HOME :: CHAPTER 4 :: 4.4 THE LILLIE-LOEB DISPUTE OVER SPERM-EGG BINDING :: THE INVENTION OF ARTIFICIAL PARTHENOGENESIS
The Invention of Artificial Parthenogenesis
Chapter 5 in Controlling Life: Jacques Loeb and the Engineering Ideal in Biology, Oxford University Press, NY. Pp. 93-117.
Philip J. Pauly, 1987
This chapter is reproduced here thanks to the generosity of the author and of the Oxford University Press.
From the late 1880s, Jacques Loeb had been living out his own scientific epic. The Urwald of Chicago was the setting for scientific creation. His 1896 cornerstonelaying speech was a declaration of the path he proposed to take. In taking up physical chemistry, the direction of this path became clearer; but he only believed that he had found his own way in 1899 when he developed what he called "artificial parthenogenesis." The personal importance of this achievement was evident in his announcement of his result to Ernst Mach, in what would be his last confessional letter, that "it is in the end still possible that I find my dream realized, to see a constructive or engineering biology in place of a biology that is merely analytical." Artificial parthenogenesis brought Loeb scientific fame, and even popular notoriety, as a modern Faust. He used that heroic stature to continue his quest in what he considered the true Eden of California. Loeb worked alone on the shores of the Pacific for seven years, returning east periodically to proclaim the new scientific era.(1)
Artificial parthenogenesis was an invention: "the artificial production of normal larvae (plutei) from the unfertilized eggs of the sea urchin."(2) Loeb found that by treating sea urchin eggs with appropriate inorganic salt solutions he could initiate embryological development, a process which up to that time had required the sperm of the male urchin. Physical chemistry could be a tool for altering the basic process of reproduction.
The nature of the control over life in artificial parthenogenesis was different from that exhibited by Loeb's earlier work on heteromorphosis. In one sense it was less dramatic, since it led to no visibly new forms, but only to that familiar embryological object, the sea urchin pluteus. But that was no disadvantage. Heteromorphosis could be downgraded as merely the creation of monsters, an imitation of nature's mistakes with no real biological significance. Such an objection could not so easily be made against artificial parthenogenesis. It was a change in the mode of production (or reproduction), an improvement upon nature in that it eliminated an unnecessary element (the sperm), and offered the possibility of replacing half the sea urchin population with a mixture of salts.
The invention of artificial parthenogenesis represented an attack on the privileged status of natural modes of reproduction. A race of fatherless sea urchins was a nonnatural product completely outside the orderly structure and gradual evolution of the animal economy. Nature contained only a very restricted set of modes of reproduction; an engineering biology could lead to expansion of possibilities far beyond those that might appear through the process of evolution, or through a biology whose framework was bounded by evolutionism. For instance, artificial parthenogenesis proved that it was possible to separate the sperm's function as initiator of development from its role as carrier of the male hereditary characters.(3) Loeb believed that it would be possible to manipulate each independently, and perhaps create hitherto unknown sets of hybrids by superimposing artificial parthenogenesis on eggs fertilized by foreign sperm that were normally unable to initiate development of those eggs.(4)
Sea urchins were only the beginning. Loeb concluded the short announcement of his discovery by asking "whether we may expect to produce artificial parthenogenesis in mammalians.... I consider it possible that only the ions of the blood prevent the parthenogenetic origins of embryos in mammalians, and I think it further not impossible that a transitory change in the ions of the blood may also allow complete parthenogenesis in mammalians."(5) The possibility of human parthenogenesis, the sort of basic social reformation to which science could provide the key, was evident from the start. Artificial parthenogenesis was a vindication of Loeb's hopes and a model for science to come in which biologists would consciously work to reconstruct the natural order to make it more rational, efficient, and responsive to the ongoing development of engineering science.
Background: Loeb and Thomas Hunt Morgan on Sea Urchin Development
The history of Loeb's development of artificial parthenogenesis illustrates the nature and significance of the relationship between his general intellectual program and the details of his scientific practice. Artificial parthenogenesis was not a difficult technical problem requiring great experimental sophistication. Loeb's successful experiments were in fact performed with improperly labeled salt solutions, and many quite different techniques were developed in the following years.(6) Rather, the crucial element was the idea that artificial parthenogenesis was a biological problem at all. The perception that a biologist should try to produce fatherless sea urchins was the difficult thing; the actual production was relatively simple. Loeb discovered artificial parthenogenesis because he was seeking to control life on its most basic level; it was a natural consequence of his conviction that biology was and should be an engineering science concerned with transforming the natural order.
In later years-as artificial parthenogenesis was incorporated into class lectures and textbooks-the context of its discovery was obscured. As in many other cases, for pedagogical purposes the history of the discovery was written as the history of techniques; in this framework, Loeb's achievement seemed to be the straightforward culmination of many efforts, especially those of T. H. Morgan. As a result, some of Morgan's students, as well as those of Loeb's critics at the University of Chicago, came to believe that Loeb had illegitimately "reaped the credit for Morgan's discovery."(7) Contrasting the papers of Loeb and Morgan, however, makes clear the differences in their aims.
Loeb began experiments on embryonic development in 1892, immediately after escaping from his unsatisfying position under Morgan at Bryn Mawr. Teaching embryology had not changed his view that cytology was boring, and he sought to show that simple physiological concepts were sufficient for dealing with development and that investigations of cytological complexity were both superficial and unnecessary. Loeb began experiments on sea urchin eggs on the basis of his attempts in the preceding year at Naples to extend to animals Sachs's demonstration of the dependence of plant growth on osmotic pressure. Since growth of hydroids, like that of plants, was a function of osmotic pressure (which depended upon the concentration of salts in the medium), Loeb reasoned that it might also be possible to control the rate of development of sea urchin eggs by altering the concentration of salts in the seawater.(8) Increasing the salt concentration, which changed the osmotic equilibrium and hence the amount of water in the egg, did retard cleavage; but the more important phenomenon occurred when the eggs were taken out of the salt solution and returned to normal seawater. They did not merely begin to segment again at the usual rate, but divided all at once into many cells, nearly catching up to the stage of development of the control eggs left in normal seawater. E. G. Conklin, who had recently graduated from Johns Hopkins, examined the eggs kept in solution cytologically for Loeb, and found many "distinct nuclei."
It appeared to Loeb that while nuclear cleavage processes had continued in the salt solution, the removal of water from the cytoplasm lowered its irritability and hence its ability to respond to the stimuli coming from the nucleus. When irritability returned with the return to seawater, the cytoplasm responded to the stimuli from many nuclei and segmented immediately into as many cells as there were nuclei. Hence cell division was a process centered in the nucleus: "The segmentation of the protoplasm of the egg, and probably in every cell, is only the effect of a stimulus exercised as a rule by the nuclei." Control of cell division was the result of control of nuclear division; cytologists' attempts to explain cell division through the complex mechanics of intracellular structures were probably a waste of time. The physiological concept of stimulus and response provided all the necessary complexity.
In a paper that appeared two years later, T. H. Morgan attacked this attempt to downgrade the importance of cytology. While agreeing that immersion in concentrated salt solutions retarded cleavage and that rapid segmentation occurred on return to seawater, Morgan denied that the nuclei continued to divide during the period in the salt solution. Cytological examination showed that "eggs put in the salt-solution come to rest both for the protoplasm and the nucleus," and remained that way "throughout all the time of immersion." Only when the eggs were replaced in seawater would the nucleus undergo "a rapid and irregular division (fragmentation) and the nuclear pieces migrate to the periphery of the egg." Detailed investigation had demonstrated, Morgan believed, that Loeb's physiological speculation was simply incorrect.(9)
In the summer of 1895 Loeb repeated his old experiments and those of Morgan with the assistance of W. W. Norman, his friend from Naples-now professor at the University of Texas and Loeb's pro forma Ph.D. student. They found that the nature of the salt made some difference in the result, so they substituted magnesium chloride for sodium chloride; Norman sectioned the eggs and figured the mitotic divisions of the nuclei of eggs kept in the salt solution-confirming Loeb's prediction. Loeb argued that Morgan's inability to repeat Loeb's experiment had been due to use of contaminated solutions.(10)
Morgan soon conceded that "in my experiments on the eggs of Arbacia I did not use chemically pure sodium chloride," and that "the nucleus may sometimes divide" while in the salt solution, as Loeb had originally claimed. While admitting the insufficiencies in his chemical techniques, he still maintained that his cytological examinations showed that the phenomena in eggs in salt solutions were not as simple as Loeb had described. He argued now that the salt stimulated the egg to produce certain abnormal structures, most particularly a number of "artificial astrospheres," which appeared even in unfertilized eggs kept in salt solutions; on return to sea water these structures faded away and did not take part in the rapid cell division.(11)
The fact that he had obtained artificial astrospheres in both fertilized and unfertilized eggs led Morgan to study the effects of salt solutions in more detail. His contrasts between normal development and the phenomena in salt-soaked eggs were expanded in a long paper that he published in June 1899. There he showed that although nuclei of fertilized eggs did divide to some extent while in a salt solution, the division was "so extremely abnormal, irregular, and imperfect as to preclude any comparison between the stages that take place in such eggs and in those in normal eggs."(12) Morgan argued that Loeb was wrong in his claim that the salt merely decreased the "irritability" of the protoplasm, and deluded in thinking that it was possible to pin down exactly the effects of particular agents on well-defined aspects of embryonic development. It was impossible to predict the nature of the changes from the nature of the external "stimulus." The major causes of embryonic phenomena were internal, and could only be understood through detailed cytological study of the "vital structure" of the egg.(13) It was necessary to analyze the actions of centrosomes, astrospheres, chromosomes, spindles, and so on under many different circumstances in order to gain insight into the nature of development; this was what Morgan did in this paper, for almost one hundred pages.
One of the examples of cellular complexity was the creation of astrospheres by both fertilized and unfertilized eggs in response to stimulation by the salt solution. A second was the fact that on return to seawater not only fertilized but unfertilized eggs began to divide. Morgan described the cytological transformations in these eggs, in which single nuclei sometimes contained "hundreds of chromosomes," searching for information regarding the functions of centrosomes, chromosomes, and artificial astrospheres.(14)
Such transformations were not unexpected. In 1895 the German zoologist Richard Hertwig discussed problems similar to those that interested Morgan, namely, the origins of the centrosome and its role in cell division. He pointed out that old unfertilized sea urchin eggs sometimes would begin a nuclear division as they degenerated, and that this process could also be initiated by "stimulating" the eggs with strychnine (poisoning, therefore, being the result of overstimulation).(15) Morgan recognized the similarity of his results to those of Hertwig, and shared the German's opinion regarding their pathological nature. Although he obtained more cell divisions than Hertwig, he saw little connection between these phenomena and the actual development of embryos. He pointed out that "these pieces never acquire cilia and do not produce any form that resembles any stage of the normal embryo." Rather, there were similarities to the growth of tumors, another pathological phenomenon in which the key aspect of embryological development-the development of form-was missing.
Hertwig's and Morgan's experiments producing "pathological" division of unfertilized stimulated eggs were analytical efforts wholly analogous to Roux's killing of one of the two frog blastomeres. The difference was in the insertion of a different sort of "bomb" in a different part of the organic factory. For all these men the causes of cell division and differentiation were internal to the egg. Yet influenced by Hans Driesch's critique of Roux's appeals to mechanical causation, Morgan was very conservative in his conclusions in 1899 regarding the nature of these determinants. The preceding year he had argued that phenomena of regeneration were better explained in terms of "intelligence" than in terms of chemistry and physics, and in this paper he again alluded to the problem of the limits of mechanistic explanation:
It is the vital structure of the egg on which the result largely depends, for if one kind of stimulus is as capable as another of starting the development of the egg, then we have accomplished very little in the way of explanation if we have only determined what these stimuli may be. I trust that I shall not be misunderstood as to the way in which I understand the doctrine of vitality as applied to living things. I have in no sense denied that the vital action is in reality a complicated series of unknown chemical and physical changes, but, also, I have not affirmed that it is so.(16)
Morgan considered it at least possible that "vital action" depended upon Driesch's indeterministic entelechy.
The Production of Parthenogenetic Sea Urchins
Loeb began his experiments in the summer Morgan's long paper appeared. Both this paper, and his own recent work on ionic inducement of rhythmic contractions in skeletal muscle and ionic balance in the survival of Fundulus embryos, led him to believe (as he later said) that "by changing the ions contained in a tissue we can impart to it qualities which it does not ordinarily possess."(17) From the start, apparently, he was searching deliberately for the means to produce parthenogenetic sea urchins; the problem was to determine the appropriate mixture of ions. Working with solutions of sodium chloride, potassium chloride, calcium chloride, and magnesium chloride, Loeb first decided that since the magnesium salt was the least toxic it was the best agent for artificial parthenogenesis.(18) Such reasoning was diametrically opposed to that of Hertwig and Morgan, who saw such pathological cell division as the result of poisoning-of "overstimulation" analogous to the effects of large doses of alcohol and morphine on humans. Loeb then experimented to find the proportions of salt and seawater and the duration of soaking that would maximize development. Despite the fact that his eggs never formed the usual prominent "fertilization membrane" and as a result often broke up into fragments, Loeb remained unconcerned that his productions might be mere pathological curiosities or monsters; he claimed in the title of the article that these were "normal" larvae. He drew no sharp line between pathological and normal processes, examining the pathological in order to illuminate the normal, as did Morgan; what mattered to him was the end result.
Loeb rapidly published the fact of his discovery, along with his speculations concerning the possibilities of producing mammals parthenogenetically, in October 1899; in the following spring he published the detailed account of his experiments. They were presented in chronological order as a model for scientific problem solving, and as a dramatic struggle to find the solution before the supply of spawning sea urchins ran out for the year. He described how he systematically varied the proportion of magnesium chloride in the solution, then the time of immersion, and finally the absolute number of other ions, searching for the combination that best promoted cell division. Further adjustment of the proportions of magnesium chloride and seawater enabled development finally to go beyond the blastula to the pluteus stage, which was as far as larvae could be raised in the laboratory. He then assured himself that these processes were not due to inadvertent fertilization by spermatozoa; this was evident from the lack of a fertilization membrane in the parthenogenetic eggs, the lack of development of control eggs kept in seawater, and the toxicity of magnesium chloride solutions for sperm.
Artificial parthenogenesis demonstrated that the fertilizing properties of the sperm were separate from the transmission of hereditary qualities and that it was necessary to "transfer the problem of fertilization from the realm of morphology into that of physical chemistry." Ions, changes in temperature, and enzymes or other as yet unknown materials in the sperm could all initiate developmental processes. Perhaps this was due to the fact that all these agents could alter the physical state of proteins.(19)
While presented as a model of scientific problem solving, Loeb's development of artificial parthenogenesis was in fact not an example he would have liked emulated. Osmotic pressure, relative proportions of the ions, and the gross amount of magnesium were all being simultaneously changed in his successful treatments. In his initial report he argued that, by analogy with his experiments on Fundulus, the key factor was the change in the proportions of the ions that resulted from the addition of magnesium chloride. In the more detailed paper he attributed it to increase in osmotic pressure and a specific effect of magnesium, since the other solutions did not work.(20)
Error in interpretation was not the biggest problem, however. After publishing the preliminary paper and writing up the full report, Loeb was anxious to continue this important research. European scientists could work through the winter at Naples, but he would have to wait until the next summer to find more sea urchin eggs at Woods Hole. One marine station where urchins spawned in the winter was that of Stanford University; in the winter of 1900 Loeb traveled cross-country to Pacific Grove, California, in order to continue his work. (21) For weeks he tried to repeat his previous experiments with a different genus of urchin, but without success. Worry mounted rapidly. Zoologists such as Morgan and Whitman had long criticized his lack of expertise in the hard details of cytology; he had now made radical claims for control of reproduction. If they were spurious results—blobs of protoplasm interpreted as blastulae—he would look ridiculous. He checked and double-checked his work, and finally found the answer: an assistant had made the standard salt solutions he had used at Woods Hole to the wrong concentrations. Loeb had been using mixtures quite different from what he thought—and reported—he had. (22)
Now, with proper mixtures, he was able to get larvae by putting eggs for a period of 30 to 120 minutes in a variety of hypertonic solutions (solutions with an osmotic pressure higher than that of sea water), including solutions of sugar and urea. Artificial parthenogenesis was not as complicated as the survival of Fundulus; it was merely another example of the importance of osmotic pressure in life phenomena.While inserting a few remarks about the effect a hypertonic solution might have on the egg "for those who enjoy the speculative side of biology," he now felt that it was best, after being wrong twice, "to supply the lacking experimental data in this field of biology before we begin to theorize." (23)
Some important conclusions can be drawn from the fact that Loeb was able to succeed at artificial parthenogenesis in spite of his experimental errors. The ease of his success demonstrates that the development of experimental techniques was much less significant than the perception that artificial parthenogenesis was worthwhile. Loeb's experiments were quite similar to Morgan's; the difference was in their understanding of what a biologist should do. Loeb's activity indicates clearly what his main motivation was. Theories of the processes underlying artificial parthenogenesis were primarily of rhetorical significance; they were introduced and discarded in rapid succession. It mattered little that his technique was sloppy and that the data were often inconsistent. Loeb was guided by the desire to produce fatherless sea urchins, not to reduce fertilization to a particular chemical process. Analytical difficulties were subordinated to the search for control of development. Loeb invented artificial parthenogenesis because of his search for an engineering biology, and artificial parthenogenesis served as an exemplar for the further development of that approach.
Handling the News
In late 1899, a number of newspapers reported Loeb's first announcement of artificial parthenogenesis (figure 10). The headline in the Boston Herald was typical: "Creation of Life. Startling Discovery of Prof. Loeb. Lower Animals Produced by Chemical Means. Process May Apply to Human Species. Immaculate Conception Explained. Wonderful Experiments Conducted at Woods Hole." Loeb rapidly moved from being a professor known only in narrow academic circles to a major scientist and eminent, even notorious, public figure. By 1902 his actions were front-page news, and he was the subject of a profile in the mass-circulation McClure's Magazine.
|Figure 1 Early newspaper account of artificial parthenogenesis.|
Artificial parthenogenesis entered popular consciousness with the new century. It was the first major manipulation of the reproductive process to reach the public; and because Loeb worked on such "lowly" organisms as sea urchins, it was a form of sex that was open to newspaper discussion. Yet there was considerable interest in its human implications. Journalists speculated on the possibilities of human parthenogenesis, and they solicited comments from a wide range of American biologists. Artificial parthenogenesis in many respects set the images and the tone for the discussion of reproductive technology in the public mind for a long time to come; Loeb, on the cover of Harper's Weekly as one of a series of "Americans of To-Morrow," became the epitome of a new kind of biologist. The content of these reactions is thus important both as an index of sentiment at the time and as an influence on future perceptions.(24)
The initial scientific reaction to Loeb's announcement was doubt. According to Loeb, Morgan-far from claiming credit for anticipating him-announced publicly that the work "must be a mistake."(25) It was questionable whether Loeb had in fact produced blastulae; and if he did, it was quite possible that the eggs had merely been fertilized by stray sperm. Debates were kept alive through the winter of 1900 by Loeb's changing claims and by his problems in replicating his initial result.
That situation changed rapidly by the summer of 1900. Respected European scientists such as Curt Herbst, working at the Naples Zoological Station, confirmed Loeb's claim to have initiated basic developmental processes, and excluded the possibility of inadvertent fertilization by using boiled seawater. Loeb published the more elaborate descriptions of his experiments. E. B. Wilson worked out the cytology of the process. Loeb extended his work to worms, and others succeeded that summer with starfish. By 1910 the French biologist Eugene Bataillon induced artificial parthenogenesis in frogs by piercing eggs with a needle. Morgan's Experimental Embryology, though not inclusive, listed an average of twelve papers on artificial parthenogenesis published each year between 1900 and 1910; at Woods Hole the popularity of such studies grew to the point that when the young evolutionist Francis B. Sumner first came there in 1903 he perceived it as a "cult."(26)
E. G. Conklin (now at the University of Pennsylvania), in announcing Herbst's private letter confirming Loeb's work, considered artificial parthenogenesis "one of the greatest discoveries in biology."(27) In a popular article a few months later, Wilson, reviewing recent work in experimental embryology, considered Loeb's discovery "foremost in interest." It opened up a variety of manipulative possibilities—of controlling sex and "creating wholly new organic forms by varying slightly the conditions of development."(28) Some biologists were less enthusiastic about such claims, arguing privately and in the newspapers that parthenogenesis was a common property of invertebrates, but at least initially this skepticism was a minor theme.(29) In 1901 Loeb was a finalist for the first Nobel Prize in Physiology or Medicine.(30)
Artificial parthenogenesis was the first product of Woods Hole to become major news. Loeb stimulated discussion with hints about mammalian parthenogenesis in his first paper, and with other comments that were picked up by a reporter:
The development of the unfertilized egg, that is an assured fact. I believe an immaculate conception may be a natural result of unusual but natural causes. The less a scientist says about that now the better. It is a wonderful subject, and in many ways an awful one. That the human species may be made artificially to reproduce itself by the withdrawal of chemical restraint by other than natural means is a matter we do not like to contemplate. But we have drawn a great step nearer to the chemical theory of life and may already see ahead of us the day when a scientist, experimenting with chemicals in a test tube, may see them unite and form a substance which shall live and move and reproduce itself.(31)
While Loeb soon became more reticent, other biologists elaborated on these themes. Whitman's protege F. R. Lillie, recently returned to a faculty position at Chicago, explored the application of parthenogenesis to humans in some detail. He believed it possible in principle, and agreed that "there is no doubt that the investigators in morphological anatomy will continue their experiments with such discoveries in view." The problem, as Lillie saw it, was to determine the appropriate conditions. "When asked how long it probably would be before the feat of reproducing vertebrates artificially is accomplished, Dr. Lillie asked in turn: 'When do you think the north pole will be discovered?' and added: 'The solution of the problem seems extremely remote, indeed."' But it would be a problem for future generations.(32)
There were a variety of themes expressed, and it took some time for a stable popular image of Loeb's work to develop. Biologists-Loeb among them-were uncomfortable with the imagined details of human parthenogenesis, and expressed this with smutty humor. They talked about "maiden ladies" giving up sea bathing on hearing of Loeb's discovery, and joked that vacations at the seashore did often result in conception. Loeb supposedly responded to the criticism that he had not excluded all sperm in his first experiments by earnestly assuring the "gentlemen" that he had filtered and sterilized the seawater, and "in wiping out the dishes used only ladies' handkerchiefs."(33) The potential superfluousness of the male was a second theme. Women were said to have written Loeb asking him to "give them children." The French biologist Yves Delage reported receiving letters "signed with women's names, congratulating him in dithyrambic terms for having finally freed the woman from the shameful bondage of needing a man to become a mother."(34)
One of the more bizarre interpretations, deriving in part at least from Loeb's use of the term "immaculate conception," centered on the relation between artificial parthenogenesis and the virgin birth of Jesus. One religious paper noted that "it is interesting to know that the science of this recent day is entertaining as a possibility what Christian faith accepts as actual in the case of the most notable birth that ever occurred."(35) In the other direction were speculations that it would soon be possible "to raise domestic animals and children born without help of a male through an operation which would be regulated scientifically and almost commercially, similar to raising the fry of trout."(36)
By 1902, as coverage shifted from the daily papers to magazines, interpretations stabilized. The person who crystallized an image of Loeb, his work, and his aims, and presented it to a wide public was Carl Snyder. This young journalist, who would later become a major figure in applied economics, interviewed Loeb for McClure's Magazine, the popular progressive monthly edited by Lincoln Steffens. Snyder's article gained even wider circulation through being reprinted in the English Fortnightly Review and collected with Snyder's other articles on cosmology, physics, and psychic research in a book that appeared in English, German, and Italian.(37)
Snyder visited Loeb in his Chicago laboratory, pictured with a rolltop desk and simple equipment of reagents and finger bowls. He likened Loeb to a "busy and businesslike surgeon," remarkable on the one hand for his "clear-cut features and rather piercing eyes," and on the other because he was "so cautious in his statements, so candid as to precisely what he has achieved." Snyder described Loeb's work of the preceding decade on such subjects as tropisms, heteromorphosis, and salt action, and he quoted extensively from Loeb's papers. He considered artificial parthenogenesis, in some ways "the most vital discovery in the history of physiology," a near approach to "the manufacture of life in the laboratory."
In his discussion with Snyder, Loeb was "candid" enough to talk of his "many periods of profound discouragement," and he retailed his complaints about the domination of American society and American science by "politicians." He argued that the forces controlling biological phenomena were the same as those active in the inanimate world, and complained that hardly one step had been made in understanding electrophysiology from the time of Galvani to the present. He was perhaps too candid. He told Snyder that his purpose was "to go to the bottom of things. I wanted to take life in my hands and play with it." Snyder described him reaching out, "as if to catch this elusive phantom," as he spoke. "I wanted to handle it in my laboratory as I would any other chemical reaction-to start it, stop it, vary it, study it under every condition, to direct it at my will!"(38)
"Playing with life" was the image that struck Snyder, and that he used as a recurring motif in his article. He connected Loeb's "dream" to the activities of Prometheus and Faust, two notable legendary technologists. He did not mention the other analogous figure, Frankenstein, but other journalists soon introduced that comparison. While most of the more serious journalists pulled back from the idea that Loeb was "manufacturing" life through artificial parthenogenesis, the image of Loeb as Faust or Frankenstein continued through the rest of the decade (figure 11).(39)
Given Loeb's work, his claims, and their implications, it is not surprising that he was equated with these two other well-known German scientific "creators." What is remarkable, however, is that neither the tragedies nor the moral messages of these dramas were discussed. Scientists commenting on Loeb's work restricted themselves to discussing the feasibility of artificial parthenogenesis in different species, and apart from one vocal Yale instructor, R. W. Hall, who expressed concerns about eventual degeneration of a line of parthenogenetic humans, they said nothing about its desirability.(40) Snyder reported Loeb's materialism and argued that his work "seemed to topple the whole structure of our ideas of life." But apart from calling Loeb an iconoclast, Snyder presented a straightforward, mildly enthusiastic account. A San Francisco reporter, apparently the first to compare him to Frankenstein, was well aware of the moral of Mary Shelley's book, but argued blithely that Frankenstein's creation was a monster only because Shelley "premised the necessity of a soul where she should have predicated the fermentation of the enzyme."(41)
A range of speculations is possible regarding this lack of concern about the problematic implications of reproductive technology. The possibility of realization was remote, conventions of science "reporting" militated against serious criticism, and Americans in the Progressive Era were notoriously "innocent" in their enthusiasm for science and technology.(42) The crucial point, however, is that the problems were by no means self-evident; more particularly, at this time, reproductive manipulation was not differentiated from any other technological possibilities in its moral status.
Loeb was not opposed to articles such as Snyder's describing his work. He kept up acquaintance with Snyder for many years, and later in the decade junior colleagues and proteges published articles such as "The Search for the Origin of Life" and "Dynamics of Life; the Achievements of the Scientific Frankenstein."(43) But on a more mundane level, Loeb's sudden emergence as a celebrity was painful. He was unable to deal with the "lack of etiquette" of newspaper journalists who reported lectures prior to scientific publication as bulletins from the laboratory and begged him for scoops. Some borrowed students' notebooks and eavesdropped on his classroom, reporting his "sharp reprimand to his quiz class, which, he asserted, was given to too much frivolity."(44) Just as disturbing were the gross exaggerations in a wide variety of newspapers. Loeb was not glad to read that he had demonstrated the possibility of the "immaculate conception" of Jesus, nor that artificial parthenogenesis was equivalent to spontaneous generation.
At the December 1901 meeting of the American Society of Naturalists, reporters turned two of Loeb's papers-one on prolonging the life of starfish eggs through chemical treatment, the other an argument for the role of ions in nerve conduction-into major news items. Headlines such as "Secret of Endless Life" and "Science Claims It Can Stay Death" trumpeted Loeb's experiment as the first step toward bringing about human immortality. Matters grew completely out of control as the yellow papers began to search through Loeb's old work and began to publish such articles as a description of some of Loeb's old heteromorphosis experiments, with a drawing of a cow whose head and forelegs were transposed; the caption read, "What would happen if Professor Loeb were to subject a cow to the treatment which his most recent discovery is said to have made possible."(45)
The yellow papers, in most cases elaborating on generalities from a few firsthand stories, were incorporating Loeb into the most extreme elements of the American tradition of medico-scientific puffery. The "Endless Life" story generated articles joking about Jonathan Swift's Struldbrugs, perpetual motion machines, and whether death was a germ or a German.(46) Reports that Loeb had claimed that electricity and not heat was the source of vital energy (based on a paper that argued that salts should be viewed not as food, but as necessary to maintain the body's ionic balance) led to a flood of letters claiming priority and use of Loeb's name in testimonials; a fullpage advertisement in the San Francisco Examiner had the extended headline, "Dr. McLaughlin Wants Rockefeller's Million. The Man who proved that he was five years ahead of Professor Loeb in discovering that Electricity is Life now says that he can cure John D. Rockefeller, and agrees to give $5000 to charity if he fails."(47)
While the more established publications recognized that these claims were exaggerated, and one paper even gave Loeb the backhanded compliment that it did him an injustice to assume that his discoveries were "epochal or even very important," he was in some circles on the verge of being considered a quack. Foreign journalists considered his reputed claims about eternal life typical American "humbug."(48) Some scientists, generally not among the disciplinary leaders, criticized what they supposed were his ideas and interest in cheap publicity. An editorialist for the Journal of the American Medical Association, for example, cited the newspaper reports on artificial parthenogenesis and the lecture by "A. B. Conklin," but not Loeb's own papers; he dismissed Loeb's work as "a crude idea apparently based on misconception of the facts." Ludwig Hektoen, editor of the journal and Loeb's junior colleague at Chicago, apologized for the article, explaining that the editorial resulted from "strong feeling among good physicians against newspaper notoriety."(49)
Loeb was embarrassed by the claims attributed to him and by the implication that he was bypassing scientific channels of communication. What most distressed him, however, was the public ridicule, in part by scientists, to which he was unable to respond. He was sensitive to his position as a foreigner, a Jew, and an atheist. The treatment he had received at the hands of Hitzig, resurfacing at this time with Hitzig's publication of a fifteen-page polemic against the threepage summary Loeb gave of his 1886 results in his book on neurophysiology, was never far from Loeb's consciousness.(50) And his sense of personal insecurity, present from childhood as a Jew in a Catholic town, surfaced in such circumstances. This combination of associations is important for explaining the violence of Loeb's reaction in January 1902 to what Robert Kohler called an "innocent peccadillo" on the part of his junior colleague Albert P. Mathews.(51)
Mathews had graduated from Columbia in 1898 with a Ph.D. in physiology and zoology. He worked with Loeb at Woods Hole in 1899, and the following year published on artificial parthenogenesis, attributing the discovery ambiguously to both Loeb and Morgan. Loeb was impressed with Mathews and in 1901 brought him from Harvard to Chicago as a physiological chemist, but tension arose almost immediately as Mathews began to follow too closely in Loeb's footsteps. At the 1901 Naturalists' meeting Mathews presented a paper on the colloidal basis of nerve stimulation that, according to Loeb, made too extensive use of the work of Loeb and his students. At first, while complaining to President Harper, Loeb grudgingly excused what he considered Mathews's inappropriate conduct. His attitude shifted in the next two weeks, however, as reporters and healers learned about his own papers at the meeting and began to badger him about his supposed discovery of endless life and the electrical basis of vital energy. He began to suffer what he thought were heart palpitations, and in this agitated state became increasingly upset at the apparently unrelated overaggressiveness of Mathews. On January 22, 1902, he pressured Harper to set up a committee to investigate what he considered Mathews's unsound work and unbalanced personality- but he still claimed to want peace in the department.(52)
Three days later, however, Loeb blew up completely. He learned that Mathews, in addition to appropriating his ideas, was also responsible for his other problems. Mathews had initiated the newspaper publicity by sending reports of both his and Loeb's papers to his brother, a reporter for the New York Sun. Loeb's usually tight, crabbed handwriting became increasingly dark, large, and disjointed as he detailed his objections to Harper, concluding that Mathews's action was "more than I can be expected to endure. It is impossible for me to enter into a controversy with Daily papers or Magazines."(53)
As the senior physiology professor, Loeb could have pushed Mathews quietly aside. He was less interested in Mathews's position, however, than in his own vindication from blame. He requested an investigation by the university senate, which concluded that "the conduct of Dr. Mathews in publishing premature and immature statements is reprehensible," and which set up guidelines for future intra-departmental relations.(54)
One indication of the overwhelmingly positive response to Loeb's development of artificial parthenogenesis was the change it made in his professorial status. He had been an associate professor at Chicago since 1894, and some years later considered himself near the bottom of the promotion list and was exploring other positions. In February 1900, however, he was promoted to full professor, and a year later to "head professor," with membership in the university senate. Whatever their reservations, the Chicago administration began to recognize that Loeb was a "rare bird"-someone involved with important research.(55)
In spite of these changes, Loeb's moods fluctuated, with periods of dissatisfaction and depression. He felt himself intellectually isolated in physiology; when Frank Mall asked about nominating him for the Nobel Prize, he replied that he could not win, since he belonged to no established physiological "Schule"-"and this is almost as bad as to be without ancestors in China." The improvement in his status at Chicago meant increased administrative responsibilities and the trauma of dealing with situations like that of Mathews. He was now also involved in providing routine physiological instruction to the students of Rush Medical College, which had been cobbled together with the University of Chicago without adequate financial support. He complained about "staleness"; almost all his summers had been spent in Woods Hole, where he usually rushed to complete as much work as possible in the three months available, and where arguments over control and financing had produced a politically charged atmosphere.(56) Loeb thought increasingly about the uninterrupted periods for work on marine invertebrates he had enjoyed at Stanford's small Hopkins Marine Station in Pacific Grove, California, in the winters of 1898 and 1900. He confessed to Mall that he longed to live in California in order to develop the possibilities of artificial parthenogenesis full-time; he wanted to work on marine organisms because, he joked, they were the only animals, besides man, "whose life is entirely absorbed in assimilation and reproduction."(57)
Literature Cited, Including Footnotes
1. JL to Ernst Mach, 28 December 1899, EM; on the myth of the hero in late nineteenth-century German science see Frank J. Sulloway, Freud, Biologist of the Mind (New York: Basic Books, 1979), pp. 445-495.
2. JL, "On the Nature of the Process of Fertilization and the Artificial Production of Normal Larvae (Plutei) from the Unfertilized Eggs of the Sea Urchin," American Journal of Physiology 3 (1899):135-138; reprinted in JL, Studies in General Physiology (Chicago: University of Chicago Press, 1905), pp. 539-543.
3. JL, "On the Artificial Production of Normal Larvae from the Unfertilized Eggs of the Sea Urchin (Arbacia)," American Journal of Physiology 3 (1900):434-471; reprinted in Studies, pp. 575-623, esp. pp. 620-623.
4. JL, "Experiments on Artificial Parthenogenesis in Annelids (Chaetopterus) and the Nature of the Process of Fertilization," American Journal of Physiology 4 (1901):423-459; reprinted in Studies, pp. 646-691, esp. pp. 681-682.
5. JL, "On the Nature of the Process of Fertilization," in Studies, p. 543.
6. Ethel Browne Harvey, The American Arbacia and Other Sea Urchins (Princeton: Princeton University Press, 1956), pp. 198-201, listed dozens of ways to induce artificial parthenogenesis.
7. Donald Fleming, "Introduction," JL, The Mechanistic Conception of Life (Cambridge: Harvard University Press, 1964).
8. JL, "Investigations in Physiological Morphology. III. Experiments on Cleavage," Journal of Morphology 7 (1892):253-262; reprinted as "Experiments on Cleavage," in Studies, pp. 253-264.
9. T. H. Morgan, "Experimental Studies on Echinoderm Eggs," Anatomischer Anzeiger 9 (1894):149
10. JL, "Ueber Kerntheilung ohne Zelltheilung. Briefliche Mittheilung an den Herausgeber," Archiv fur Entwicklungsmechanik der Organismen 2 (1895):298-300. W. W. Norman, "Segmentation of the Nucleus without Segmentation of the Protoplasm," ibid. 3 (1896): 106-126.
11. T. H. Morgan, "The Production of Artificial Astrospheres," ibid. 3 (1896):340.
12. T. H. Morgan, "The Action of Salt Solutions on the Unfertilized and Fertilized Eggs of Arbacia, and of Other Animals," ibid. 8 (1899):484.
13. Ibid., pp. 526-527.
14. Ibid., pp. 460-479.
15. Richard Hertwig, "Ueber die Entwicklung des unbefruchteten Seeigeleiers," Festschrift zum siebensigsten Geburtstage von Carl Gegenbauer am 21 August 1896, 2 vols. (Leipzig: W. Engelmann, 1896), 2:21-86, esp. pp. 57-63; cf. Morgan, "Action of Salt Solutions." pp. 522-524.
16. Morgan, "Action of Salt Solutions," pp. 526-577: idem, "Some Problems of Regeneration," Biological Lectures Delivered at the Marine Biological Laboratory of Wood 's Hole in the Summer Sessions of 1897 and 1898 (Boston: Ginn & Co, 1899). pp. 205-206.
17. JL, "Artificial Production of Normal Larvae," in Studies, p. 581; idem, "On Ions Which Are Capable of Calling Forth Rhythmical Contractions" (1899), in Studies, pp. 518-538; idem, "On Ion-Proteid Compounds and their Role in the Mechanics of Life Phenomena" (1900), in Studies, pp. 544-558, esp. pp. 547-548. This last paper was written after the discovery of artificial parthenogenesis, but the experiments were performed earlier.
18. JL, "Artificial Production of Normal Larvae," in Studies, pp. 581-599.
19. Ibid.. pp. 620-622.
20. JL, "On the Nature of the Process of Fertilization," in Studies, pp. 541 -542; idem, "Artificial Production of Normal Larvae," in Studies, pp. 621-622.
21. JL, "On Artificial Parthenogenesis in Sea-Urchins," Science I I (1900):612-614; reprinted in Studies, pp 624-626.
22. JL, Studies, p 639n. (dated 1903).
23. JL, "Further Experiments on Artificial Parthenogenesis and the Nature of the Process of Fertilization," American Journal of Physiology 4 (1900):178-184; reprinted in Studies, pp. 638-645; quotation p. 644.
24. Harper's Weekly, 13 December 1902, p. 1924; Howard Mumford Jones, The Age of Energy: Varieties of American Experience, 1865-1915 (New York: Viking Press, 1970), pp. 306-307.
25. JL to E.P.Lyon,2 May l917,LPbox8. 1
26. T. H. Morgan, Experimental Embryology (New York: Columbia University Press, 1927), pp. 734-749; see also E. Newton Harvey, "Methods of Artificial Parthenogenesis," Biological Bulletin 18 (1910):269-280; Francis B. Sumner, The Life History of an American Naturalist (Lancaster, Pa.: Jacques Cattell Press, 1965),pp. 170-171.
27. E. G. Conklin, "The Fertilization of the Egg and the Early Differentiation of the Embryo," University Medical Magazine 13 (I900):18-19, 63.
28. E. B. Wilson, "Aspects of Recent Biological Research," International Monthly 2 (1900):86-88.
29. "Professor Loeb's Discovery," Topeka Daily Capital, 26 November 1899; "Loeb's Theory Combatted," Boston Evening Transcript, 2 October 1900.
30. JL to F. P. Mall, 21 October 1900, CDE-FM; Claire Salomon-Bayet, "Bacteriology and the Nobel Prize Selections,1901-1920," Science, Technology, and Society in the Time of Alfred Nobel, ed. C. G. Bernhard, et al (Oxford: Pergamon Press, 1982), p. 392; he was also a finalist in 1906, 1909. and 1917. The second American to reach that stage was the bacteriologist Theobald Smith, in 1908. Salomon-Bayet discusses how bacteriology dominated prize selection prior to 1920.
31. "Creation of Life," Boston Herald, 26 November 1899; see also "Science Nears the Secret of Life," Chicago Tribune, 19 November 1899, p. 33.
32. "Reproduction of Humans," Boston Evening Transcript, 2 October 1900.
33. Fleming, "Introduction," p. xxiv; [W. J. V. Osterhout, compiler], "Anecdotes of Jacques Loeb," LP box 59.
34. Fleming, "Introduction," p. xxiv; Yves Delage and Marie Goldsmith, La Parthenogenese naturelle et expenmentale (Paris: E. Flammarion, 1913), p. 302. None of these letters, unfortunately, can be found in LP.
35. Detroit Christian Herald, 26 April 1900.
36. Delage and Goldsmith, La Parthenogenese, p 302.
37. Carl Snyder, "Bordering the Mysteries of Life and Mind," McClure's Magazine 18 (2 March 1902):386-396; reprinted in Fortnightly Review 77 (1902):1010-1023; also in idem, New Conceptions in Science (New York: Harper, 1903).
38. Ibid., p. 388.
39. Enos Brown, "The Creation of Life by Artificial Means," Scientific American 92 (1905):459-467; Anna Drzewina, "Dynamics of Life; the Achievements of the Scientific Frankenstein," Scientific American Supplement 67 (1909):156-157.
40. "Loeb's Theory Combatted," Boston Evening Transcript, 2 October 1900.
41. "Illustrious Biologist Joins Faculty of State University," San Francisco Examiner, 12 November 1902.
42. Henry F. May, The End of American Innocence (New York: Knopf, 1959), pp. 174-175, discusses Loeb briefly.
43. Martin Fischer, "Professor Jacques Loeb," University of California Chronicle 5 (1902):349-351: S. S. Maxwell. "Search for the Origin of Life," Public Opinion 38 (1905):947-950: J. B. MacCallum, "The Recent Work of Professor Loeb," Independent 59 (1905):315-319; Drzewina, "Dynamics of Life." There is substantial correspondence between Loeb and both Snyder and Drzewina in LP.
44. "America Disgusts Dr. Loeb." Chicago Inter-Ocean, 16 February 1902; "Topics in Chicago," New York Tribune 5 October 1902,11:14; author's interview with Leonard B. Loeb. 30 July 1977, Pacific Grove, California.
45. "Secret of Endless Life." Chicago Inter-Ocean, 31 December 1901; "Science Claims It Can Stay Death," New York Journal, 1 January 1902; "Can Make Feet and Head Change Places," New York Journal, clipping, n.d. Dozens of clippings of this sort are in LP boxes 57-58.
46. "But Who Wishes to Live Forever?" Detroit Free Press, 1 January 1902; Kansas City Journal, 2 January 1902; St. Louis Post-Dispatch, 3 January 1902.
47. "Anticipated Loeb's Discovery," New York Tribune, 3 January 1902; San Francisco Examiner, 15 February 1903.
48. "Nothing Wonderful," Rochester Chronicle, I January 1902; Octave Uzanne, "Le vie a perpete," Echo de Paris, 9 January 1902. Loeb was worried enough about the European response that he sent a disclaimer to Nature 63 (1901):372, that he was "in no way responsible for the journalistic idiosyncrasies of newspaper reporters."
49. "Artificial Parthenogenesis," Journal of the American Medical Association 34 (1900):1009-1010; Ludwig Hektoen to JL, 24 April 1900, LP box 6.
50. JL, Comparative Physiology of the Brain and Comparative Psychology (New York: G. P. Putnam,1900), pp. 265-267; Eduard Hitzig, "Alte und neue Untersuchungen uber das Gehirn: III. Das Versuch Loebs," Archiv fur Psychiatrie und Nervenkrankheiten 34 (1901):24-38.
51. Roben E. Kohler, From Medical Chemistry to Biochemistry (Cambridge: Cambridge University Press, 1982), pp. 300-301; see also S. S. Cohen, "Some Struggles of Jacques Loeb, Albert Mathews, and Ernest Just at the Marine Biological Laboratory," Biological Bulletin 168 (suppl.) (1985): 130-132.
52. JL to W. R. Harper, l l January 1902, 22 January,1902, UChicPP 45:6; JL to Svante Arrhenius, 19 August 1909, SA; JL to Simon Flexner, 30 December 1922, LP box 4.
53. "Secret of the Nerves Found," New York Sun. 31 December1901;--Why Our Grandchildren May Reasonably Expect to Live 150 Years, by Professor Jacques Loeb," New York Journal 12 January 1902, magazine; "A New Theory of Nerve Action," Literary Digest 24 (18 January 1902):81-82; JL to Harper, 25 January 1902, UChicPP 45:6.
54. JL to Harper, 26 January 1902, UChicPP 45:6; "University of Chicago Senate Minutes," 15 March 1902, UChicPP.
55. JL to Mall, n.d. [1897?], CDE-FM; "University of Chicago Board of Trustees Minutes," 20 February 1900, 30 April 1901, UChicPP; Simon Flexner to Mall, 29 August 1902, CDE-FM.
56. JL to Mall, 21 October 1900, 28 November 1900, CDE-FM; JL to Harper, 8 March 1900, UChicPP 17:13; Kohler, Medical Chemistry, pp. 145- 148; F. R. Lillie, The Woods Hole Marine Biological Laboratory (Chicago: University of Chicago Press, 1944), pp. 43-62.
57. JL to Mall, 13 February 1901, CDE-FM.
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