason upon the of this inheritance of morphological characters. 36. mutatis mutandis..Influence of Environment on Animals The on the heliotropism of 223 animals is identical ^\^th the heli- otropism of plants. be extended. of peculiarities (e. therefore. that is necessary for this transmission is the presence of a pho- tosensitive substance in the eyes (or in the skin) of the animal.g. The chromatophores are at first scattered irregularly over the yolk sac and show their characteristic ramifications (Fig. tropic c) The morphogenetic reactions of certain tissue-cells and the effects of these reactions. There is at that time no definite . is An example of the found in the tiger-like in the marking of the yolk sac of the of Fundulus and marking is writer found that the former in part. erroneous to try to explain these heliotropic reactions of animals on the basis in plants. The least due to the creeping of the chromatophores upon the blood-vessels. —Since plant-cells show not heliotropic reactions identical with those of animals. space permitted. vision) which are not found We may through the otropism. The writer has sho\\Ti that the experiments of effect of acids on the heliotropism copepods can be It is.

and Driesch has showTi that these migrate actively to the place of their destination. seem to indicate the in possibility that in the adult the some forms at least. and that the oxygen tions of the blood may be the cause of the spreading of the Certain observa- chromatophores around the blood-vessels. they nevertheless reached their destination. Herbst pointed out that this might be a case of chemotropism. until finally all the chromatophores cells form a sheath around the vessels and no more pigment are found in the meshes between the vessels (Fig.224 relation The Mechanistic Conception of Life between blood-vessels and chromatophores. a more rigid structure and are prevented from acting in the way indicated. first marking the embryo The writer is inclined to believe that we are here dealing with a case of chemotropism. Driesch has found that a tropism underlies the arrangement of the skeleton in the pluteus larvae of the sea-urchin. 37) and forms a complete sheath aromid the vessel. migrate to the peripherj^ of the egg. 38). mission of certain markings. Nobody who has not actually watched the process of the creeping of the chromatophores upon the blood-vessels would anticipate that the tiger-like stages of facts coloration of the yolk sac in the later in this development was brought about way. possibly led there under the influence of certain chemical substances. As soon as a ramification of a chromatophore comes in contact with a blood-vessel the whole mass of the chromatophore creeps gradually on the blood-vessel (Fig. In the developing eggs of insects the nuclei. Similar of can be observed in regard to the itself. When Driesch scattered these cells mechanically before their migration. together with some cytoplasm. The writer has expressed . It seems to the writer that such observations as those made on Fundulus might simplify the problem of the hereditary transchromatophores have. The position of this skeleton of the cells is predetermined by the arrangement mesenchyme cells. caused by the oxygen surrounding the egg.

if therefrom may be determined by forces lying outside the it Since these forces are ubiquitous and constant appears as we were is dealing exclusively with the influence of a gamete. on which they would starve. investigated the conditions for the laj^ng of e. Salatnandra atra and In both forms the eggs are fertilized in the in the uterus. eggs in two forms of salamanders.eggs (from twelve to . body and begin to develop for a few larvae in the uterus. the latter being forced by an outside influence to creep to the surface of the egg.. a large is Smce there is room only number of eggs perish and It this number the greater the longer the period of gestation. man}.Influence of Environment on Animals the opinion that the formation of the blastula 225 may be caused general^ by a tropic reaction of the blastomeres. and not upon the fat. Kammerer has S. —For the preservation of species the instinct of animals is to lay their eggs in places in which the food and can develop of is young larvae paramount importance. AVhen a piece of meat and of fat of the its same animal are placed side by side. maculosa. find their A simple example its of this instinct the fact that the common fly lays eggs on putrid material which serves as food for the young larvae. elapsed since they were fertilized. very few young of advanced stage and these are in a rather development. thus happens that when the animals retain their eggs a long time. d) Factors which determine place and time for the deposition of eggs. the fly will deposit eggs upon the meat on which the larvae can grow. When the animal lays its eggs comparatively soon after copulation. These examples maj^ of definite groups of cells suffice to indicate that the arrangement and the morphological effects resulting cells. owing to the long time which ones are born. Here we are dealing with the effect of a volatile nitrogenous substance which reflexly causes the peristaltic motions for the laying of the egg in the female fly.g. all while in reality that is necessary for the gamete to transmit a certain form of irritability.

if serve as a basis for theories of evolution. law to phenomena of heredity. If they are kept in water or in proximity to water and in a moist atmosphere they have a tendency to lay their eggs earlier and a comparatively high temperature enhances the tendency to shorten the period of gestation. while in later stages they gills and breathe through their lungs. maculosa bears young ones in an earlier stage. show that we are dealing with the direct effects of definite outside forces. must. atra bears young which are already developed and beyond the stage of aquatic life. that . While we may speak of adaptation when all or some of the variables which determine a reaction are unknown.226 The Mechanistic Conception of Life seventy-two) are produced and the larvae are of course in an earty stage of development. has been termed adaptation. while S. gills In the early stage the larvae possess and can therefore live in water. as shown especially by Bateson and his pupils. Vll. that if new species may arise by mutation and the wide not universal applicability of Mendel's not permanently. it is obviously in the interest of further scientific progress to connect cause and effect directly whenever our knowledge allows us to do so. These discoveries It is true place before the experimental biologist the definite task of pro- ducing mutations by physico-chemical means. CONCLUDING REMARKS The discovery of DeVries. for the time being. however. If the salamanders are kept in comparative dr^mess they show a tendency to lay their eggs rather enhances this tendency. the fact that S. Kammerer showed that both forms of Salamandra can be induced to lay have no their eggs early or late. Kammerer's experiments. according to the physical conditions surrounding them. Since Salamandra atra is late and a low temperature found in rather dry alpine regions with a relatively low temperature and Salamandra inaculosa in lower regions with plenty of water and a higher temperature.

since each species has.^ What was said in regard to the production of new species still physico-chemical means may be repeated with problem by more justification in regard to the second of transformation. the regenera- tion of broken crystals and the regeneration of lost limbs by a crustacean were declared identical will not appeal to the biologist. but the writer wishes to apologize to these authors for his inability to convince himself of the validity of their claims at the present moment. which result in the s^mthesis or group of of a definite compound compounds. The enzymes nucleins have the peculiarity of acting as ferments or for their own synthesis. This determines the continuity of a species. But it shows us that whoever claims to have succeeded in making living matter from inanimate will have to prove that he has succeeded in producing nuclear material which acts as a ferment for its own synthesis and thus reproduces this.g. thus far succeeded in it although nothing warrants us in taking is for granted that this task 1 beyond the power Since this was written the beautiful experiments of Kammerer as well as those of Tower seem to have furnished proof that external conditions can cause hereditary changes in animals. Nobody has of science. produced living beings. namely. nucleins. the making of living from inanimate matter. itself. namely. We know that growth and development in animals definite although and plants are determined by series of catenary complicated chemical reactions. Thus a given type of nucleus will continue to synthesize other nuclein of its o^vn kind. or the plays on words by which. probably. purely morphological physicists have The which imitations of bacteria or cells now and then proclaimed as artificially e.. He thinks that only continued breeding of these apparent mutants through several generations can afford convincing evidence that we are here dealing with mutants rather than with merely pathological variations. also its own specific nuclein or nuclear material.Influence of Environment on Animals 227 certain authors claim to have succeeded in this. .

.

21. of. ff Acid poisoning. 15. Bees. Catalyzer. influences of. 80 flf. F. Diflfusion of salts. Arrhenius. death and. 5. 229 . Ascidians.. 218. 58. Baer. Area striata. 220. Antagonism of acids and salts. 202. 99. Consciousness. treatment of egg. 196 Chemical symmetry. Amphitrite. 198. 25. Correns. 198. 16 Chun. 41. Associative memory. 30. 145. 208. Ants. Artificial production of living matter. Catalysis of esters. antagonism. Darwin. 146. 85. Coordinated 70. 138. Antedon rosacea. 132. color adaptation. 198. Blaauw. Antagonistic salt action. changes in. 177 flf. 127 flf. Ctenolabrus. 131. eflfects of. Butyric acid. 39. von. 38. Bases as membrane-forming stances. in reflexes. Bateson. of the egg. 18 Cannon. 116 flf. 92. experimental control flf." 72. 42. Chaetopterus. 176 flf. Chlorostoma. Copepod. Cytolytic agents. 11. Butschli. 226. 226. 15. Agalma. 5. 49. 114. 144. 58. 92. Boveri. Cerianthus membranaceus. "Center of coordination. 38. Bischof. Crabs. 6 fif. Bataillon. 73. Biology. 115. Charles. 117. 68. 58. 7. 43. 47. 208. Asterina. Change in intensity of light. Depth migration. 93. 91. 220. 4. 43. Action of potassium cyanide. through influence of temperatiu*e. Artificial Cohen. 220. beginning of scientific. Death and development. 198. 79. 4. on heliotropism. K. 179 Antagonism of three salts. 203. sub- De Vries.. Biedermann. 4. of.. 27. 37. 60. Contents of life. 209 fif. 145. 53." 113. 213. of butterflies. 179 ff. 222. Ciona intestinalis. dififerent chemical processes. flf. 38. 144 fif.. 43. 43. 72. 17.. 182 flf. Asterias capitata. 11. " Aura seminalis. production of double and Compulsory movements. 156 Activation of the egg. 48. Cuma Rathkii. 5. 198.. 189. 208. Color. flf. 134. Bancroft. Chemical agencies. E. Cytolysis. 173. 172 217. 28 flf. 198.. Aphides. dififerent 54. 55. Balanus perforatus. Changes in color of butterflies produced through influence of temperature. INDEX Abraxas. acids and salts. 16. fiddler. Arabacia. 51. 146. 68. 157. 164. 211. 208. 211. 213 fif. 29. 41. 29. Asterias forbesii. 162. Artificial parthenogenesis. flf. 26 flf. 135. Callianira. 27. Antennularia antennina. 51.. Acids. movements 62. mechanical destruction of. 195. Bolina. 100 flf.. 10. 40. flf. Artificial causation of positive heliotropism. Chromosomes. Baltzer. 4. 199 flf. Ciamician. 46. multiple monstrosities in sea-urchins. 10. Darwin. Cosine law of illumination. Asterias ochracea. Difference of salt permeability of various membranes. 37 flf. Cinerarias.. color blindness. 41 flf. 189 flf. 147. Chemotropism. Daphnia. Bunsen-Roscoe law. Delage. 54 flf. 107. 136. Bohn. Claparede. chemical processes. 49. 10. Cooperative action of salts causing impermeability of the egg membrane. Atwater. 190. 208. Bardeen. 209 flf. 55. 51. Animal instincts. 208. Bunsen. Development. 4. 220. mechanical causation of. 40. Beginning of scientific biology. Barry. 46. Cortical layer.. 11. 196. Cooke. 114. 132 flf. 19. 222. Davenport. 224. 10. Berzelius. of unfertilized egg.

50 ff. 122. Growth and Guyer. 25. Heteromorphosis. growth and. 215.water. 5. heterogeneous. v. 213 effects of. Hybridization. Haberlandt. 207 ff. 121. factors which determine time and place for the deposition of. 217. 101. 89. 4. 186. Dzierzon. mem- brane. 15. 215. 125. Lillie. poisonous 189. 217 fl. cortical layer of unfertilized. 7. 49. to cytolytic agents. 95. Emil. Experiments on. Fertilization. Instincts. 215. Gravitation. 213. Hippolyte cranchii. Fusion of normally double organs. varying susceptibility of. 147 ff. Illumination. 60. Increased sensitiveness of unfertilized egg to cytolytic agents. 136. Light. 204. by animals Hagedoorn. 29. 212. 214. 148. Effects of chemical agencies. 107. 220. 172 fl. Effect of retarded oxidations on poisonous salt action. Huyghens. Intensity of illumination. 7. 15. 36 selection of intensity of. Goldfarb. 8. 44. Heterogeneous hybridization. 207. 15. E. Fischer. 39. 42. animals. HgiiscIigii 79 Herbst. Hydra. 175. 214. 147. 116 fl. Ethics. 225. Keeble. activation of. 204 ff Eggs. 213. 113. of. Froschl. of. 211. 223. 215. Koeppe. 213. Ferments of oxidation. 142... 116. Heredity. photochemical action of. 222. 108. 5. on the egg of the frog. Eudendrium racemosum. of light. 198. Holmes.. Laplace. 199. 54 fl. 199. 220 fl. 58. 30. 25. 3. 179 heteroclitus. 148 fl. Infusoria Coelenterates. animals. 220. inten- Factors place 225. production of twins from. 69. Hartmann. ff. Galvanotropism. on hydroids. 199.. 41 . 132. 190 ff. 21. influence of acids on. 163 fl. 101. 35. 51. 26 fl. 10. . 92. 105. Kammerer. 20 Leeuwenhook. 43. Hoeber. 224. Fundulus 175. on the mechanism 218. 10. Geotropism. 56 Gies. Drosophila.. 52 ff. Dumas. 24. 27 ff.. Cosine law sity of. 43.. fl... 58. 170 ff. 5. 221. otoliths of. heliotropic reactions. 25. 16. 49. of organisms. 10. 24. 134. 142. 204. 95. Fischer. F. 31 fl. 18. Law of segregation. Influence of membrane formation in causing the egg to develop.. 217. change in Intensity of. Kupelwieser. Ear. Knaffl. Gemmill. Hertwig... 187. 131 fl. Lillie. 217. 114. 44. action of distilled water on. 17. Jennings. 6ff. fertilization 6. 12. 196 fl. 101. 16. Heliotropic 35. 41. Jacobi. 189. 60. localization. and orientation to center of gravity of the earth. Huxley. 15 fl. 123. 199. 227. 5. Esters. 208. artificial causation of positive. Fiddler crabs. 85 fl. 52. 57. Egg. fl.. 177. Hydroids. 196 fl. 93 fl. sensitiveness of unfertilized. of gravitation. Holmes. 220. Kellogg. 74. S. 198.. Lavoisier. R. 148. Influence of temperature. 181. effects Groom. 15. EmiUsion theory of membrane formation. 15. experiments on the mechanism of. Harvey. 215.. Henking. 21. 142. which for determine deposition time of and eggs. 21. 4. R. 144. Liebig. 207. 100 mechanics of in Life. ff. 150 ff. 113. von. 62. His. increased . to body extracts of same species. 213. 224. 163 ff. immunity of. Fovea centralis. 100. 213. 223. . of heliotropic reactions. 55. Immunity of eggs to body extracts of same species. 4. France. Hardy. maternal character of heterogeneous. Growth. treatment of. 196 fl. " Life principle.. 145. 211. 230 The Mechanistic Conception of Life Harmonious character 23 fl. Hypertonic sea. 218. 136. 225. von. 189. Dorfmeister. Enzymes. 170 ff. 174. Godlewski. Hypotricha. O. Hertwig. contents of. Gammarus. 213. Heliotropism.... butyric acid." 14. Experimental control of animal instincts. Hybrids. 116. 115. 96 ff. the Immiuiity.. 115. Doncaster. Experiments. 99. 60. 218. 55. Handovski. 226. Driesch. light. 216. 115. Gamble. catalysis of. 134. 5.. 104. J.

diflfercnce of permeability of various membranes. substances. Radl. 79. Roscoe. action of. 177. 210. 144. Photochemical action of Photochemical. C. 36. Oxidation. 186 flf. 100 flf. 218 flf. 38. law of. 104. 190 flf. Mathews.. Ostwald. Planarians. Pennaria. Movements. 150 flf. 16. and their relation to the egg after fertilization. Pauli. 127 flf. Role of water in segmentation. Morgan. 91. membrane formation. 10. 30. 109... 186 flf. 59 flf. 216. Merrifleld. 24. 38. 216. 99. acid. 16. 53. 225. 199. diffusion of. 95. 92 flf. purposeful character of. Maxwell. 15 flf. hypertonic. poisoning. Neuberg. 198. 99. 148. . Ostwald. "Lysin theory. 214. 24. Polarization. 107. 190. 48. Pycnopodia. 20 fl. 114. 59. Porthesia chrysorrhoea. Mytilus. B. 52. Mead. Sachs. 35. 158. 35. 152. 145. 147 flf. Mechanical causation of cytolysis. 48. 220. influence of. Parker. Monstrosities. 7. Organisms. Munk. eflfects. 109. Moenkhaus. Osmotic phenomena in muscles. 117 flf. 19.. 211. Pouchet. Sea-water. 38. 181. osmotic phenomena in. 18 ff. Overton.. Roux. harmonious character of. 226. 188. Membrane of fertilization. Protective solution. Mechanical destruction of cortical layer. Memory. Pettenkofer. 57. coordinated movements 70. 20. 99. 139 ff. Polygordius. on Schopenhauer. Meyerhof. . 131 fl. 208. 185. 11. Pawlow. 215. physiological. 65 ff. Lysins. 214. 117. in causing egg to develop. flexuosa.. 21. 179. artificial. 222.54. Poulton. 60. 177.. 207. Mendel. Reflex. artificial production of. 172. 204 flf. Production of twins from one egg through a change in the chemical constitution of the sea-water. 177 ff. Wilhelm. causing impermeability of egg membrane.. 119. 29. 115. 226. 62. 4. 20. Wolfgang. Phylloxera. 89. fusion of normally double. 117. 5. 176 ff. 38.. 52. Maternal character of heterogeneous hybrids. Potassium cyanide. 4. cooperative action of. Montgomery. 177. 27. 30. 134. membrane formation." 5. 99. 107. 38. 156. Ringer. . 50 flf. 117. varying. 5. 14. 92. 21. 7. poisonous salt action." 143. 49. 179 flf. 17. 24. Salamandra maculosa. 57. Loeb. 13. 25. 66. Norman. membrane-forming substances. Purposeful character of reflexes. Lyon. Salts. 147. 79. 132. Organs. Morphology. 160 flf. Poisoning. light. Plasmolysis.. Poisonous action of distilled water on Gammarus. 99. 16. Mendel's laws. antagonism of acids and. ferments of..Index Living of. Natural selection. 182 ff. A. 173. S. 58. 178. 57. 208. Physiological morphology. Procter. Mechanics of growth in animals. Pfluger. ISIuscles. 12. Mach. 16. 47. compulsory. eflfect of retarded. 173. 4. 55. 164. 172.. Nemec.. 189 flf. associative. 73. Parthenogenetic development. 4. artificial production Localization experiments. 114. 49. Polynoe. salt.. 199 flf. 4. Rubner. Minkowski. 148 flf. 225. Menidia. Salt . 41. Peter. 179. Photosensitive surfaces. 170 flf. McClung. Rayleigh. 145. bases as. 210. Ranke. Macromysis 27. P. 198. 58. Robertson. emulsion theory of. Luther. 19. Oxidations. in. in their relation to life and death. 35. 208. 80. Margelis. 8. Parthenogenesis. 15. 39. 144. Leo. Photosensitiveness. 11. ff. Pyrrhocoris. 39. Prevost. 49. Miescher. in animals. 209. 7. Nereis. Schmiedeberg. 116 flf. 7. 39. 66. Lucas. Salamandra atra. 49. S.. Lottia. Osterhout. T. 156 flf. Reflexes. 216. 213. 116 Segregation. "Riddle of life. in sea-urchins. 191. 208. Otoliths of ear and orientation to center of gravity of the earth. Schultze. 39. Reinke. 23 flf. antagonism of three salts. 39. 216. 45. 157. 178. varying susceptibility of eggs to. 16.. 39. 231 matter. 17. 181.

protective. 204 fertilization. 208.. 12 ff. Stevens. 4. 215. 50 16. mals. Vision. of intensity of light Tyrosin. Wasteneys. 4. Water. Standfuss. Snyder.. of." 104. D. Symmetry.. 201. 211. 55. Spermatozoon. Weismann. 157. . 132 ff. from egg. 139 ff. Wilson. 19. 91 ff. 189. 140. 144. Tunicates.232 The Mechanistic Conception of Life by ani- Selection. 208. Sipunculides. 23. "Speziflsche Bildungsstoffe. 222. " Unterschiedsempfindlichkeit." 54. 19. 161 ff. "Will. 172. Virchow. 17. Zimtz. 90.. Warburg. Temperature 135ff. CD. 13. 60. Tower. Strongylocentrotus franciscanus. 39. 182. 177 fif. Spontaneous raovements. 113 ff. Voit. 220. Waller. 144. . 96. natural. 97. 161 fl. 36 69. 17. Twofold action of spermatozoon in 12 ff. Strongylocentrotus purpuratus. Sumner.. Traube. ff." 35 ff. 157. 92. 197. 43. 188. 80. production of. role of." 24. 210. 45. Twins. ff. 51. 205. 127. X-chromosomes.. Volvox. 8. 40. 160. susceptibility of eggs of different species to parthenogenetic development. as a cytolytic agent.. 141. fif. 189. Solution. Varying Stockard.. Spermatozoa. 70. Vanessa antiope. 211. 71. twofold action in fertilization.. Tubularia mesemhryanthemum. 188. 170 ff. Uexkuell. 11. 54. 95. poisonous action of distilled water on. Spyrogyra. 38. 138ff. 162. 128.. 211. Shearer. 91. 26. Tropisms. Wiener. 92. 207. Sex determination. 198. Twin formation. 13. 190.. Vanessa levana. 99. Van Duyne. 18 ff. Stereotropism. 132 ff. 79 ff.. Vanessa prorsa. 62. 227. in segmentation. chemical. Stieghtz. 52flf. 18. distilled. 39. 70. 223. "Zielstrebigkeit. A. 205. 223. 211. 206.. Vaucheria. 181. Spallanzani. 113. 72. Varying photosensitiveness in animals. van't Hoff.

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