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Huxley, Haeckel, and the Oceanographers: The Case of Bathybius haeckelii Author(s): Philip F. Rehbock Source: Isis, Vol. 66, No. 4 (Dec., 1975), pp. 504-533 Published by: The University of Chicago Press on behalf of The History of Science Society Stable URL: http://www.jstor.org/stable/228925 . Accessed: 15/05/2013 17:26 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. . The University of Chicago Press and The History of Science Society are collaborating with JSTOR to digitize, preserve and extend access to Isis. http://www.jstor.org This content downloaded from 150.135.114.195 on Wed, 15 May 2013 17:26:47 PM All use subject to JSTOR Terms and Conditions
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Huxley, Haeckel, and the Oceanographers: The Case of Bathybius haeckeliiAuthor(s): Philip F. RehbockSource: Isis, Vol. 66, No. 4 (Dec., 1975), pp. 504-533Published by: The University of Chicago Press on behalf of The History of Science SocietyStable URL: http://www.jstor.org/stable/228925 .

Accessed: 15/05/2013 17:26

Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at .http://www.jstor.org/page/info/about/policies/terms.jsp

.JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range ofcontent in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new formsof scholarship. For more information about JSTOR, please contact [email protected].

.

The University of Chicago Press and The History of Science Society are collaborating with JSTOR to digitize,preserve and extend access to Isis.

http://www.jstor.org

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Huxley, Haeckel, and the

Oceanographers: The Case of

Bat hybius haeckelii

By Philip F. Rehbock*

INTRODUCTION

AT THE NORWICH MEETING of the British Association for the Ad- vancement of Science in August 1868, Thomas Henry Huxley announced

the results of his microscopical examination of some specimens of North Atlantic bottom sediment. Among the constituents of these seemingly unexciting speci- mens Huxley had found some curious "granule-heaps," surrounded by extensive masses of viscous matter, which he described variously as "lumps of a transparent, gelatinous substance" and a "colourless, and structureless matrix."' Tests of the substance convinced Huxley that he was dealing with a new type of organism in the form of virtually undifferentiated protoplasm. He gave the organism the generic name Bathybius, after its oceanic habitat, and called the species B. haeckelii, after his friend and fellow champion of Darwinism, Ernst Haeckel.

Bathybius haeckelii was to live a brief but eventful life of some seven years. In 1875 the scientists aboard H.M.S. Challenger, then nearing the end of their epochal voyage for the establishment of the new science of oceanography, discovered that Bathybius was nothing more than an inorganic precipitate. Despite his obvious embarrassment, Huxley readily admitted his error. Haeckel, however, who had applauded Huxley's original discovery, would not give up so easily. For him Bathybius was both an important member of his new class of unicellular organisms, the Monera, and a keystone in his mechanistic philosophy of life.

Received May 1974: revised/accepted June 1975. *Department of General Science, University of Hawaii, Honolulu, Hawaii, 96822. I am greatly

indebted to Professor Camille Limoges of the Institut d'Histoire et de Sociopolitique des Sciences, Universite de Montreal, who suggested the topic of this study and whose inspiration and criticisms were essential at each stage of the research. I would also like to thank Professor William Coleman of the History of Science Department and Professors Jeremy B. C. Jackson and Steven M. Stanley of the Earth and Planetary Sciences Department, The Johns Hopkins University, for their careful reading and comments. I also wish to thank the National Science Foundation for support of this study. An abbreviated version of the paper was presented at the Joint Atlantic Seminar for the History of Biology on April 7, 1973. This paper was awarded the 1974 Henry Schurman prize of the History of Science Society.

'T. H. Huxley, "On Some Organisms Living at Great Depths in the North Atlaritic Ocean," Quarterly Journal of Microscopical Science, 1868, N.S. 8:203-212 (p. 205). See also T. H. Huxley, "On Some Organisms which Live at the Bottom of the North Atlantic, in Depths of 6000 to 15,000 Feet," Report of the British Association for the Advancement of Science, 1868, p. 102.

504

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BATHYBIUS HAECKELII 505

Another decade would pass before he would abandon Huxley's creation. Previous narrations of the Bathybius story have been characterized by a brevity

and jocularity which obscures the seriousness of the controversy and the zeal of the debators. To be sure, there is a comical tone to the affair, deriving from the comments of the participants, especially Huxley, and from the subject matter itself (reminiscent of stories of sea serpents and other abyssal curiosities). But there is also a grave aspect, manifested in the immense efforts expended on laboratory investigations, ocean dredgings, and publications in scientific journals, all in behalf of a nonexistent organism.

Loren Eiseley has called Bathybius "one of the most peculiar and fantastic errors ever committed in the name of science."3 At first sight it may seem strange that some of the most distinguished biologists of the nineteenth century should have been allied with such a "fantastic" error. Professor Eiseley claims it to be "the product of an over-confident materialism, a vainglorious assumption that the secrets of life were about to be revealed."4 This analysis is incomplete, however, for it lays the blame on a philosophical predisposition while ignoring the unique sequences of events in several scientific disciplines whose timely convergence made Bathybius"'discovery" possible.

Bathybius was simultaneously a candidate for the lowliest form of protozoologi- cal life, the elemental unit of cytology, the evolutionary precursor of all higher organisms, the first organic form in the fossil record, a major constituent of modern marine sediments, and a source of food for higher life forms in the otherwise nutrient-poor deep oceans. Among biological entities Bathybius was probably unsurpassed in the variety of scientific specialties from which confirmation seemed forthcoming. Its eventual refutation came from outside this group of specialties, from the knowledge and techniques of chemical analysis. The present study attempts to trace out the full course of Bathybius' birth, development, and demise, emphasizing the complex milieu of scientific theories which allowed the "organism" its brief existence.

PRENATAL CONDITIONS OF BATHYBIUS

The idea that life on earth arose originally in the sea is one of the oldest speculations of natural history. Variations on this theme appear in the philosophy of the Milesian Anaximander5 and more recently in the speculative biology of the Naturphilosophe Lorenz Oken (1779-1851). Oken believed that life began as a primitive mucous substance which evolved from inorganic constituents existing in shallow marine waters. He equated the individual vesicles of this mucus with the smallest organisms known at the time, the infusoria. All other

2E.g., Susan Hubbard, "Beer, Bologna and Bathybius," Oceans, 1969, I(No. 3):23-26. 'Loren Eiseley, The Imm itse Journey (New York: Vintage Books, 1959), pp. 34-35. 4 Ibid. The prevalence of this spirit of the maturity of science in the latter half of the nineteenth

century has been discussed in a suggestive essay by Lawrence Badash, "The Completeness of Nineteenth-Century Science," Isis, 1972, 63:48-58.

5John Burnet, Early Greek Philosophy (3rd ed., London: A. C. Black, 1920), pp. 7071.

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506 PHILIP F. REHBOCK

organisms, according to Oken, were made up of, were "metamorphoses" of, these infusoria.6

Oken's writings, though mystical in tone and erroneous in many details, were strikingly anticipatory of later developments in biology, including the cell theory of Schleiden and Schwann and the protoplasm theory which followed it.7 The early history of these theories has been well described elsewhere.8 For present purposes it should be recalled that by mid-century the initial conception of the cell, a unit characterized by a definite boundary or cell wall, was being questioned and new light was being cast on the nature of the cell's contents. Franz Unger's demonstration of the identity of the cell substances of plants ("sarcode") and animals ("protoplasm") in 1850 initiated an era of intensive protoplasmic research.

The decade of the 1860s witnessed the most intensive investigations and speculations about protoplasm. Two events of particular interest marked the beginning and the end of this decade. In 1861 the German microscopist Max Schultze (1821-1874) published his essay "Uber Muskelk6rperchen und dass was man eine Zelle zu nennen habe,"9 in which he described his examination of membraneless protoplasm in marine invertebrates. Schultze's work convinced most scientists that the truly essential portion of the cell was not its outer membrane but its contents, the protoplasm and nucleus.'0 And in 1868 T. H. Huxley, by then a protoplasm devotee himself, placed the protoplasmic theory before the public in his famous lecture "On the Physical Basis of Life." "'

Huxley's lecture generated much interest because, along with his strong support of the protoplasmic theory and its physicochemical basis, he brought serious criticisms against both vitalists and positivists. 12 In his espousal of protoplasm Huxley failed to mention, however, that in an earlier essay he

6Lorenz Oken, The Elenents of Physiophilosophy, trans. Alfred Tulk (London: Ray Society, 1847), pp. 185-189. The 1st German ed. (Lehrbuch der Naturphilosophie) appeared in 1809. See also Lorenz Oken, Die Zeugung (Bamberg: Joseph Anton Goebhardt, 1805), p. 2.

7Oken pointed out these relationships himself in the preface to the 3rd ed. of his Lehrbuch (1847), pp. xi-xii.

8See John R. Baker, "The Cell Theory: A Restatement, History and Critique," Q. J. Micros. Sci., 1948-1955, 89, 90, 93, 94, 96; William Coleman, "Cell, Nucleus, and Inheritance: An Historical Study," Proceedings of the American Philosophical Society, 1965, 109:124-158; Gerald L. Geison, "The Protoplasmic Theory of Life and the Vitalist-Mechanist Debate," Isis, 1969, 60:273-292; Arthur Hughes, A History of Cytology (London: Abelard-Schuman, 1959).

9Max Schultze, "Uber Muskelkorperchen und dass was man eine Zelle zu nennen habe," Archiv fur Anatomie, Physiologie und wissenschaftliche Medizin, 1861, pp. 1-27.

'0Geison ("Protoplasmic Theory," p. 276) states that "the publication of this paper, more than any other single event, marked the birth of the protoplasmic theory of life." Erik Nordenski6ld (The History of Biology, New York: Knopf, 1928, p. 404) goes even further, claiming that Schultze's work "laid the foundations on which cell research has since been built, and this marks a new era in the science of cytology."

"T. H. Huxley, "On the Physical Basis of Life," Fortnightly Review, 1869, N.S. 5:129-145. I have used the republication of this essay in T. H. Huxley, et al., Half-Hours with Modern Scientists (New Haven, Conn.: Charles C. Chatfield, 1871), pp. 7-35.

'2One of Huxley's most vocal opponents in the vitalist-mechanist controversy was the London microscopist and Kings College professor of physiology Lionel S. Beale (1828-1906). Beale countered Huxley's "Physical Basis" in his Protoplasm, or Life, Matter and Mind (London: J. Churchill and Sons, 1870). Geison ("Protoplasmic Theory," pp. 285-290) discusses this controversy at length. See also Sydney Eisen, "Huxley and the Positivists," Victorian Studies, 1964, 7:337-358.

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BATHYBIUS HAECKELII 507

had argued for a much different conception of the cell, one which emphasized the cell membrane ("periplast") at the expense of the intracellular material ("endoplast"). This argument appeared in his 1853 review of cytological research, "The Cell-theory." 13 Thus, sometime between 1853 and 1868 Huxley's ideas on the fundamental nature of the cell made a full turnabout.

There is apparently no direct evidence of how or when Huxley's reversal came about. Perhaps he was convinced by Schultze's work of 1861 or by the German mycologist Anton de Bary's researches on the membraneless myceto- zoans (slime molds) published in 1864.14 It is also possible that he changed his mind as a result of his discovery of the purely protoplasmic Bathybius in the summer of 1868. In any case the timing of events testifies that Bathybius must at least have given confirmation to his change of mind, for it was only three months after his announcement of its discovery to the British Association that he lectured "On the Physical Basis of Life." In the intervening fifteen years since "The Cell-theory" he had published nothing on the subject of cells, his principal concerns having been vertebrate palaeontology and the defense of Darwinism.

If Bathybius appeared in some minds to be the culmination of half a century of cytological research, it was seen as the conclusion of an even longer period of investigations in protozoology. The earliest attempt to describe and systemati- cally classify those life forms which exist at or below the threshold of human vision was that of the Dane Otto Frederik Muller (1730-1784). Muller's Animalcula infusoria fluviatilia et marina,'5 published posthumously in 1786, represented the "state of the art" at the time Oken wrote of his infusorial mucous vesicles. The nineteenth-century researches of Christian Ehrenberg, Felix Dujardin, and Karl von Siebold greatly refined infusorial conceptions and classifications, and by 1845 unicellular organisms of irregular form and simple organization were referred to as the Protozoa.'6

At the protozoan level of complexity the difficulty of differentiating between plant and animal soon became obvious. Ernst Haeckel encountered this problem

'3T. H. Huxley, "The Cell-theory," British and Foreign Medico-Chirurgical Review, 1853, 12:221- 243. This was apparently the first instance of Huxley's going astray in cytology, Bathybius being the last. In the 1853 essay Huxley not only clung obstinately to the importance of the cell wall, while others were realizing that it was not an essential constituent; he also abandoned the nucleus as a significant factor, although it was well accepted and had been since the 1840s. See Baker, "Cell Theory," 1949, Part II, 90:106. For reasons unknown, the date of "The Cell-theory" has been confused, though there can be little doubt that the correct year was 1853. The Royal Society Catalogue of Scientific Papers and Leonard Huxley's Life and Letters of Thtomas Henry Huxley (New York: D. Appleton, 1900), Vol. I, p. 152, both give the year as 1858.

14Heinrich Anton de Bary, Die Mycetozoen (Schleimpilze) (2nd ed., Leipzig, 1864). See Baker, "Cell Theory," Part II, 90:97. Huxley mentioned de Bary in the "Hunterian Lectures on the Invertebrata," Q. J. Micros. Sci., 1868, N.S. 8:126-129, 191-202 (p. 127).

150. F. Miller, Animalcula infusoria fluviatilia et marina (Hanniae, 1786). '6Nordenskibld (History of Biology, pp. 426-430) treats this period of protozoology in some

detail.

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508 PHILIP F. REHBOCK

in his Generelle Morphologie der Organismen (1866) and solved it by establishing a third kingdom, the Protista, for the questionable, unicellular organisms.'7 The Generelle Morphologie was Haeckel's first attempt to found a taxonomy of nature upon evolutionary principles. His taxonomy took the form of genealogical trees (Stammbaume), with man at the apex and the Protista at the base. One of Germany's most zealous evolutionists, Haeckel believed that the unicellular organisms were the foundation of all higher forms of life and that their study was therefore of critical importance. His first monograph, on the planktonic Radiolaria, was based on these conceptions and secured him a professorial post at Jena in 1862.18 Subsequently extending the roots of his Stammbaume toward organisms of greater and greater simplicity, Haeckel created the class Monera of the Protista. The Monera included the lowliest forms of life, lacking in nucleus and exterior membrane: "an entirely homoge- neous and structureless substance, a living particle of albumen, capable of nourishment and reproduction." 19

Huxley had read and applauded the Generelle Morphologie and had even expended considerable effort, fruitlessly, to have it translated into English.20 He was not entirely convinced about the Protista, however. In the Hunterian Lectures, early in 1868, his opinion was that some of the Protista were decidedly plants and others animals. Only the Monera were acceptable as "intermediate ground," such as the organism Haeckel had named Protogenes: "The simplest bit of living matter possible," whose lack of a nucleus "proves the absence of any mysterious power in 'nuclei' and shows that life is a property of the molecules of living matter, and that organization is the result of life, not life the result of organization." 21

Haeckel pursued his researches on the Protista throughout the 1860s in the belief that he was at the same time closing the gap between cytology and protozoology. In the spring of 1868, barely a few months before Huxley was to discover Bathybius, Haeckel published his "Monographie der Moneren," an elaboration of the class established in the Generelle Morphologie. The seven species of Monera which had been described up to that time were of interest, Haeckel claimed, because of the completely structureless homogeneity of the substance composing them and, more importantly, because of the similarity of this substance to the protoplasm of all plant and animal cells. Haeckel had been a firm proponent of the protoplasm theory and was delighted that his Monera gave it additional support:

The Protoplasm or Sarcode theory, that is, that the albuminous contents of animal and vegetable cells (or more correctly, their "cell-matter"), . . . are identical, and

17 Ernst Haeckel, Generelle Morphologie der Organismen (Berlin: Georg Reimer, 1866), pp. 198-206. 18 Ernst Haeckel, Die Radiolarien (Berlin: Georg Reimer, 1862). I)Ernst Haeckel, "Monographie der Moneren," Jenaische Zeitschrift fur Medicin und Naturwissen-

schaft, 1868, 4:64. I have employed the translation by W. F. Kirby and E. P. Wright, Q. J. Micros. Sci., 1869, N.S. 9:27-42, 113-134, 219-232, 327-342 (p. 28).

20Georg Uschmann and Ilse Jahn, eds., "Der Briefwechsel zwischen Thomas Henry Huxley und Ernst Haeckel," Wissenschaftliche Zeitschrift der Universitht Jena (Math.-Nat. Reihe), Jg. 9 (1959/1960), H. 1/2, p. 13.

21 Huxley, "Hunterian Lectures on the Invertebrata," pp. 126-127.

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BATHYBIUS HAECKELII 509

that in both cases this albuminous material is the original active substratum of all vital phenomena may perhaps be considered one of the greatest achievements of modern biology, and one of the richest in results. ... By no phenomenon is the correctness of this theory so thoroughly proved, and, at the same time, in so simple and unassailable a manner, as by the vital phenomena of the Monera, by the processes of their nourishment and reproduction, sensitiveness and motion, which entirely proceed from one and the same very simple substance, a true "primitive slime." 22

Not surprisingly, the Monera were to become Bathybius' closest relatives. And Huxley's recent acquaintance with the "Monographie der Moneren" was undoubtedly responsible, at least in part, for his ability to see Bathybius as a living organism. But for Haeckel, Bathybius had a more momentous implication (an implication which occurred to Huxley but which he never accepted publicly): the possibility that it arose by spontaneous generation out of inanimate matter on the ocean floor.

Of the various theories which formed a background for the acceptance of Bathybius, the idea of spontaneous generation had probably the longest, most eventful history. John Farley has recently provided new clarity in our under- standing of the spontaneous generation controversy by emphasizing that two distinctly different concepts were involved and often were confused by scientists and later by historians of science.23 From the seventeenth through the mid-nine- teenth centuries "spontaneous generation" usually referred to the apparent development or "heterogenesis" of intestinal worms, infusorians, yeasts, bacteria, and certain insects within an organic environment where no such organisms were thought to have been previously present. By the mid-nineteenth century the researches of J. J. S. Steenstrup, Ehrenberg, Pasteur, and many others had convinced most scientists that the need to invoke heterogenesis had been eliminated. At the same time a second meaning of spontaneous generation was becoming increasingly common, namely "abiogenesis," or the evolution of living organisms out of inorganic matter.24 The popularity of this notion, first in Germany (1860s) and then in Britain (1870s), derived its energy chiefly from Darwinism and from the German preoccupation (inherited from the earlier Naturphilosophie) with a Weltanschauung or universal system of knowledge.

Darwin was able to provide a mechanistic explanation for the diversity of organisms and their alteration through time, but as to the source of the first primitive organisms on earth, he refused to comment. Here Haeckel entered the picture. He insisted that the complete mechanistic Weltanschauung demanded an abiogenetic starting point as an a priori necessity. And he set out in search

22 Haeckel, "Monographie der Moneren," pp. 223-224. 23 John Farley, "The Spontaneous Generation Controversy (1700-1860): The Origin of Parasitic

Worms," Journal of the Histoiy of Biology, 1972, 5:95-125, and "The Spontaneous Generation Controversy (1859-1880): British and German Reactions to the Problem of Abiogenesis," J. Hist. Biol., 1972, 5:285-319.

24 Abiogenesis continues to be the subject of speculation and experiment in the twentieth century. Well-known works include A. I. Oparin, The Origin of Life (New York: Macmillan, 1938), John Keosian, The Origin of Life (New York: Reinhold, 1964), S. L. Miller, "A Production of Amino Acids Under Possible Primitive Earth Conditions," Science, 1953, 117:528-529.

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510 PHILIP F. REHBOCK

of ever-simpler protozoans, hoping to narrow the gap between life and nonlife.25 The simpler the organism, the smaller the gap between it and its abiotic environment, and thus the easier to conceive of a bridging of the gap by a purely chemical process of spontaneous generation or "archegony." With his work on the Protista and then the Monera, Haeckel saw the gap closing:

. . . the acceptation of a genuine archegony (once or repeated) has at present become a logical postulate of scientific natural history. Most naturalists who have discussed this question rationally believed that they must designate simple cells as the simplest organism produced thereby, from which all others developed themselves. But every true cell already shows a division into two different parts, i.e., nucleus and plasm. The immediate production of such an object from spontaneous generation is obviously only conceivable with difficulty; but it is much easier to conceive of the production of an entirely homogeneous, organic substance, such as the structureless albumen-body of the Monera.26

With his pedigree of the natural world now extending down to the Monera, the next logical step Haeckel hoped for in his research was the discovery of the on-going formation of living protoplasm in the Monera from strictly nonliving matter. Such a discovery was not only desirable for the sake of biological completeness; it was essential to Haeckel's monistic philosophy and his struggle against the vitalist conception of nature. Within a few months of his publication of the "Monographie der Moneren" Haeckel's biological and philosophical aspirations were to be triumphantly fulfilled, though only temporarily, by Huxley's Bathybius.

The role of Bathybius in protozoology, cytology, and abiogenesis gave it a position of credibility for biologists. The immediate source of its discovery, however, was not biology but the study of marine geology, then becoming active in Britain. Since the 1850s there had been a growing interest, especially among the British, in the acquisition of bathymetric data for the purpose of laying telegraph cables. A spin-off from the new technology of sounding the depths was the collection of small bottom samples via the sounding device. One of the most common constituents of these samples was the calcareous shells of the protozoan group known as foraminifera, mainly of the genus Globigerina. Early studies of these organisms had been made by Christian Ehrenberg in Germany and by Jacob Whitman Bailey in the United States, but no one was able to state decisively whether these foraminifera lived on the bottom or merely fell there from the surface after their death. Ehrenberg

251t is especially interesting that Haeckel chose to find the missing evolutionary links by a study of existing life forms rather thap fossil forms. He was, of course, trained as a biologist, not as a palaeontologist. But more important, he may have considered the study of fossilized microorganisms unduly difficult and even unnecessary, if, as he believed, these same organisms were still being spontaneously generated at the present time (see below).

26Haeckel, "Monographie der Moneren," p. 30.

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BATHYBIUS HAECKELII 511

believed them to be benthic-that is, bottom dwellers- while Bailey regarded them as planktonic.27

In the summer of 1857 Huxley, then palaeontologist at the London School of Mines, looked forward to receiving some fresh data on the Globigerina question. H.M.S. Cyclops, under the command of Captain Joseph Dayman, had just completed a line of soundings from Great Britain to Newfoundland and back, in preparation for a cable laying. Huxley had earlier supplied the ship's medical officer with instructions for obtaining, observing, and preserving whatever deposits might be brought up by the sounding device. In his instructions Huxley drew special attention "To the preservation of the freshly brought up soundings in a tolerably strong alcoholic mixture, so that the presence or absence of soft parts in them might be determined at any future time, and under more convenient circumstances." 28 The use of alcohol as a preserva- tive (a practice dating back to Robert Boyle 29) was, on the one hand, a reasonable precaution, as there was no opportunity to examine the numerous samples aboard ship. On the other hand, alcohol was to become one of the chief culprits in the generation of Bathybius, as will shortly be seen.

The samples reached Huxley in the autumn of 1857. His preliminary examination of them was published as an appendix to Captain Dayman's report of the results of the soundings.30 Huxley's examination had been only a superficial one, apparently just enough to permit the preparation of a report to the hydrographer of the Admiralty, who had sanctioned the collection. As Huxley stated in the report, "I am desirous of briefly laying their results before you, reserving for a future occasion the communication of the full details, and of the illustrations, which, in accordance with the authorization of the Admiralty, I am now having prepared to accompany them.'

In his initial examination Huxley found the samples to be uniformly composed of "an excessively fine, light brown, muddy sediment." Microscopic inspection revealed large amounts of foraminifera, "fully nine-tenths, as I imagine, by weight." Although he was unable to determine with certainty whether they were normally bottom dwellers or not, he was, like Ehrenberg, inclined to

27 Margaret Deacon, Scientists and the Sea, 1650-1900 (London: Academic Press, 1971), p. 297. For a contemporary account of the foraminifera question, see Matthew Fontaine Maury, The Physical Geography of the Sea and its MHeteorology, ed. John Leighly (Cambridge, Mass.: Belknap Press, 1963; from the 8th ed., 1861), pp. 297-304. The belief that foraminifera did not and could not live at the bottom of the sea was based in part on the "azoic zone" hypothesis, suggested in the 1840s by Edward Forbes (1815-1854). The leader of Britain's proto-oceanographers in the second quarter of the nineteenth century, Forbes theorized that the diminishing quantity of bottom fauna at greater and greater depths indicated that no life existed below approximately 300 fathoms. The azoic-zone hypothesis is discussed at length in Ch. V of my Ph.D. dissertation, "Organisms in Space and Time: Edward Forbes (1815-1854) and New Directions for Early Victorian Natural History" (The Johns Hopkins University, 1975).

28Joseph Daymnan, Deep-Sea Soundings in the North Atlantic made in H.M.S. "Cyclops" in June and July, 1857 (London: H.M.S.O., 1858), p. 63.

29Robert Boyle, "A Way of Preserving Birds taken out of the Egge, and Other Small Foetus's," Philosophical Transactions of the Royal Society, 1666, 1:199-201. See also Hughes, History of Cytology, p. 14.

30Daymnan, Deep-Sea Soundings, App. A, pp. 63-68. 31 Ibid., pp. 63-64.

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512 PHILIP F. REHBOCK

believe that they were. In addition to the predominance of foraminifera in the samples, Huxley noted considerable quantities of "curious rounded bodies . . . looking at first sight somewhat like single cells of the plant Protococcus." Since these minute particles were easily dissolved by acid, Huxley decided they could not be organic; ". . . I will, for convenience sake, simply call them Coccoliths."32 The Coccoliths were to have an interesting and error-ridden history and a controversial relationship to Bathybius.

For reasons not fully clear, Huxley set aside these bottom samples for over ten years. Presumably he was satisfied that little else was to be gained by further study with his existing instruments; and other projects of higher precedence, notably his defense of Darwin's theories, were soon to place heavy demands on his time. Nevertheless, his observations, though buried in Dayman's report, were not altogether ignored by other microscopists. In 1861 the London physician and marine enthusiast George Charles Wallich (1815-1899) published several articles on his deep-sea dredging experiences of the preceding year.33 Wallich noted "great numbers" of the Coccoliths Huxley had described but found that they existed, not only in the free state observed by Huxley, but attached to the outer surface of spherical bodies "in such a manner as to leave no doubt of that being their normal position."34 Wallich thought these bodies, which he named "Coccospheres," were the larval form of some type of foraminifer. He claimed, moreover, that when the outer shell of the Coccosphere was crushed it was found to contain a protoplasmic substance, "a homogeneous, gelatinous, and almost colourless matter, exhibiting no visible trace of organization, and, in all probability, consisting of sarcode. The wall of the cell may be distinctly seen under a high power; but from the minuteness of the entire structure, I have hitherto found it impossible to do more than attest its existence." 35

Little more was heard on the subject of Coccoliths and Coccospheres36 until Huxley gave his reknowned lecture "On a Piece of Chalk," in 1868. In that lecture, presented to working men at Norwich during the British Association meeting, Huxley eloquently demonstrated the identity of the chalk deposits to present deep-sea mud. This was an opportunity to bring together, for the layman, the discoveries of Coccoliths and Coccospheres and to reiterate his

32 Ibid., p. 64. 33Wallich accompanied H.M.S. Bulldog during her soundings between the Faroe Islands,

Greenland, and Labrador. He published a full account of his findings in The North Atlantic Sea-Bed: Comprising a Diary of the Voyage on Board H.M.S. "Bulldog" in 1860 (London: John Van Voorst, 1862).

34G. C. Wallich, "Remarks on Some Novel Phases of Organic Life, and on the Boring Powers of Minute Annelids, at Great Depths in the Sea," Annals and Magazine of Natural History, 1861, 3rd Ser., 8:52-58 (pp. 52-53). (I have not been able to examine Wallich's first work on this subject, "Notes on the Existence of Animal Life at Vast Depths in the Ocean," published privately in 1860.)

35Wallich, "Remarks," p. 53. 36In 1865 Wallich reported he had found Coccospheres "free-floating in tropical seas," but

this important observation, mentioned only in a footnote to an article on another subject, seems to have been missed by his contemporaries. G. C. Wallich, "On the Structure and Affinities of the Polycystina," Transactions of the Microscopical Society, 1865, N.S. 13:57-84 (p. 81).

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BATHYBIUS HAECKELII 513

views on the benthic habitat of Globigerina. Huxley added that he had recently studied the Coccoliths a second time, convincing himself that "they are produced by independent organisms, which, like the Globigerina, live and die at the bottom of the sea."37 The "independent organismn" alluded to was his newly found Bathybius haeckelii.

Yet another organism, not mentioned by Huxley but of much greater antiquity than the chalk deposits, was shortly to become allied with Globigerina and Bathybius. The "dawn animal of Canada," Eozoo'n canadense, extended the time dimension of Bathybius back to the Precambrian and gave scientists of North America a role in Bathybius' history. The Eozoo'n geological configuration, a complex layering of limestone and serpentine, had been discovered in 1858 in the Laurentian limestones of Canada.38 The case for its organic origin was not pursued until 1864, when a campaign was begun by John William Dawson (1820-1899), palaeontologist, principal of McGill College, and civic leader of Montreal.39 In England Dawson enlisted the support of the respected micro- scopist, physiologist, and subsequent organizer of the Challenger expedition William Benjamin Carpenter (1813-1885). Carpenter informed the Royal Society of Dawson's "revolutionary" Eozoo'n, a distant relative, he thought, of modern foraminifera.4" Soon after Dawson's description of it in the Journal of the Geological Society (1865)41 the scientific forces began to align themselves: for Eozoon canadense, the Precambrian protozoan; and for eozo6n, the mineralogical phenomenon.

At first the heavy artillery seemed to be on Dawson's side. Max Schultze's examination of an Eozoon specimen left him with "no serious doubt" as to its foraminiferal nature;42 in the fourth edition of the Origin (1866) Darwin added his impression that "it is impossible to feel any doubt regarding its organic nature;"43 and Huxley commented, early in 1868, that Eozoon was

37T. H. Huxley, "On a Piece of Chalk," AMacmillan's Magazine, 1868, 18:39-408. I have utilized the version in Huxley's Discourses Biological and Geological (New York: Appleton, 1896), pp. 1-36 (pp. 17-18).

38William E. Logan, "On the Laurentian Limestones," Canadian Naturalist antd Geologist, 1859, 4:300.

39Dawson's interesting career has been chronicled in Charles F. O'Brien's "Sir William Dawson: A Life in Science and Religion," Memoirs of the American Philosophical Society, June 1971, 84. The Eozo6it controversy is covered more thoroughly by O'Brien in "Eozoon Canadense, 'The Dawn Animal of Canada,'" Isis, 1970, 61:206-223. See also M. E. Mitchell, "On Eozoon Canademse" and C. F. O'Brien, "Comments," Isis, 1971, 62:381-383.

40W. B. Carpenter, "On the Structure and Affinities of Eozoon Canadense," Proceedings of the Royal Society of London, 1863-1864, 13:545-549.

41J. W. Dawson, "On the Structure of Certain Oiganic Remains in the Laurentian Limestones of Canada," Quarterly Journal of the Geological Society of Loadon, 1865, 21:51-59.

42 George P. Merrill, The First One Hundred Years of Americant Geology (New Haven: Yale University Press, 1924), p. 572.

43 The Origin of Species by Charles Darwin: A Variorum Text, ed. Morse Peckham (Philadelphia: University of Pennsylvania Press, 1959), p. 515.

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514 PHILIP F. REHBOCK

"fairly proved to be an encrusting Foraminifer." 44 The leading opponents of Eozodn were William King and Thomas H. Rowney, mineralogists at Queen's College, Galway, Ireland.45 The controversy was truly international. For two decades the scene of action shifted among England, Ireland, Germany, Canada, and the United States. Most of the activity centered on a heated, not always becoming exchange of articles between Dawson and Carpenter on the one side and King and Rowney on the other. The exchange subsided after 1874,46 but resolution of the issue (in favor of the mineralogists) had to wait until the twentieth century.47

In the beginning the Eozoon controversy was a purely geological one. It might have subsided sooner had it not been for the new interest in the habitat of the foraminifera and the discovery of Bathybius. In some minds there was a reinforcement of belief in each of the organisms by the other two. All three organisms seemed to be of protozoanal simplicity, to be distributed over wide geographic areas, and to be associated with calcareous bodies. If the foraminifera were bottom dwellers, then they might well be related to Bathybius; and if the Eozoon fossil configuration had been originally formed on the floor of the primitive oceans and later elevated to its Canadian location,48 then it was easy to imagine Eozoon as the Precambrian ancestor of modern foraminifera and of Bathybius.

By the mid-1860s, then, several lines of biological and geological research seemed to be approaching a common ground. In cytology, Schultze's work had affirmed that an amorphous protoplasm was the essential component of the cell. Haeckel's study of the protozoa had yielded a number of species that seemed to contain little more than this same protoplasm. Further, Haeckel advanced the notion-by a logic he felt certain would be empirically con-

44Huxley, "Hunterian Lectures," p. 129. 45Their opposition was first expressed in a letter to the editor, Reader, 1865, 5:660. King and

Rowney included an "annotated history" of the controversy in their book An Old Chapter of the Geological Record . . . (London: John Van Voorst, 1881). Their chronology (1858-1880) mentions over 40 participants, representing Canada, the United States, Britain, Germany, Spain, France, and Russia.

46See W. B. Carpenter, "Final Note on Eozoon Canadense," Ann. Mag. Nat. Hist., 1874, 4th Ser., 14:371-372, in which the pointlessness of continued debate is emphasized.

47 O'Brien, "Eozoain canadense, 'The Dawn Animal of Canada,' " p. 223. For recent pronouncements on Eozoon, see also Percy E. Raymond, "Pre-Cambrian Life," Bulletin, Geological Society of America, 1935, 46:375-392 (pp. 377-378); Thomas H. Clark and Colin W. Stearn, Geological Evolution of North America (2nd ed., New York: Ronald Press, 1968), pp. 32, 402, 408.

48 The idea of massive shifts in the relative positions of the ocean floor and the continents was a popular one at this time. It was invoked by some, including Huxley ("On a Piece of Chalk") and C. Wyville Thomson (The Depths of the Sea (London: Macmillan, 1872), Ch. 10) to explain the similarity of modern ocean sediments to the ancient chalk formations. This idea, known as the "continuity of the chalk," was a favorite one of Thomson's, chief scientist of the Challenger expedition. His first pronouncement on the subject was a lecture in April 1869. A recent discussion of the concept appears in Susan Schlee, The Edge of an Unfamiliar World (New York: Dutton, 1973), pp. 96-98, 142-143.

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BATHYBIUS HAECKELII 515

firmed-that this protoplasm, the evolutionary basis of all subsequent life, had been generated from nonliving matter. Concurrently, the early marine geologists were pondering the origin of certain minute calcareous constituents of the newly won ocean sediments, while palaeontologists were seeking a cause for equally minute and strikingly similar structures of the Precambrian era. The completion of this rather complex web awaited the discovery of an organism composed entirely of undifferentiated protoplasm, capable of acquiring a calcium shell, and living in an environment where conditions were sufficiently homoge- neous through space and geological time to minimize the tendency for evolution or extinction.

In retrospect it would seem that the completion of this web with an organism such as Bathybius might have been effected by any microscopist working in the tradition of mechanistic biology. Haeckel and his colleagues were probably the most likely candidates. Haeckel's overly eager investigations of the Monera had already led him into classifications later proven erroneous.49 Carpenter would have been a second possible discoverer because of his interests in Eozoon, his study of foraminifera, and his connections with ocean dredging. The ill-fated discovery came to Huxley, however, because of the supremacy of the British navy in deep-sea sounding during the third quarter of the nineteenth century, and because of Huxley's access to the Admiralty through his position at the School of Mines and his former naval commission as assistant surgeon (1846- 1854).

THE BIRTH OF BATHYBIUS

... there are periods in the history of every science when a false hypothesis is not only better than none at all, but is a necessary forerunner of, and preparation for, the true one.

-T. H. Huxley50

In the spring of 1868 Huxley returned to the study of his sediment specimens. Why he did so is far from certain, but several conjectures are possible. He had recently obtained a much stronger microscope (1,200 diameters) manufac- tured by Andrew Ross,5' which he said "renders obvious many details hardly

49 See below, n. 114. 50"The Cell-theory," pp. 248-249. 5'The study of microorganisms was severely hampered by the state of microscope technology

until the mid-nineteenth century, when the achromatic compound microscope had become clearly superior to the single lens. Andrew Ross, one of the leading British microscope producers, constructed his instruments on the highly successful design of Joseph Lister, from about 1831 until his death in 1859. His son, Thomas Ross, then took over the business and during the 1860s began producing binocular microscopes. It is possible that Huxley's new microscope was one of these binocular models, but since he makes no mention of such an improvement, I assume that he continued to use the elder Ross' monocular type. The latter is depicted in Plt. I of John Quekett's A Practical Treatise on the Use of the Microscope (2nd ed., London: H. Baillier, 1852). See also Hughes, History of Cytology, p. 11; S. Bradbury, "The Quality of the Image Produced by the Compound Microscope: 1700-1840," in S. Bradbury and G. L'E. Turner, Historical Aspects of MVicroscopy (Cambridge: Heffer, 1967), p. 170: S. Bradbury, The Evolution of the Microscope (Oxford: Pergamon Press, 1967), pp. 194, 211.

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516 PHILIP F. REHBOCK

decipherable with the 1/6th inch objective which I used in 1857."52 Perhaps the sediments were merely a convenient subject on which to test the powers of the new instrument. It seems more likely, however, that he was seeking answers to some old problems. Were there any soft parts remaining in the Globigerina shells which would attest to their having been alive when the sounding device snatched them up? This could confirm Huxley's belief in their benthic existence. Were the Coccoliths actually just miniature plates which had come loose from the surface of Coccospheres, as Wallich thought, or did they normally exist independently of the Coccospheres, as Huxley had first suggested? 53

Were there Coccospheres in his samples which he had missed during his 1857 examination? If so, did they actually contain soft parts ("sarcode") as Wallich insisted? Were the sediments accumulating now on the ocean floor identical to those formed in the Cretaceous period? And what of Eozoon? Did any of the foraminifera in his sediments resemble the alleged Precambrian fossil? Any or all of these questions could have influenced Huxley's decision to reexamine the sediments, but his subsequent remarks indicate that some questions were more important than others.

Upon concluding his reexamination, Huxley reported his findings, verbally to the British Association in August 1868,"4 and in writing in the Quarterly Journal of Microscopical Science the following October.55 The reexamination revealed, in addition to the numerous familiar sediment constituents, some new members: (1) two distinct types of Coccoliths which Huxley named "Discoliths" and "Cyatholiths;" (2) the Coccospheres described by Wallich; and (3) protoplasm-like "granule-heaps" and "transparent gelatinous matter," which appeared to contain no nucleus and "no trace of a membranous envelope."56 He described this substance further as a

... deep-sea "Urschleim," which must, I think, be regarded as a new form of those simple animated beings which have recently been so well described by Haeckel in his "Monographie der Moneren." I proposed to confer upon this new "Moner" the generic name of Bathybius, and to call it after the eminent Professor of Zoology in the University of Jena, B. Haeckelii.57 [See Fig. 1.]

It is important to note that at this time Huxley was quite guarded in his statements about Bathybius. To affirm that it was living protoplasm was as far as he would go, a reluctance which was to lessen his embarrassment later

52Huxley, "On Some Organisms Living at Great Depths," p. 205. 53 Huxley referred to the Coccoliths as "curious rounded bodies" in the Hunterian Lectures

of 1868, indicating at least that he still regarded the question of their nature an open one ("Huriterian Lectures," p. 129).

54 Huxley, "On Some Organisms which Live at the Bottom of the North Atlantic." Huxley presented his "On a Piece of Chalk" at this same meetinig.

55Huxley, "On Some Organisms Living at Great Depths." This paper appeared just prior to his lecture "On the Physical Basis of Life."

56Huxley, "On Some Organisms Living at Great Depths," p. 206. 57Ibid., p. 210. During the latter half of the nineteenth century descriptions of the cytoplasm

of various animals frequently mentioned a "reticular or net-like structure," or "granules within an amorphous matrix." See Hughes, History of Cytology, pp. 112-120.

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BATHYBIUS HAECKELII 517

Figure 1. Bathybius haeckelii. Ernst Haeckel's drawing (B3iologische Studien, 1870) as it appeared in C. Wyville Thomson, The Depths of the Sea (New York: Macmillan,

1873), p. 412.

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518 PHILIP F. REHBOCK

on. Referring to Bathybius, the Coccoliths, Coccospheres, and other ingredients of the ooze, he insisted: "I have hitherto said nothing about their meaning, as in an inquiry so difficult and fraught with interest as this, it seems to me to be in the highest degree important to keep the questions of fact and the questions of interpretation well apart."58

Huxley did propose his theory that the Coccoliths and Coccospheres were structural components of Bathybius, analogous to the spicules of sponges. In fact, he seemed more concerned with the various calcareous bodies which had occupied Wallich than with the "new 'Moner."' He agreed that there was probably a connection between the Coccoliths and Coccospheres themselves; but instead of the former being merely cast-off sections of the latter, Huxley thought it equally likely that the Coccoliths existed independently and at some stage "coalesced" to form Coccospheres. In the ten-page report of the sediments only about two pages involved Bathybius itself; the remainder was concerned with the nature of and opinions on Coccoliths and Coccospheres. This emphasis is an indication that Huxley's main interest in reexamining the sediments had been to advance the dialogue with Wallich. Issues which were to become associated with Bathybius later (the nature of Eozo6n, the habitat of Globigerina) were not mentioned.

Huxley's first action after publishing his discovery was to write to Haeckel. Referring to his paper in the Quarterly Journal, he advised Haeckel:

It is about a new 'Moner' which lives at the bottom of the atlantic to all appearance, and gives rise to some wonderful calcified bodies....

I have christened it Bathybius Haeckelii and I hope that you will not be ashamed of your godchild. I will send you some of the mud itself with the paper. ..59

In his congratulatory reply, Haeckel wasted no time in pointing out the connection of Bathybius with Naturphilosophie:

I am, of course, most especially delighted by "Bathybius Haeckelii" and am very proud to be the godfather at its christening!

The Coccospheres are extremely remarkable; their relationship to the Discoliths and Cyatholiths is indeed still very puzzling. It will probably be some time before we are able to clear up all these lowest Protista of the ocean floor, and to establish something definite about the connections of all the uncertain protistic and protoplas- mic primitive organisms. But the "Urschleim" of Oken is clearly coming more and more back into favor, and my theory of plastids [cells with or without nuclei] can truly rejoice about that! "Vive Monera"!60

58Huxley, "On Some Organisms Living at Great Depths," p. 210. In the discussion following the paper, Huxley denied that his work gave any confirmation of the possibility of spontaneous generation. See "The British Association for the Advancement of Science," Journl of Science, 1868, 5:501-545 (pp. 529-530).

59Uschmann and Jahn, "Briefwechsel . . . Huxley . . . Haeckel," p. 18. 60 Ibid.

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BATHYBIUS HAECKELII 519

Despite Haeckel's enthusiasm, which was eventually to make him the most prolific writer on Bathybius, Huxley's British colleagues were the first to take up the subject publicly. In December 1868 Carpenter reported to the Royal Society on the dredging operations which he and Wyville Thomson had undertaken in H.M.S. Lightning the previous summer. Their brief excursion into the seas north of Scotland had yielded Coccoliths, Coccospheres, and, in the last dredging at 650 fathoms, a "particularly viscid mud.' This mud was examined by Huxley (though apparently not by Carpenter) and found to contain Bathybius, thus confirming his earlier conclusions.

Carpenter's report raised the primary question which was to reappear in nearly all later publications on Bathybius, the problem of its food source:

In what manner the materials for its protoplasm . . . are obtained, is a most perplexing problem. All the evidence we at present possess in regard to the alimentation of the Rhizopods, leads to the belief that, in common with higher Animals, they depend on Organic Compounds previously elaborated by Vegetative agency under the influence of the light and heat of the Sun. But every form of Vegetable life that is visible to the naked eye seems entirely wanting at great depths in the ocean. . .. It may be that the Bathybius . . has so far the attributes of a Vegetable, that it is able to elaborate Organic Compounds out of the materials supplied by the medium in which it lives, and thus to provide sustenance for the Animals imbedded in its midst.62

Despite the problems of its nourishment and its taxonomic status as animal or plant, Carpenter did not doubt Bathybius' existence. And he was enthusiastic in pointing out its probable geological age and relationship to Dawson's Eozoon:

The discovery of this indefinite plasmodium, covering a wide area of the existing Sea-bottom, should afford a remarkable confirmation, to such (at least) as still think confirmation necessary, of the doctrine of the Organic origin of the Serpen- tine-Limestone of the Laurentian Formation. For if Bathybius, like the testaceous Rhizopods, should form for itself a shelly envelope, that envelope would closely resemble Eozoon. Further, as Prof. Huxley has proved the existence of Bathybius through a great range not merely of depth but of temperature, I cannot but think it probable that it has existed continuously in the deep seas of all Geological Epochs.63

Carpenter may have been willing to accept Bathybius in spite of the theoretical difficulties it brought with it, but Wallich was far more skeptical. In a January

61W B. Carpenter, "Preliminary Report of Dredging Operations in the Seas to the North of the British Islands, carried on in Her Majesty's Steam-vessel 'Lightning,'" Proc. Roy. Soc. Lon., 1868, 17:168-200. See also Carpenter's "On the Rhizopodal Fauna of the Deep Sea," Proc. Roy. Soc. Lon., 1869-1870, 18:59-62.

62 Carpenter, "Preliminary Report," pp. 190-191. Henry Alleyne Nicholson, Professor of Natural History at St. Andrews, picked up this theme in the first edition of his A Manual of Zoology (Edinburgh/London: William Blackwood, 1870). He cited Bathybis as a possible exception to the rule that plants have the power to convert inorganic compounds into organic mnatter while animals lack this ability. But he added, "The water of the ocean, however, at these enormous depths, is richly charged with organic mnatter in solution, and this conjecture is thereby rendered doubtful" (p. 10).

63 Carpenter, "Preliminary Report," p. 191 n.

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520 PHILIP F. REHBOCK

1869 article Wallich discussed his researches on Coccoliths and Coccospheres at length. While Huxley had conjectured that these small calcareous bodies might constitute the skeletal structure of Bathybius, Wallich was convinced they were parts of independent organisms, having no direct relationship to Bathybius. As evidence of this independence he stated that in many bottom samples he had observed Coccospheres and Coccoliths "in profusion" where there was "scarcely a trace of muddy or slimy substance"-that is, Bathybius.f Wallich concluded by expressing "with great deference" his grave doubts about Huxley's supposed organism. He was inclined to regard it instead as a slime of decompos- ing plant and animal matter: "analogy, and the bulk of direct evidence, is in favour of the supposition that this widely distributed protoplastic matter is the product, rather than the source, of the vital forces which are already in operation at the sea bed."65

The following month Huxley had occasion to defend his discovery before a meeting of the Royal Geographical Society. The author of a paper on some new Gulf Stream data had claimed that no living matter was to be found on the bottom in the region he had sounded. After the paper's reading, Huxley countered that he had examined the soundings himself and had submitted them to Professor Edward Frankland for organic analysis. Frankland had found them to contain more than 1' per cent organic matter, identifiable in two forms: the shells of foraminifera and the "confused network" of Bathybius. These facts Huxley regarded as "now definitely acquired by science." 66

Moreover, recent soundings in the South Atlantic and Indian oceans had confirmed the existence of Bathybius there at depths up to 2,800 fathoms, proving its distribution to be truly global. But, he admitted, how animals could live at such depths, "how they acquired their store of food, was one of the most curious questions of organic chemistry; one which we could not solve at present."67

In the summer of 1869 Carpenter and Thomson carried out a second dredging cruise. Fresh Bathybius was obtained from the Bay of Biscay and described as being "capable of a certain amount of movement." 68 The notion of movement

64G. C. Wallich, "On the Vital Functions of the Deep-sea Protozoa," Monthly Microscopical Journal, 1869, 1:32-41 (p. 36).

65 Ibid., p. 39. Wallich added that Eozo6n had been used, he felt, prematurely to support Bathybius; too little was yet known about the former to be sure that it was not just as highly differentiated as ordinary foraminifera and therefore too complex to be allied to Bathybius. Ten years later Wallich, bitter over the lack of recognition of his deep-sea discoveries, sought to have Thomson and Carpenter expelled from the Royal Society for their plagiarism and misrepresentation of his ideas. When it was suggested that Huxley (then Royal Society secretary) be a party to the adjudication of the complaint, Wallich insisted that he could not accept any decision of Huxley's, because he and Huxley had earlier taken opposite sides on Bathybius. This dispute appears in a series of letters between Wallich and Joseph Hooker, 1877-1878 (Royal Society Miscellaneous Correspondence, Vol. XI, MSS 126-149). Wallich spelled out his position on Bathybius at length in "On the True Nature of the So-called 'Bathybius' and its Alleged Function in the Nutrition of the Protozoa," Ann. Mag. Nat. Hist., 1875, 4th Ser., 16:322-339.

66Huxley, [untitled discussion,] Proceedings of the Royal Geographical Society, 1869, 13:110. 67 Ibid. 68Thomson, The Depths of the Sea, p. 411. I have quoted throughout from the 2nd ed. (1874)

which is, so far as I know, identical to the 1st (1873).

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BATHYBIUS HAECKELII 521

was repeatedly emphasized by Haeckel as indubitable proof of its vitality when Bathybius later came under attack.69 No explanation has been given for this alleged movement, but one might suspect it to be a combination of the observers' imagination and the motion of the ship causing the substance to shift about sluggishly. Since all of the specimens taken ashore were preserved in alcohol (including all that Huxley and Haeckel ever observed), no movement was noted in the laboratory, nor was any expected.

On the Continent the first study of Bathybius to be reported was that of the Munich geologist Professor C. W. Gumbel. Huxley sent a sample of Bathybius to Giimbel and in return received a letter describing the latter's experiments. Giimbel had subjected the ooze to various acids and stains, determining to his satisfaction that he was dealing with organic matter. Within the calcareous bodies of the Coccoliths he discovered what he thought to be a type of cellulose; this led him to favor the theory that the Coccoliths, if not Bathybius itself, were plants rather than animals.

By 1870 Haeckel had completed the first of several analyses (as conjectural as they were chemical) of Bathybius. "Bathybius und das freie Protoplasm der Meerestiefen," a chapter in his "Beitrage zur Plastidentheorie," appeared in the Jenaische Zeitschrift for that year. Even in the opening paragraphs Haeckel showed little of the cautious regard for speculation which Huxley had demon- strated. The great quantity and vast extent of Bathybius over the ocean floor was, for Haeckel, a fact of far-reaching importance in all of biology:

One cannot regard closely this highly noteworthy fact without the deepest astonishment and one is involuntarily led to recall the protoplasm (Urschleim) of Oken. This universal protoplasm of the older Naturphilosophie, which originated in the sea and was to be the foundation of all life, the productive material of all organisms, this famous and notorious protoplasm whose broad meaning was already established implicitly through Max Schultze's protoplasm theory-has become a complete reality through Huxley's discovery of Bathybius.71

Haeckel went on to describe the history of Huxley's discovery and to extend his experimental results by the use of carmine stain.72 The action of the stain, plus the earlier observation by Carpenter and Thomson of what Haeckel called

69Ernst Haeckel, "Bathybius und die Monerein," Kosmos, 1877, 1:293-305 (pp. 298, 301, 303). 70C. W. Giimbel, "Vorkiufige Mittheilungen uberTiefseeschlamm," NeuesJahrbuchfiirMineralogie,

1870, 6. heft, pp. 753-767; and "The Deep-Sea Soundings and Geology," Nature, 1870, 1:657-658. Giimbel was also a supporter of Eozooin caniadense and of the related species Eozo6n bavaricum, which he had discovered himself. See his "On the Laurenitian Rocks of Bavaria," Can. Natur., 1866, N.S. 3:81-101.

7'Ernst Haeckel, "Beitrage zur Plastidentheorie," Jenia. Z. Med. Naturwiss., 1870, 5:499-519 (p. 500).

72 The carmine stain, derived from the cochineal insect, Coccus cacti, of Mexico, was used in microscopic work as early as 1770. It was probably the most highly valued stain among the early histologists of the nineteenth century. See H. J. Coinn, The History of Staining (Geneva, N.Y.: Biotechnical Publications, 1933), pp. 27-29.

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522 PHILIP F. REHBOCK

"characteristic protoplasmic movements"73 fully convinced Haeckel that Bathy- bius was a true organism. It seems likely, however, that Haeckel would have been committed to this view regardless of any experimental confirmation, because of the way Bathybius fit into his monistic philosophy.

Haeckel described his specimens in much greater detail than Huxley had. In particular he noted that the "granule-heaps" (collections of minute particles) were the predominant form in his samples and that the "gelatinous matrix" emphasized in the British accounts was relatively rare. As the granule heaps gave the proper response to stains while the matrix did not, Haeckel decided that the former were the true Bathybius and the latter merely a result of decomposition: "a plasma-product which arises from the death of Bathybius."74

Having dispensed with the drudgery of description, Haeckel then turned to the thrilling implications of the new organism: "The fact that huge masses of naked, living protoplasm cover the greater ocean depths in completely preponderant quantities and under entirely peculiar circumstances, stimulates such numerous reflections that one could write a book about them."75 The kernel of Haeckel's contribution to the Bathybius story was his certain conviction that it had originated on the ocean floor by spontaneous generation. The only other simple organism believed to inhabit that region was Globigerina, and Haeckel was convinced that "no kind of genetic connection" between the two could be demonstrated.76 Furthermore, the nourishment and reproduction of Bathybius would be very difficult to explain if one insisted that it, like other animals, subsist on organic nutrients. There was only one logical alternative for Haeckel: "Is protoplasm perhaps originating continually through spontane- ous generation? Here we stand before a series of dark questions, the answers to which can only be hoped for from subsequent researches."77

Later in the same year Haeckel amplified his position on spontaneous generation. The monistic explanation of the physical world, toward which science had been striving in the nineteenth century, entailed the hypothesis of spontane- ous generation as the only logical link between the organic and inorganic realms.78 And among all the Monera Bathybius was by far the most significant indication of the correctness of that hypothesis. Bathybius, because of its simplicity and its vast distribution, could be traced to no other source:

It must not be forgotten that, by admitting the living existence of the Monera and especially Bathybius, we find ourselves confronted by the following alternatives:

73Haeckel, "Beitraige zur Plastidentheorie," p. 517. 74 Ibid., p. 508. 75Ibid., p. 517. 76 Ibid., p. 518. 77Ibid., p. 519. 78This had been a point of emphasis in the Generelle Morphologie and was repeated here. Haeckel,

"Nachtrage zur Monographie der Moneren," Jena. Z. Naturwiss., Med. 1870-1871, 6:22-44 (p. 37). (A partial translation appears in E. R. L[ankester], "Ernst Haeckel on the Mechanical Theory of Life and Spontaneous Generation," Nature, 1871, 3:354-356.) Haeckel claimed here that spontaneous generation, or abiogenesis, "is, in fact, a necessary and integral part of the universal evolution theory. It is the natural bridge which places in continuity Kant's and Laplace's theory of the mechanical origin of the universe and the earth, with Lamarck's and Darwin's theory of the mechanical origin of animal and vegetal forms" (p. 355).

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Either the Monera were once and for all, at the beginning of organic life on the earth, produced by Archigenesis [spontaneous generation] -in which case they must have reproduced in a direct line unchanged for many millions of years; or else, in the course of the earth's history, they have been produced by recurring acts of Archigenesis, and in this case there is no reason why this process should not occur at the present time.79

Haeckel preferred the latter alternative because it seemed to present fewer theoretical difficulties.80 But in either case the Monera were a great step forward in the acceptance of the theory of spontaneous generation and a critical link in the understanding of the origin of life.8'

Huxley would not commit himself to Haeckel's theory of Bathybius' inorganic origins, although he was not opposed to abiogenesis in principle. In the lecture "Biogenesis and Abiogenesis" of 1870, he argued that the evidence pointed to biogenesis for "all known forms of life." 82 Though he felt certain that abiogenesis had not yet been accomplished in man's presence, he was much more guarded about what might have occurred in the geologic past:

. . .if it were given me to look beyond the abyss of geologically recorded time to the still more remote period when the earth was passing through physical and chemical conditions, which it can no more see again than a man can recall his infancy, I should expect to be a witness of the evolution of living protoplasm from not living matter. I should expect to see it appear under forms of great simplicity, endowed, like existing fungi, with the power of determining the formation of new protoplasm from such matters as ammonium carbonates, oxalates and tartrates, alkaline and earthy phosphates, and water, without the aid of light. That is the expectation to which analogical reasoning leads me; but I beg you once more to recollect that I have no right to call my opinion anything but an act of philosophical faith.83

The lecture on abiogenesis would have been an obvious place to discuss the significance of Bathybius had Huxley felt as confident in the matter as Haeckel did. But Bathybius was not mentioned there. Nor was it mentioned in Huxley's presidential address to the Geological Society of the same year,

79Haeckel, "Nachtrage zur Monographie der Moneren," pp. 41-42. 80This is one of several conceptions Haeckel apparently borrowed from Lamarck. See Lester

F. Ward, Haeckel's Genesis of iMan (Philadelphia: Edward Stern, 1879), p. 17. 81 Haeckel, "Nachtrage zur Monographie der Moneren," p. 42. Ward emphasized the difference

between the generation of life in a fluid containing organic matter ("plasmagonia") and generation in a medium consisting only of inorganic elements ("autogonia"). He criticized Haeckel for insisting that the Monera came about through autogonia (without the intervening stage of organic matter), which he thought "would doubtless be too great a saltus for Nature to take." Ward, Haeckel's Genesis, pp. 46-47.

82Huxley, Discourses Biological and Geological, p. 254. 83Ibid., p. 255.

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524 PHILIP F. REHBOCK

even though at that time he felt certain enough of Eozoon to proclaim himself on Dawson's side.84

Only Dawson and Carpenter would support a direct relationship between Bathybius and Eozoon. Hoping to further his case for Eozoon, Dawson published a history of its discovery, entitled The Dawn of Life, in 1875. In the description of the fossil specimens, Dawson pointed out their relationship to Bathybius:

The lowest layer of serpentine represents the first gelatinous coat of animal matter which grew upon the bottom, and which, if we could have seen it before any shell was formed upon its surface, must have resembled, in its appearance, at least, the shapeless coat of living slime found in some portions of the bed of the sea, which has received from Huxley the name Bathybius.. .85

Dawson's interest in Eozoon and Bathybius was more than just geological. Unlike the other participants in these discussions, Dawson was until his death a forceful opponent of Darwinism. When others saw Eozoon as filling an important gap in Darwin's imperfect geological record,86 Dawson insisted that the great hiatus in time and morphological complexity between Eozoon and the later Cambrian forms of life eliminated any possibility of the latter being evolutionary descen- dents of the former.87 Eozoon thus became as important to Dawson's creationism as Bathybius was to Haeckel's monism.88

By associating Bathybius with the mechanistic interpretation of life, Haeckel forced the proponents of vitalism into the fray. Lionel Beale devoted several pages to the subject in the second edition of Protoplasm (1870). Not having investigated Bathybius personally, Beale admitted he was dependent on others' comments, mainly Wallich's. The thought of global masses of abyssal protoplasm he regarded as "fanciful and improbable"; Bathybius was to be relegated to "a complex mass of slime with many foreign bodies and the debris of living

84 Huxley, "Anniversary Address of the President," Q. J. Geol. Soc., 1870, 26:xxix-lxiv. He remarked that "if the Eozoon be, as Principal Dawson and Dr. Carpenter have shown so much reason for believing, the remains of a living being, the discovery of its true nature carried life back to a period which, as Sir William Logan has observed, is as remote from that during which the Cambrian rocks were deposited, as the Cambrian epoch itself is from the tertiaries. In other words, the ascertained duration of life upon the globe was nearly doubled at a stroke" (p. xliv). But Huxley could not accept a genetic relationship between Eozooin and Bathybius, because while the former might be foraminiferal, the latter he felt definitely was not. He did admit, however, that the existence of Bathybiws lent credence to the organic nature of Eozo6n, because both seemed to have been "susceptible of apparently indefinite growth." And he thought it "possible that such organisms might have gone on living from the earliest geological times." See "Discussion" followiing King and Rowney, "On the So-called 'Eozoonal Rock,'" Q. J. Geol. Soc., 1869, 25:115-118 (p. 118).

85J. W. Dawson, The Dawn of Life (Montreal: Dawson Brothers, 1875), pp. 65-66. 86Huxley, "Anniversary Address," p. xliv. 87 Dawson, The Dawn of Life, p. 227. 88Dawson's attitude toward Haeckel's interpretation of Bathybius is clear from the following:

"Haeckel, the prophet of this new philosophy, waves his magic wand, and simple masses of sarcode spring from inorganic matter, and form diffused sheets of sea-slime, from which are in time separated distinct Amoeboid and Foraminiferal forms. Experience, however, gives us no facts whereon to build this supposition, and it remains neither more nor less scientific or certain than that old fancy of the Egyptians, which derived animals from the fertile mud of the Nile." (Ibid., p. 218.)

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BATHYBIUS HAECKELII 525

organisms which have passed away."89 Ironically, Beale's armchair approach had brought him closer to the truth than all the most careful observations of those he had read.

The British were not the only ones to acquire their own Bathybius. Haeckel's colleague Oscar Schmidt (1823-1886) published an article on his encounter with it in 1870. An expedition to the Adriatic gave him his material:

The first freshly examined sample of the bottom from 170 fathoms convinced me that I had Bathybius-mud before me. Its yellowish-grey colour and its exceedingly characteristic greasy nature were so well known to the officers that I was unanimously assured by them that this "primitive mud" predominates from the upper parts of the Adriatic Sea....90

The final deep-sea specimen in the Bathybius tradition was discovered in August 1872 by the polar explorer Emil Bessels (1847-1888), of Heidelberg. As the American North Pole Expedition aboard Polaris passed through Green- land's Smith Sound, Bessels dredged up a new genus at 90 fathoms. Though similar in consistency to Bathybius, the new "Moner" was simpler in that it lacked Coccoliths. For this reason Bessels thought it an older form than Huxley's genus, and assigned the generic name Protobathybius; the specific name, robesonii, honored George M. Robeson, then U.S. Secretary of the Navy and an important backer of the Polaris expedition.9' Though less well known than Bathybius, Protobathybius (see Fig. 2) also had a curious history. It was hailed as an important conifirmation of the existence of abyssal Monera by supporters of Bathybius, and when the latter was discredited, the former retained its reputation as a genuine organism. Rather than being disproved, Protobathybius apparently was just forgotten when beliefs in deep-sea protoplasm generally became unpopular.

With the announcement of Protobathybius in 1873, Bathybius reached the peak of its credibility. It is no surprise, therefore, that the oceanographers of H.M.S. Challenger expected to ascertain more exactly its geographic distribution during their four-year cruise of the world's oceans (1872-1876). What they found, however, was something quite different.

89Beale, Protoplasm, pp. 22-25 (see n. 12 above). 90Oscar Schmidt, "On Coccoliths and Rhabdoliths," Ann. Mag. Nat. Hist., 1872, 4th Ser.,

10:359-370 (p. 361). This is a translation by W. S. Dallas, from the Sitzungsberichte der Kaiserlichen Akademie der Wissenschaft, Wien, 1870, 62:669-682.

9'Though discovered in 1872, Protobathybius robesonii was not reported by Bessels until a year later, when the crew of the Polaris, rescued by British fishing vessels, had returned to the United States. "Report to the President of the United States of the Action of the Navy Department in the matter of the Disaster to the United States Exploring Expedition toward the North Pole, accompanied by a report of the Examination of the Rescued Party, etc.," Report of the Secretary of the Navy (Washington: G.P.O., 1873), pp. 283-628 (p. 546). See also E. Bessels, "Haeckelina gigantea: ein Protist aus der Gruppe der Monothalamien," Jena. Z. Med. Naturwiss., 1875, 9:265-279 (p. 277 n.); E. Bessels, "Lettre sur l'expedition polaire Americaine, sous les ordres du Capt. Hall," Bulletin, Societede Geographie de Paris, 1875, 9:291-299; E. Bessels, Die Amerikanische Nordpolexpedition (Leipzig: Wilhelm Engelmann, 1879), pp. 320-322; A. S. Packard, Life Histories of Animals, Including Man (New York: Henry Holt, 1876), pp. 3-4.

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526 PHILIP F. REHBOCK

I~~~~~~~~~~~~~~

Figure 2. Protobathybius robesonii. From A. S. Packard, Jr., Life Histories of Animals, Including Man (New York: Henry Holt, 1876), p. 3.

DEATH ON THE HIGH SEAS

In the autumn of 1872, as the Challenger was being readied for her oceano- graphic cruise, her scientific director, Wyville Thomson, published an account of his earlier dredging expeditions with Carpenter. The Depths of the Sea set forth the state of the pre-Challenger marine sciences and is thus a guidepost for the history of oceanography. In a review of the research and speculations on Bathybius haeckelii up to that time, Thomson revealed some ambivalence about Bathybi us' existence. He recognized that the many descriptions of it were not all in agreement. Although he saw nothing illogical about a vast abyssal Moner, he felt that much of what had been taken for Bathybius was probably a diverse organic residue of some sort, "a formless condition connected either with the growth and multiplication or with the decay-of many different things.""2 But in spite of his skepticism, Thomson still seemed to think that

92Thomson, The Depths of the Sea, p. 415.

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BATHYBIUS HAECKELII 527

a primitive creature of some kind lived in and contributed to the abyssal environment: "as no living thing, however slowly it may live, is ever perfectly at rest, but is continually acting and reacting with its surroundings, the bottom of the sea becomes like the surface of the sea and of the land-a theatre of change, performing its part in maintaining the 'balance of organic nature."'93

With respect to the Coccoliths, Thomson could be less equivocal:

. . .the balance of opinion is in favour of the view that the coccoliths are joints of a minute unicellular alga living on the sea-surface and sinking down and mixing with the sarcode of Bathybius, very probably taken into it with a purpose, for the sake of the vegetable matter they may contain, and which may afford food for the animal jelly.94

Though Thomson did not cite the source of this opinion, it was probably related to the work of the retired army surgeon and marine biologist Henry John Carter (1812(?)-1895). Carter had noticed a peculiar type of cell associated with sponges and ascidians in his microscopical researches on seashore life. In April 1870 he obtained some Atlantic deep-sea mud from Carpenter and immediately noted that the Coccoliths therein were identical to the cells he had previously examined. Noting their abundance in the alimentary canals of certain large ascidians he had been observing, Carter decided they must be plants. He then compared them to pulverized pieces of a branching calcareous alga (Melobesia calcarea) and found enough similarity to convince himself that the Coccoliths should be renamed Melobesia unicellularis.5

But in 1872 it was Bathybius, not the Coccoliths, that predominated in the minds of the Challenger scientists. Would the voyage yield new information about its nature and geographical extent? During the first three years of the voyage they looked for it eagerly each time the dredge was hauled on board. Thomson's earlier experience with the original organism and his generally skeptical attitude made it less likely that every viscous ooze brought up would

93Ibid., p. 411-412. Carpenter exhibited a similar skepticism in the 5th ed. of The Microscope and Its Revelations (London: J. & A. Churchill, 1875), pp. 466, 795.

94Thomson, The Depths of the Sea, p. 414. See also Thomson, "Preliminary Notes on the Nature of the Sea-bottom, Procured by the Soundings of H.M.S. 'Challenger' during her Cruise in the 'Southern Sea' in the Early Part of the Year 1874," Proc. Roy. Soc. Lon., 1874-75, 23:32-49 (p. 38).

95H. J. Carter, "On Melobesia unicellularis, better known as the Coccolith," Ann. Mag. Nat. Hist., 1871, 4th Ser., 7:184-189. Carter's work was not the last word on the subject, however. Though it was generally agreed by the late 1870s that the Coccoliths were of algal origin, their exact systematic position and ecological relations remained in question until the turn of the century. By then algologists had determined that Carter had chosen the correct kingdom but the wrong phylum for the Coccoliths. The branching Melobesia is a benthic red alga (Rhodophyta), whereas the Coccolith plate structure is produced by the planktonic biflagellate Coccolithophoridae, a major family of the golden-brown algae (Chrysophyta). See Gilbert W. Prescott, "History of Phycology," in G. M. Smith, Manual of Phycology (Waltham, Mass.: Chronica Botanica, 1951), pp. 1-9; G. F. Papenfuss, "Classification of the Algae," in A Century of Progress in the Natural Sciences, 1853-1953 (San Francisco: California Academy of Sciences, 1955), pp. 115-224; E. Yale Dawson, Marine Botany, an Introduction (New York: Holt, Rinehart & Winston, 1966), Ch. 16. I am indebted to Dr. Robert Fournier of Dalhousie University for correcting my interpretation of Carter's contribution.

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528 PHILIP F. REHBOCK

be labeled Bathybius, as seems to have happened with the Germans Schmidt and Bessels. And indeed, no fresh Bathybius could be found.

Finally, in the early months of 1875, as the Challenger was en route from Hong Kong to Yokohama, new events brought about the demise of Bathybius. The ship's chemist, John Young Buchanan (1844-1925), thinking that samples of bottom water must contain some portion of the organism, tried to isolate it by evaporating bottom water and heating the residue. No sign of carbonization or burning was observed.96 Meanwhile, John Murray (1841-1914), who had been responsible for data relating to the physics and geology of the ocean basins, had been preserving numerous bottom samples in alcohol, in accordance with Huxley's instructions.97 Some of these, he noted, "assumed a very mobile or jelly-like aspect."98 Buchanan described it as a "substance like 'coagulated mucus,' which answered in every particular, except the want of motion, to the description of the organism."99

Because of the failure of the evaporation and burning test, Buchanan became convinced that the substance must be inorganic. When he then analyzed the material, he found it to be calcium sulfate ("sulfate of lime"). The chemical had been precipitated out of the sea water present in the sediment by the addition of the preserving alcohol. Further experiments, described by Murray, showed that the essential ingredients were small amounts of sea water and large amounts of alcohol. When the proportion of alcohol to sea water was relatively small, the precipitate took on a crystalline form; this was presumably what Haeckel saw as "granule-heaps." Larger amounts of alcohol caused the precipitate to remain amorphous; this was Huxley's "structureless matrix." Staining of the precipitate did indeed show the appearance which Haeckel thought to be decisive proof of its organic nature.'00

The Challenger scientists were initially incredulous at this discovery, except for Buchanan, who seemed to take delight generally in debunking the theories of others.'0' Thomson wrote to Huxley on June 9, 1875, relating the new developments. The following excerpts from Thomson's letter document this crucial turning point in Bathybius' history and give a hint of the spirit in which this research was carried on.

There is another matter on which / I have some little hesitation / in writing at present because / I feel that our information / is imperfect, but yet I think

96J. Y. Buchanan, "Preliminary Report to Professor Wyville Thomson, F.R.S., Director of the Civilian Scientific Staff, on Work (Chemical and Geological) done on board H.M.S. 'Challenger,'" Proc. Roy. Soc. Lon., 1876, 24:593-626 (p. 605). See also J. Y. Buchanan, Accounts Rendered of Work Done and Things Seen (Cambridge: Cambridge University Press, 1919).

97Huxley had been a member of the committee of the Royal Society which had drawn up the instructions for Challenger's scientific staff. See Challenger Reports, Narrative, Vol. I, pt. 1, p. li.

98John Murray, "Preliminary Report to Professor Wyville Thomson, F.R.S., Director of the Civilian Scientific Staff, on Work Done on Board the 'Challenger,'" Proc. Roy. Soc. Lon., 1876, 24:471-543 (p. 530).

99Buchanan, "Preliminary Report," p. 605. ??Murray, "Preliminary Report," pp. 530-531.

101 Hugh Robert Mill, "Obituary to Mr. J. Y. Buchanan, F.R.S.," Nature, 1925, 116:719-720. See also George E. R. Deacon, "Early Scientific Studies of the Antarctic Ocean," Bulletin de l'Institut Oceanographique, Monaco, No. special 2 (1968), pp. 269-279.

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BATHYBIUS HAECKELII 529

/ as it is one which has been given / undue importance of late, and / bears upon a question in which / you are officially mixed up / you should be told exactly / how it stands. None of us / have ever been able to see / a trace of Bathybius, although / it has been looked for throughout / with the utmost care . . .Murray who / has worked at it most carefully, / and Suhm & Moseley who are / well up to the use of the microscope / all deny that such a thing exists / . I have / gone over Haeckel's papers & all / the other notes we have on / board about it and I think / so also, but I do not feel / absolutely certain yet as to / the constitution of the flocculent / precipitate. It seems to me / possible that a trace of organic / matter may combine with / the lime sulphate to give it / that very peculiar form, and / as there will certainly be a / vigorous discussion about it / I am inclined to check any / hasty publication. Murray / and Buchanan are anxious / to bring it out at once for / they expect the question to / arise at the British Association and / they would rather have the / first word. I told them, or / rather I told Murray, that I / would not sanction a paper / being sent home till I had / written to you, as you took / a fatherly interest in the beast? / and got your answer.'02

Though Thomson had not given up all hope of finding Bathybius alive, Huxley was ready to admit his mistake and call off the search, as evidenced by his immediate publication of a portion of Thomson's letter in Nature. After the letter he added:

Prof. Thomson speaks very guardedly, and does not consider the fate of Bathybius to be as yet absolutely decided. But since I am mainly responsible for the mistake, if it be one, of introducing this singular substance into the list of living things, I think I shall err on the right side in attaching even greater weight than he does to the view which he suggests.'03

Soon thereafter Huxley wrote to Michael Foster:

I have just had a long letter from Wyville Thomson. The Challenger inclines to think that Bathybims is a mineral precipitate! in which case some enemy will probably say that it is a product of my precipitation. So mind, I was the first to make that 'goak.' Old Ehrenberg suggested something of the kind to me, but I have not his letter here. I shall eat my leek handsomely if any eating has to be done.'04

LINGERING BATHYBIAL MEMORIES

A thorough exposure of Bathybius was made by Murray and Buchanan in their "Preliminary Reports" of the Challenger results, presented to the Royal

102 Huxley Papers (Imperial College), Vol. 27, MSS 315-317. I thank the Archivist, Imperial College, for permission to quote from the Huxley Papers.

103Huxley, "Notes from the Challenger," Nature, 1875, 12:315-316. Later published in Q. J. Micros. Sci., 1875, 15:392.

104 Leonard Huxley, Life and Letters, Vol. I, p. 480. Ehrenberg's attitude toward Bathybius was expressed in his "Mikrogeologische Studien iuber das kleinste Leben der Meeres-Tiefgriinde aller Zonen und dessen geologischen Einfluss," Abhandlungen der koniglichen Akademie der Wissenschaften zu Berlin, 1872, pp. 131-397 (pp. 362, 376). One letter of the Huxley-Ehrenberg correspondence on Bathybius survives in the Huxley Papers (Imperial College), Vol. 15, MS 172; Ehrenberg expressed therein his doubts about Bathybius and requested samples be sent to him soon.

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Society in 1876.105 British naturalists then fell silent on the issue until the 1879 meeting of the British Association. In the opening address-a speculative discussion of theories of living matter-the president, George Allman, resur- rected Bathybius as an example of protoplasm. He was not convinced that the Challenger staff had overturned the findings of the venerable Huxley.'06 Had the latter's discovery not been confirmed by Bessels' Protobathybius?'07

Huxley was surprised that Allman would use the highly questionable Moner as a basis-for his theorizing. In his artful reply to the address, Huxley expressed his regret at the apparent error he had initiated:

The president, in the early part of his address, alluded to a certain thing-I hardly know whether I ought to call it thing or not-of which he gave you the name Bathybius, and he stated, with perfect justice, that I had brought that thing into notice; at any rate, indeed, I christened it, and I am, in a certain sense, its earliest friend. . . I thought my young friend Bathybius would turn out a credit to me. But I am sorry to say, as time has gone on, he has not altogether verified the promise of his youth. In the first place, as the president told you, he could not be found when he was wanted; and in the second place, when he was found, all sorts of things were said about him. Indeed, I regret to be obliged to tell you that some persons of severe minds went so far as to say that he was nothing but simply a gelatinous precipitate of slime, which had carried down organic matter. If that is so, I am very sorry for it, for whoever else may have joined in this error, I am undoubtedly primarily responsible for it. But I do not know at this time of my knowledge how the matter stands... Therefore my own judgment is in an absolute state of suspension about it.108

Despite his alleged uncertainty about the final status of Bathybius, Huxley preferred (or, for lack of dredging opportunities, was forced) to let the matter rest. There is no record of his having researched it further. He did not include it among the Monera in his Manual of the Anatomy of the Invertebrated Animals (1877).109

105 Buchanan, "Preliminary Report," pp. 604-605; Murray, "Preliminary Report," pp. 529-531. See also Lord George Campbell, Log-Letters from "the Challenger" (London: Macmillan, 1876), pp. 446-448. The Challenger expedition also settled the issue of the habitat of Globigerina. Unlike many other foraminifera which are bottom dwellers, Globigeriria lives in the upper layers, an important member of the zooplankton. Huxley relented to this view in his lecture "On Some Results of the Expedition to H.M.S. Challenger" (1875). See Huxley, Discourses Biological and Geological, pp. 82-92.

106G. J. Allman, "Presidential Address to the British Association," Rep. B.A.A.S., 1879, pt. 1, pp. 1-30 (pp. 3-4). Interestingly, Allman was Thomson's immediate predecessor as Professor of Natural History at Edinburgh.

107 Ibid., pp. 3-4. Bessels did not mention whether or not he preserved his Protobathybius specimens in alcohol. Apparently he observed the substance right after removing it from the dredge. The Polaris did carry alcohol for preservation purposes: Bessels testified to the Navy Department that the ship's commanding officer had become intoxicated several times from drinking it (Report of the Secretary of the Navy, p. 541).

'08"The British Association at Sheffield," Nature, 1879, 20:405. Huxley's recantation evoked the following response from Michael Foster: "Bye the bye, you did that Bathybius business with the most beautiful grace-I wish you would sell me-a little morsel of that trick." Huxley Papers (Imperial College), Vol. 4, MS 215.

'09T. H. Huxley, A Manual of the Anatomy of the Invertebrated Animals (London: J. & A. Churchill, 1877), pp. 78-87.

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BATHYBIUS HAECKELII 531

Bathybius died more slowly in Germany, where it had been nursed by philosophic doctrine as well as scientific orthodoxy. Haeckel's principal reply to the Challenger results was an essay in Kosmos (1877) entitled "Bathybius und die Moneren." 110 Here he presented a painstaking review of the history of his Monera, of Bathybius, and of the criticisms of both. The real error committed by Bathybius investigators, he insisted, was one of premature generali- zation. Thomson, Carpenter, and Bessels had definitely observed fresh Bathybius in the North Atlantic; "' its "amoeboid movements" were proof of that. But the Challenger expedition had been unable to find it elsewhere. The obvious conclusion for Haeckel was that he and others had been wrong to assume its distribution to be worldwide, when in fact it was, like most organisms, quite limited." 12

Haeckel's criticism was especially directed, not at the Challenger scientists, of whom he spoke highly, but at Huxley, who had forsaken his own discovery in the face of limited negative evidence. After quoting Huxley's retraction in Nature, Haeckel commented:

These are the words of Professor Huxley, which stirred such concern and, according to widespread opinion, have dealt the death-blow to poor Bathybius. But the more the real parent of Bathybius shows himself inclined to give up his child as hopeless, the more I feel bound, as its godfather, to look after its rights, and if possible, to reestablish recognition for its diminished life processes."13

These comments by Haeckel were the last substantive scientific additions to the Bathybius episode, although he kept it alive in his popular writings into the 1880s."14 In England it flared up again in 1887, when the Duke

"0Haeckel, "Bathybius und die Moneren," pp. 299-300. This essay also appeared as a chapter in Haeckel's Das Protistenreich (1878), a work which I was able to examine only in the French translation by Jules Soury, Le Regne des Protistes (Paris: C. Reinwald, 1879), Ch. 5.

I"'Haeckel regarded Bathybius and Protobathybius as probably the same organism; "Bathybius und die Moneren," p. 301.

112This attitude was also taken by Edmond Perrier in Les colonies aninales et la formation des organismes (Paris: G. Masson, 1881), pp. 61-63. For other opinions, see Otto Biutschli, "Protozoa," in H. G. Bronn, Klassen und Ordnungen des Thier-reichs (Leipzig/Heidelberg: C. F. Wintersche, 1880-1882), Band 1, pp. 180-181; H. A. Nicholson, A Manual of Zoology (6th ed., Edinburgh/Lon- don: William Blackwood, 1880), p. 62.

13Haeckel, "Bathybius und die Moneren," p. 300. 114The last of Haeckel's works to support Bathybius which I was able to examine was his Pedigree

of Man and Other Essays (London: Freethought Publishing Co., 1883). English translations of his A'nthropogenie continued to mention it up to 1897. It was omitted entirely from his Systematische Phylogenie der Protisten und Pflarnzen (1894). Finally in The Wonders of Life (a translation of Die Lebenswunder, 1904, by Joseph McCabe, London: Watts and Co.), Haeckel admitted that Bathybius "seems according to the latest investigation, not to have the significance ascribed to it" (p. 214). Nevertheless, this had no effect on the importance of his Monera and archegony. Although the previous twenty years of research had reduced his non-nucleated Monera to only two types, the Chromacea (blue-green algae) and bacteria, their position in Haeckel's philosophy was no less central. In the meantime his theory of archegony had become more sophisticated (ibid., pp. 355-356, 369). Interestingly, after brief disfavor the Monera concept has undergone a revival of popularity in the twentieth century. See Edmund B. Wilson, The Cell in Development and Inheritance (3rd ed., New York: Macmillan, 1925), pp. 24-25; Herbert F. Copeland, "The Kingdomof Organisms," Quarterly Review of Biology, 1938, 13:383-420; R. H. Whittaker, "New Concepts of Kingdoms of Organisms," Science, 1969, 163:150-159; Lynn Margulis, Origin of Eukaryotic Cells (New Haven: Yale University Press, 1970).

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532 PHILIP F. REHBOCK

of Argyll found it a convenient subject for satirizing the dogmatic side of science, which for him included the followers of Darwin and Huxley:

A fine new Greek name was devised for this mother slime, and it was christened 'Bathybius,' from the consecrated deeps in which it lay. The conception ran like wildfire through the popular literature of science, and here again there was something like a coming plebiscite in its favour. . . The naturalists of the 'Challenger' began their voyage in the full Bathybius faith. But the sturdy mind of Mr. John Murray kept its balance. . . The laboratory in Jermyn Street [where Huxley worked] was its unfailing source, and the great observer there was its only sponsor. The ocean never yielded it until it had been bottled."5

Huxley responded immediately. He freely admitted responsibility for the initial description and naming of Bathybius but disavowed quite legitimately any role in the subsequent claims for its antiquity and evolutionary importance. "That which interested me in the matter was the apparent analogy of Bathybius with other well-known forms of lower life. . . . Speculative hopes or fears had nothing to do with the matter." 116

Huxley's folly was utilized, with similar intent, by William Mallock, the writer and theologian, in 1890. Mallock's article solicited a delightfully typical response from the "bishop-slayer," who was by then feeling some exasperation. This reply is the last recorded event in the history of Bathybius in which its creator took part:

Bathybius is far too convenient a stick to beat this dog with to be ever given up, however many lies may be needful to make the weapon effectual.

I told the whole story in my reply to the Duke of Argyll, but of course the pack give tongue just as loudly as ever. Clerically-minded people cannot be accurate, even the liberals." 7

Although there are no corporeal remains of Bathybius, the episode left behind one reminder, an addition to the English language:

Bathybial (baJp i - bial) a. [fr. Bathybius + -al] Of or pertaining to bathybius or the depths at which it is found; belonging to or living in the deepest parts of the sea. 118

But more important than this etymological novelty are the conclusions which may be drawn from the case. That Bathybius was an error, and an embarrassing one, there can be no doubt. The stimulus it gave to dredging and the analysis

15 George Douglas Campbell, 8th Duke of Argyll, "A Great Lesson," Nineteenth Century, 1887, 22:293-309 (p. 308).

'16Huxley, "Science and the Bishops," Nineteenth Century, 1887, 22:625-641 (p. 638). "1 Leonard Huxley, Life and Letters, Vol. II, p. 171. "1 Oxford English Dictionary, 1971, supplement, p. 64.

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BATHYBIUS HAECKELII 533

of marine sediments was significant, "9 but that could not prevent its downfall. It was not an error of accuracy or of calculation, but rather an error of interpretation. Bathybius was a highly functional concept, an explanatory device which made sense in the context of mid-nineteenth-century biological and geological thinking. Independent but concurrent developments in cytology, protozoology, submarine geology, Precambrian palaeontology, microscopy, and evolutionary biology converged to produce an intellectual environment congenial to Bathybius in the 1860s. That environment had not existed a decade before; it ceased to exist a decade later. However "peculiar and fantastic" it might now appear, Bathybius was a rational construction, a snugly fitting piece in the intricate puzzle of Victorian science. There must have been many who, by the end of the century, could agree with the comment in Nature that "Bathybius is dead, but one cannot leave it without the reflection that there are few naturalists, the young and expert included, but would have given similar exDlanation of the aDDearances." 120

'19Even greater credit is afforded Bathybius by Pierre-Paul Grass'e ed., Traite de zoologie (Paris: Mosson, 1952), Tome 1, fas. 1, p. 37.

'20George Murray and V. H. Blackman, "Coccospheres and Rhabdospheres," Nature, 1897, 55:510-511.

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