RESEARCH ARTICLE

Controlling and Culturing Diversity: Experimental Zoology Before World War II and Vienna's Biologische Versuchsanstalt CHERYL A. LOGAN1* AND SABINE BRAUCKMANN2 1

UNCG, Department of Psychology and History, Greensboro, North Carolina, USA 27402 University of Tartu, Teaduskeskus, Tartu, Estonia 50091

2

ABSTRACT

Founded in Vienna in 1903, the Institute for Experimental Biology pioneered the application of experimental methods to living organisms maintained for sustained periods in captivity. Its Director, the zoologist Hans Przibram, oversaw until 1938, the attempt to integrate ontogeny with studies of inheritance using precise and controlled measurements of the impact of environmental influences on the emergence of form and function. In the early years, these efforts paralleled and even fostered the emergence of experimental biology in America. But fate intervened. Though the Institute served an international community, most of its resident scientists and staff were of Jewish ancestry. Well before the Nazis entered Austria in 1938, these men and women were being fired and driven out; some, including Przibram, were eventually killed. We describe the unprecedented facilities built and the topics addressed by the several departments that made up this Institute, stressing those most relevant to the establishment and success of the Journal of Experimental Zoology, which was founded just a year later. The Institute's diaspora left an important legacy in North America, perhaps best embodied by the career of the developmental neuroscientist Paul Weiss. J. Exp. Zool. 323A:211–226, 2015. © 2015 Wiley Periodicals, Inc.

J. Exp. Zool. 323A:211–226, 2015

How to cite this article: Logan CA,Brauckmann S. 2015. Controlling and culturing diversity: Experimental zoology before World War II and Vienna's Biologische Versuchsanstalt. J. Exp. Zool. 323A:211–226.

In an attempt to destroy Hitler's Germany, the US Air Force bombed Vienna in February and March of 1945, as the Soviet Red Army took the city. Most bombing targets were oil refineries, bridges, troop marshaling yards, and railroads; but the bombing and artillery shelling badly damaged much of the once imperial city. One important nineteenth century Renaissance-style



Correspondence to: Cheryl A. Logan, UNCG, Departments of Psychology and History, Greensboro, NC 27402. E-mail: [email protected] Received 26 November 2014; Revised 7 January 2015; Accepted 8 January 2015 DOI: 10.1002/jez.1915 Published online in Wiley Online Library (wileyonlinelibrary.com).

building, which, near the end of the War, was used to house the German army, was destroyed by incendiary bombs on April 11, 1945 (Vivarium papers, Archives of the Austrian Academy of Sciences; Taschwer, 2014). The building had been originally been built as an animal show house for the 1873 Vienna World's fair; then it became a reptile house; and finally, a showplace for “exotic” humans and their animals (Reiter, '99). But as the public lost interest, the building's fortunes and focus changed radically at the turn of the 20th century (Fig. 1). After 1900, it was transformed into a pioneering scientific laboratory equipped with unprecedented facilities for the experimental control of live organisms. In 1903, the Viennaborn Zoologist Hans Przibram, along with two other scientists, the plant physiologist Wilhelm Figdor and the botanist Leopold von Portheim, re-opened the renovated space as a scientific

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Figure 1. Die Biologische Versuchsanstalt, ca. 1910, from Fritz Knoll Papers 1/2, Postkartensammlung, Archives of the Austrian Academy of Sciences. Reprinted with permission from the Austrian Academy of Sciences.

institute. Their wealthy Viennese families, long devoted to science, provided the necessary funds. They called their new institution the Institute for Experimental Biology (die Biologische Versuchsanstalt, the BVA). The three men owned it until 1914, when Przibram and von Portheim donated it to the Austrian Academy of Sciences. Until the outbreak of World War II, the BVA drew many international visitors, its staff scientists collaborated with faculty and students from the University of Vienna, and it was governed by scientific commissions whose members included such prominent biologists as the pioneering physiologist Sigmund Exner, a student of Hermann Helmholtz and Ernst Br€ ucke, as well as the zoologist Bernhard Hatschek and the botanists Julius Wiesner and Richard von Wettstein, for whom the Wettstein system of plant phylogeny is named. The founding of marine research stations, notably Concarneau opened by Victor Coste in 1859, initiated the modernizing trend in biological research (De Bont, 2009). A decade later, the construction of international laboratories that regularly hosted visiting scientists for experimental research began with the founding in 1872 of the Naples Zoological Station (Stazione Zoologica di Napoli, SZN) by the German evolutionary zoologist Anton Dohrn. Like many Germans, Dohrn was especially taken with Darwin's new theory of evolution, and, using annelids, he formulated the theory of functional change (Breidbach, 2007; Dohrn, '75). Dohrn also valued naturalistic studies of animal habits and “ways of living” (De Bont, 2009). To integrate these with experimental analyses of evolution and with experimental zoology, Dohrn invented the “table system,” which allowed scientific institutions, universities, and governments from around the world to rent bench space and send at least one scientist per year to do experiments at the SZN. These scientists enjoyed access to modern facilities and fresh organisms for experiments, as well as exposure to the German-speaking scientists who were then revolutionizing the life sciences, especially in embryology, cell J. Exp. Zool.

LOGAN AND BRAUCKMANN theory, and neurophysiology (M€ uller, '96; Groeben, 2002; Groeben and De Sio, 2006).1 The system quickly shaped an international community of zoologists, of which the emerging American group was an important part. Americans, such as Thomas Hunt Morgan, Edmund Wilson, Charles Otis Whitman, and Charles Davenport, regularly traveled to Naples for research, as ideas and methods spawned with German inspiration spread and were transformed by concepts and techniques developed in their own institutions. Morgan and Whitman's experience, along with the impetus of the Boston Society of Natural History and the Women's Education Association, led them to transform a summer teaching lab established for Boston teachers into a modern opportunity for marine research for Americans who could not travel to Naples. The result was the founding in 1888 in Woods Hole, Massachusetts, of the Marine Biology Station (Maienschein, '85, 2007; Benson, '88; Pauly, '88). Non-marine laboratories for experimentation on live animals followed soon thereafter. The BVA opened in 1903, just as the laboratory trend also flourished in the United States. In 1904, with funds from the Carnegie Institution, Charles Davenport opened the Station for Experimental Evolution at Cold Spring Harbor, New York, a facility founded as a marine biology teaching school for high school boys; and in 1906, the Wistar Institute in Philadelphia completed its transition from an anatomy museum to an experimental laboratory when they hired the University of Chicago neurologist Henry Donaldson as the Institute's new scientific director (Clause, '93). Just a year later, at the Seventh International Zoology Congress in Boston, the pioneering British geneticist William Bateson announced: “With their base on Cold Spring Harbor, or Woods Hole and the Biologische Versuchsanstalt in Vienna, the allied armies of genetics, cytology and experimental zoology start for the grand attack; and I think when we meet at the end of another period of ten years, there will be victories to record (Bateson, '12).” Though ironically, Bateson's military metaphor badly misjudged what that decade would bring, the power and promise of the hope is clear. Each of these early laboratories transformed the biological sciences. Yet, because its vision for an organismic approach that was nonetheless fused with experimentation and its fate amid the destruction brought by Fascism differed so from the others, the distinct impact of the BVA is only recently being revealed. In what follows we summarize the philosophy, scientific facilities, and key accomplishments of the BVA scientists to present a picture of the Institute's role in promoting the field of experimental zoology. We then trace its significance through concepts and

1 Fantini (2000, p. 526) writes that by 1909, “2,200 scientists from Europe and the United States had worked at Naples and more than 50 tables per year had been rented out.“

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Figure 2. Floor plan of the BVA showing the Department of Botany and Plant Physiology with terraria, dark chambers, a laboratory for moderate temperatures, warm rooms with an aviary, a thermal laboratory, as well as thermal and cold chambers with skylights, and the Department of Zoology with a chemical laboratory, dark rooms for photography, freshwater chambers, weighing-chambers, chemical preparation areas, thermal and cold greenhouses, and many breeding cages (Austrian State Archive, AVA, U-Allg., Ktn. 128, Zl. 48.551/1905 Beilage 6). Reprinted with permission from the Austrian State Archive, Unterrichtsministerium-Allgemein.

individuals whose influence persisted well after the people who shaped it and the building itself were gone. Most of those people were Jewish. Hans Przibram had directed the BVA from its construction in 1902 until he and all other Jewish scientists and staff were finally forced out in March 1938, when Austrian Nazis took over the administration of the Institute. Thereafter, very little scientific work was done, and in 1943, Przibram and his wife were imprisoned in the Theresienstadt concentration camp, where Przibram died in 1944 (Taschwer, 2014). Until the Nazis removed him from the Directorship of the BVA, Przibram had been in close contact with the many changes taking place in American biology in the early 20th century. He noted in the Forward to the first volume of his seven volume series entitled Experimental-Zoologie, that it was no longer appropriate to simply name his series the “second edition” of his earlier work, Einleitung in der experimentelle Morphologie der Tiere [Introduction to the experimental morphology of animals], as he had planned. Instead, Przibram wrote: “ . . . in a detailed

treatment, the separation of morphological and physiological experiments proved itself to be impossible, and therefore the more apt title of the Americans, which they coined for their new journal, was preferred: “Experimental Zoology” (Przibram, 1907, p x).” Research done by pioneering Americans, such as Morgan, Davenport, Wilson, Ross Harrison, and Frank Lillie, was well represented in several volumes of his series. In fact, in Volume 1, only two journals were mentioned so often that their titles were abbreviated for efficiency: They were Roux Archiv f€ ur Entwicklungsmechanik der Organismen, in which a great deal of BVA research was published, and America's Journal of Experimental Zoology. It is, therefore, fitting that, on the occasion of the Journal's 110th anniversary, we introduce American biologists to this pioneering institution. The BVA Founding Przibram announced the opening of the new institute in a short paper published in 1903. He began by stressing the importance of J. Exp. Zool.

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Figure 3. Diagram of the seawater and ventilation facilities of the BVA (from Przibram, '08c, Fig. 1, p 11). Reproduced with the generous permission of the family of Hans Przibram.

a strong reliance on experimentation. Unlike many other scientific institutions of the time, the BVA would give clear methodological priority to controlled experiments. Though other methods were not to be excluded, the goal was not to describe or catalog life, but to determine its causes; and the causes of the emergence of form and function would have to be established experimentally. The institute would especially serve, he wrote, “the newest branches of biology, experimental morphology and developmental physiology as well as the related areas of biochemistry and biophysics (1903, p 153).” Physical Facilities. To illustrate, Przibram briefly described the exacting character of the new facilities, then under construction (see Fig. 2). All of the rooms in which animals or plants would be housed or bred had steam controlled central heating, and half of the entire building was devoted to experimental rooms in which high constant temperatures of up to 45 °C could be maintained. Most rooms had outlets for both fresh and salt water, and distilled water was readily available. The seawater facility (see Fig. 3) consisted of eight stone basins, the largest of which was nearly 3 m high by 2 m wide, which circle-like surrounded the glass wall of the building's front. To fill the facility, 1,514.2 liters of seawater were transported by train from the Marine Station in Trieste. A ventilation device primed the air by a sucking port from the outside and compressed it into cylindrical steel kettles to a pressure of 5 at, which then distributed seawater through pipes J. Exp. Zool.

running throughout the building. In the garden, four cavities of 4.5 m  3 m  1.2 m were dug, which functioned as terraria; in addition, there were a large basin and four smaller ones for algae and “higher” water plants. In the basement, three ventilation chambers were equipped for experiments with darkness and constant temperatures, and all rooms in the Institute had electrical outlets. For advanced vertebrates, an extension was later built that added 15 glass cages, and two greenhouses were built for plants. The addition totaled over 56 m2 that was added to the south side of the building (see Fig. 2). With such control, Przibram stressed, scientists could achieve one key goal of the institute: to establish the role of external factors—temperature, lighting, an environmental medium, etc.,—on changes in organic form, as well as on the persistence of those changes in descendents. Przibram closed by inviting all of the assembled scientists to apply for research space.2 It was an invitation to participate in a new approach to understanding the characteristics of life. Przibram's mathematical inclinations placed a premium on precision. So, the BVA also excelled in its wealth and abundance of equipment and measuring instruments. An important addition, original to the BVA compared with biological institutes at the universities, was the chemical laboratory, which was equipped 2 Przibram's announcement was made at the 1902 meeting of the Society for German Natural Scientists and Physicians.

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Figure 4. Motor-driven clinostat after Figdor and Portheim (from Przibram, '08a, Fig. 6, p 260). Reproduced with the generous permission of the family of Hans Przibram.

with heliostats, a nephelometer, and Wilhelm Ostwald's viscometer. In 1907, this lab became the Department of Physical Chemistry. Wolfgang Pauli (father of the Nobel Prize winning physicist of the same name), who experimented with colloids (e.g., Pauli, '07), directed the new Department. For assessing the impact of gravitation on the growth of plants, Wilhelm Figdor and Leopold von Portheim, directors of the Department of Botany and Plant Physiology, improved the clinostat originally devised by Julius Sachs (and refined by Julius Wiesner) with an electrical motor (see Fig. 4). With a clinostat, a plant that is fixed horizontally can be slowly turned on its vertical axis so that its shoots and roots are uniformly exposed on all sides of the rotational plane to the force of gravity. Thereby, a unilateral effect of gravitation was eliminated, and biased geotropic growth did not occur. The device was crucial for Figdor's experiments on the impact of light and gravitation (Figdor, '09; see Nickelsen, 2015). Fauna-Culture in the “Vivarium”. Przibram's explicit stress on biochemistry and biophysics illustrates the priority placed on the analysis of physical and chemical causes. But like the SZN, the BVA also aimed to develop experimental accounts of evolution, thus combining analyses of physical input with the complexities of organic change in ontogeny and in speciation. Attention to descent and evolution would require the successful breeding of a

wide range of species, a BVA goal that was unprecedented even among the other institutes then stressing experimentation. The Viennese citizens, who had witnessed its many incarnations popularly named the building housing the BVA the “Vivarium.” The title accurately echoes one of the institution's unique strengths. The marine stations had offered a wide range of fresh test organisms for research. Visiting scientists flocked to the SZN in part for its ready access to diverse marine organisms taken from the sea and brought directly from sorting rooms into the laboratories. But in the United States, an early emphasis on cellular and molecular problems and the increasing use, beginning around 1920, of a restricted range of organisms, such as Drosophila and inbred rat and mouse lines, led to a de-emphasis on biological diversity and increased focus on a limited number of “model” organisms (Harwood, '93; Kohler, '94; Logan, 2002; Rader, 2004). At the BVA, by contrast, the range of species available for research would rival that of the bay of Naples and the Atlantic coast of Massachusetts. Further, the BVA's integrative systems approach stressed both natural and novel adaptive outcomes, alongside the exploration of cellular and physiological mechanisms. To produce this range of adaptations, “exemplars” of many species were nurtured over their entire life histories, often in conditions that enabled them to breed regularly. In his 1913 progress report on research done at J. Exp. Zool.

216 the BVA, Przibram (1913) included a catalog of all organisms being maintained and bred for research there. Each species was listed along with its housing conditions, housing population density, feeding conditions, and the location from which the specimens had been collected. Asterisks designated species that were being successfully bred. An earlier paper had stressed invertebrates, so this version only included mollusks and tunicates among the invertebrates along side a wide range of vertebrates. Not including subspecies, 358 species were listed, of which 120 (33%) were being bred for research. All five classes of vertebrates were represented, and various species of all five were being bred, though most breeding involved reptiles and amphibians. So, along with breeding populations of snails, slugs, ascidians, salamanders, frogs, toads, lizards, and snakes, visiting researchers could work on replenishing populations of lamprey, poeciliids, perch, daces, sculpins, linnets, canaries, squirrels, guinea pigs, mice, or even a hybrid of domestic and wild cats. It was as if visitors doing research at the BVA were being offered the natural diversity of a kind of terrestrial Bay of Naples, but with access to facilities to precisely control the conditions that prevailed in experiments conducted over extended periods in the lives of the animals. The BVA zoologist Paul Kammerer had compiled the catalog of species included in Przibram's paper. Kammerer was a master at nurturing, culturing, and breeding animals, particularly amphibians and reptiles. Przibram had hired Kammerer at the laboratory's founding to maintain and produce these diverse animal colonies. In the United States, Kammerer is likely remembered by the scandal surrounding the so-called midwife toad controversy. This was an episode of acknowledged scientific fraud, which, following his suicide in 1926, destroyed Kammerer's scientific reputation.3 But his important role in establishing a wide-ranging fauna-culture for experimental zoology is rarely discussed. Debates about the scandal almost never mention the research done by Kammerer before he joined the BVA, which involved a large community of aquarium and terrarium enthusiasts. The inclusion of this aspect of Kammerer's career paints a somewhat different picture of the man's contribution to biological science. Berz and Taschwer (2010) suggest that Kammerer's stress on environmental control emerged though his youthful experience with an elaborate European network of aquarium and terrarium enthusiasts. He had published hundreds of studies on nurturing and breeding species of amphibians that both addressed the great evolutionary transition from water to land and provided that

3 Partly because the specimens were produced in 1909, but the fraud not detected until 1926, there remains much debate about Kammerer's guilt or innocence. Arthur Koestler defended Kammerer in his 1971 book, The Case of the Midwife Toad. Recent analyses reveal important issues unaddressed by Koestler; but several historians find it unlikely that Kammerer committed the fraud (see Hirschm€ uller, '91; Gliboff, 2006; Berz and Taschwer, 2010).

J. Exp. Zool.

LOGAN AND BRAUCKMANN network with vital practical information. And he brought that expertise to the BVA and to Przibram's methodological vision. Seen in the context of the BVA's innovative effort to maintain diverse materials for chronic experiments of the kind Ivan Pavlov pioneered in digestive physiology (see Todes, 2002), Kammerer's skills in animal husbandry were as important to the BVA as the work later found to be fraudulent. They made possible chronic controlled experiments, which permitted animals to be explored throughout their lives in ways that included microecologies inducing the development of novel attributes: today's neophenogenesis (see Johnston and Gottlieb, '90). Similar short-term experiments had been done on the early development of cell lineage complexes in marine invertebrates at Naples and Woods Hole. But in land-locked laboratories with terrestrial vertebrates much more was involved. Przibram's catalog of species included references for each of the species listed, citing the source of the information. Many of these citations reference Kammerer's early work, which later facilitated the maintenance of a comparative perspective manifest in the BVA's many breeding colonies of animals maintained before the First World War. Research Themes Przibram's systems perspective is well reflected in Volume 5 of his series on experimental zoology, which was entitled Function [Funktion (Verrichtung)] and subtitled Practice, Interrelationship, Adaptation [Aus€ ubung, Wechselwirkungen, Anpassung]. In it, Przibram described his understanding of the meaning of biological function, which he defined as the performance of activities that sustain organs, organisms, species, or ecosystems. He distinguished three groups of activities that ensured such performance: sensation, movement, and nourishment (in the last he included internal secretions and the development of form). The categories need not engage one another in all life forms, but in most animals they were usually “linked in such a way that an effective factor called forth specific reactions in each of the three groups (1914, p 2).” In addition to these coordinated functional complexes sustaining life internally (through neural, secretory, and compensatory processes), individuals must perform functions that adapt them to the changing conditions of the external world. Closely following Wilhelm Roux (see below), he termed them “functional adaptations.” These were his focus; they represented ontogenetic adaptations that were triggered by adjustments to the reception of physical energy—short wave electromagnetic stimulation, such as light, mechanical wave patterns, including those producing sound, and chemicals affecting the molecular constitution of the body. His nine chapters elaborate the action of such external simulation, using the notions of reception, in the sense of the detection of physical energy, and that of correlation. In this and earlier work, Przibram ('08b, '10a) outlined the union of analysis and functional integration defining the explanatory frame for much research done at the BVA, particularly the many experiments on organ transplantation and tissue regeneration.

EXPERIMENTAL ZOOLOGY IN VIENNA Roux's theory of functional adaptation was based on his definition of epigenesis as a developmental integration, or synthesis, of the great variety of organic shapes originating both in the forces of a substrate (Weismann's germ-plasm) and in the diversity of possible forms (Roux, 1885). The issue he wanted to address was which mechanisms control, or at least guide, morphogenesis by acting on the developing organism as if by reacting to tensions and tractions. For Roux, such mechanisms represented an organism's potential to regulate itself and maintain physiological function and morphological form over generations, regardless of internal or external disturbances (Roux, 1885, p 414–415, 423; Roux, '12, p 16– 19). For example, gastrulation is the self-differentiation of segments of the germ-plasm (epigenesis), which then proceeds to form an embryo that began as a mosaic of parts (preformation). Both processes manifest an organism's capacity to adapt to internal and external “stimuli,” as Roux phrased it, and to trigger individual ontogenetic development. To frame this capacity, Roux introduced the concept of the “functional adaptation” of organisms as a fundamental property of life. He later included the concept among his categories of organismic self-regulation. Roux's theory represented substantial progress compared to the, sometimes tedious, “descriptive morphology” of the 19th century. Much BVA research also emphasized developmental plasticity that could be revealed experimentally by nurturing adaptive novelty. This emphasis led to the exploration of both ontogenetic plasticity and plasticity in heredity—the inheritance of acquired characteristics, an idea of great interest that was already demonstrated experimentally in several groups of plants and animals (see Przibram, '10b). In the first of a series of studies published under the title “The environment of the germ-plasm [Die Umwelt des Keimplasmas],” Przibram ('12) wrote that the question of heritable acquisition had reached a new point in its scientific development. Research on most major groups of animals and plants had shown that environmental effects first apparent in parents could indeed persist in their unexposed offspring. But Przibram was skeptical. The issue, he stressed, was by what mechanism did this occur? Was the germ-plasm (the genes) largely insulated from the effects of the environment, or could changes in the soma initiated by the physical environment be transferred to the germ cells by some cellular process of “somatic induction” (see Logan, 2013)? August Weismann had famously rejected the latter idea as a matter of principle. However, Przibram's belief in ontogenetic plasticity shaped by changes in the physical environment made plausible its extension to genes, and, therefore, to the next generation. Stressing Roux's epigenetic functional adaptation and citing Richard Semon's theory of the regulatory role of internal traces of stimulation during development, Przibram formulated a research program that guided the BVA into the 1920s. The program centered around three crucial issues: (1) Determining the physical

217 conditions to which the gonads (and, therefore, heredity) are exposed inside the body; (2) exploring the modifications that occur in these conditions when external environmental factors change; and (3) correlating the changes in the gonads with those in the rest of the body, the internal (germinal) world and its microenvironment [Binnenwelt] (Przibram, '12). Experiments that so varied external stimulation reaching the genes could reveal the proposed processes of somatic induction. Roux and Semon had brought considerable evidence to bear on the problem, but definitive tests of somatic induction were rare. Przibram's program sought to settle the issue—to establish experimentally if and how it occurred. Semon's theory proposed physiological processes involving residues or traces of prior events by which the concept of selfregulation could be expanded and tested experimentally. But much of his evidence was retrospective, and even before the war, Przibram stressed examples that lead him to accept an effect of the physical environment on individual development, yet reject its extension to descendents. Even after Semon revised his theory, Przibram was still wary: “For the present we must remain undecided about which way to adjudicate the inheritance of acquired characters and be content with acknowledging the role of external factors, thereby we grant them not merely selective power (Przibram, 1910, p 211).” External factors, he believed, acted beyond just the selective process stressed by “neoDarwinists” (the label then describing those following Weismann); they also guided morphogenesis, Przibram's key focus. World War I virtually stopped this and other research at the BVA. As resources were diverted to the war effort, the BVA was closed from 1915 to 1916, and several scientists went to fight; after it reopened in 1917, the facility was used as a war hospital. Following the war, war reparations imposed on the former Axis countries had devastating economic consequences in Austria, as the Austro-Hungarian Empire was dissolved and “Austria” became a small, resource-poor state. As a result of these and a number of other changes, Przibram mentioned the inheritance of acquired characteristics less and less after the War. By the early 1920s, Kammerer's experimental work, on which Semon and other supporters had relied, was already controversial, and his suicide in 1926, badly discredited the concept. In this context, Przibram became less focused on whether important influences altering ontogeny in individuals could also transform their progeny; he instead addressed what he saw as the needed first step: uncovering mechanisms of plasticity in ontogenesis (see below). The BVA Departments The Institute had initially been conceived with three departments: Zoology, Botany and Plant Physiology, and Physical Chemistry. The Physical Chemistry Department was closed in 1914, and in 1913, an Animal Physiology Department was added, directed by the pioneering Austrian comparative physiologist Eugen J. Exp. Zool.

218 Steinach. In what follows, we summarize their research foci, concentrating on the two departments most relevant to the growth of the discipline of zoology.4 Zoology. With more than 70 scientists, nearly 100 students trained in doing experiments or working on their PhDs, many visiting international scientists (e.g., Oskar Carlgren, Davenport, Jacques Loeb), and over 600 publications, the Zoology Department was the BVA's largest department. Its research program closely followed the course of an “experimental morphology” similar to that proposed by Davenport (1897, 1899) and entailed in the functionalist physiological method advanced by Roux, 1885.5 Davenport had defined experimental morphology as the biological discipline that deals with how chemical and physical agents effect organic development. His program listed eight categories of external factors (chemical substances, density, molar agents, gravity, electricity, moisture, light, and heat) that experimental biology must aim at to isolate. Przibram adopted these categories to formulate his Department's twofold research program. Starting with the issue of ontogenetic adaptation—a fine-tuning of Roux's functional adaption, which Przibram subsumed under the “physiology of adaptation” (Przibram, '21, p 394),—he stressed morphogenetic relationships between development and the environment to frame an approach to which among the many acquired characters could be inherited and which could not. The other theme concentrated on the new method of the “physiology of morphogenesis” (Przibram, '10a) to examine growth processes and, in particular, regeneration. Paul Weiss, then working at the Zoology Department, later renamed and modified this field “morphodynamics,” a term that combines the physiological method with the concept of a morphogenetic field (Weiss, '26). At the turn of the 20th century, Viennese science had been steeped in efforts to modernize the idea of the inheritance of acquired characteristics (see Kassowitz, '02). And to reconcile the possibility of the inheritance of acquired characters with Darwin's selection theory, Przibram ('10b) called for a workable definition of the concept of “species” and for experiments investigating how the characteristics of a species are maintained when external conditions are changed. His goal was to render the process of speciation accessible to experimental analysis in a program of “experimental phylogenesis.” But, after the War, though a few scientists continued to address the problem, Przibram remained skeptical about plasticity in heredity, and 4 For further information on the Department of Botany and Plant Physiology, see K€arin Nickelsen (2015). 5 We limit our discussion to the experimental program, deemphasizing Przibram's focus on applying mathematical tools to biological issues, e.g., to chromosome numbers, the velocity of regeneration processes, sex determination, variation and selection (see Przibram, '08c, '23); for an example of this approach, see Karl Przibram (1913), the physicist and brother of Hans, on probability and the movements of paramecium.

J. Exp. Zool.

LOGAN AND BRAUCKMANN most experiments instead exaggerated the external influences impinging upon organisms to isolate conditions that could alter ontogeny. Using the same factors proposed by Davenport 15 years earlier, which Przibram had also used to structure his textbooks on Embryogenese (1907a) and Phylogenese (1910a), the Department's zoologists, for example, explored the impact of chemical factors on the coloration of insects (Przibram, '19), amphibians, and frogs (Kammerer, '13; Przibram, '22), of changes in light on butterfly wings (Weiss, '25) or their pupation (Brecher, '22), and the effects of exaggerated temperature conditions (Bierens de Haan, '22a, b; Congdon, '12). But World War I and its aftermath had a devastating effect on the progress of experimental biology in Vienna. Not only did research cease temporarily, but funds for electricity and heat were not available again until 1926, by which time the budget had been greatly reduced. Przibram's program, conceived before the war in better financial times, was no longer financially feasible. This situation was aggravated by his increasing dissatisfaction with Paul Kammerer's behavior. Before the war, Kammerer's knowledge of breeding techniques in vertebrates had been key to the inclusion of evolution and phylogeny in the BVA program. But during the war, the Directors of the Institute increasingly felt that he was no longer reliable. The situation worsened in the next few years. In May 1921, Przibram informed the BVA Council that Kammerer's behavior was not acceptable anymore and proposed that the BVA hire another scientist as assistant (see the Vivarium papers, Archives of the Austrian Academy of Sciences). But Kammerer's job was not filled until January 1925, when Weiss took the position, but without an institutional requirement to oversee animal husbandry. The breeding program was severely cut back to include only those animals (butterflies, locusts, rats, and mice) being used in Przibram's research. The war also changed the status of Central European science, especially in Austria, which was suddenly stripped of its empire. Central Europe had been the acknowledged leader of the mid- and late 19th century modernization of biology; but World War I helped shift the focus of scientific progress away from that region and to the now wealthy United States. With the rise of Mendelism and Morgan's school of transmission genetics, US experimental biologists became much less interested in the idea of the inheritance of acquired characteristics; the concept moved out of the scientific mainstream—no longer an international priority (Allen, '78; Harwood, '84, '93). In this context, Przibram reframed the BVA's institutional priorities. Probably not wishing to appear outdated and increasingly willing to distance himself from the controversial Kammerer, by about 1918, Przibram refocused the Zoology Department on exploring the impact of external environmental factors on individual ontogeny and abandoned experimental assessments of phylogeny and evolution. Conditions had imposed a less ambitious and more feasible program, which stressed plasticity in the ontogenetic development of animal form —an interest that had shaped Przibram's entire career.

EXPERIMENTAL ZOOLOGY IN VIENNA Brecher, Dembowski, and Przibram, for example, explored the impact of chemical factors on the mechanisms underlying color adaptations in butterfly larvae (Vanessa Io, V. urticae, P. atalanta). Their findings led them to provisionally conclude that during ontogenesis animal coloration adapts to the colors of the environment through light sensitive ferments (Bateson, '13; Przibram et al., '21) and absorptive light-sensitive chromogenes. Although they could not yet explain the chemical mechanism, the authors demonstrated that during the transition from cell expansion to cell division, the phases of labile color modifications and fixed color adaptation alternated with one another. Brecher's work on cabbage butterflies showed that exposing the pupa to lights of different wavelengths correlates with the final coloration of the larvae, implicating the effect of external light in a chain of factors internal to the caterpillar that mediated sensitivity to light. Brecher demonstrated a color- and reactionsensitive tyrosinase and a specific modification of the organism's reaction during metamorphosis from caterpillar to pupa. She interpreted this as an adaptive buffering against [“Schutz gegen”] the intensity of light (Brecher, '22, p 274). Brecher had demonstrated a cellular process underlying the adaptive ontogenetic induction of color modification. Edgar Congdon, a collaborator from Harvard University, studied the impact of temperatures ranging from 10 to 15 °C higher or lower than medium body temperature on the gonads and fetuses of mice (M. decumanus, M. musculus) and hibernating dormice (Myoxus glis). Francis B. Sumner, who supported Przibram ('17b) by providing live square crabs (Gelasimus pugnax Smith) sent from Woods Hole to Vienna in moist seaweed, was rather critical of Congdon's experimental series. He argued that the latter had used “inadequate methods of recording the internal temperature of the body (Sumner, '15, p 330).” The BVA's chambers equipped with constant temperatures were not yet available for Congdon's experiments. So Przibram repeated the work using measures of body temperature in mice and rats to more precisely assess ontogenetic changes produced at different temperatures (Przibram, '17b). Neither Sumner nor the Przibram group, however, could conclusively show whether changes in mice and rats kept at different temperatures from birth produced heritable characters. Reviewing Sumner's career, Charles Manning Child later wrote: “The early results seemed to show positive inherited effects of the different temperature environments, but later data were not entirely consistent and the work ended in uncertainty (Child, '47, p 154).” From 1885 to 1901, regeneration studies took center stage among experimental biologists and anatomists in Europe and America, as Frederick B. Churchill has argued. Even before he finished his PhD, Przibram began his career with a study of regeneration in lower crustaceans (Przibram, 1899). Describing regeneration as the organism's potential to re-establish an equilibrium of form (Przibram, '07b), Przibram made feasible the

219 use of Hans Driesch's notion of a harmonic-equipotential system (Driesch, '01) in his “physiology of morphogenesis” (Przibram, '10a). Regenerative processes, whether experimentally induced or naturally occurring, were accessible to experimentation, and, with a wealth of data and refined methods already available, important open questions awaited answers. Przibram's first point of departure was the crucial issue of whether the capacity of regeneration is inherent to all organisms, or whether it is acquired in only some species or organs (via the inheritance of acquired characters). According to Przibram ('08b), many experiments on regeneration favored the inheritance of acquired characters, and, therefore, the conditional occurrence of regeneration. For example, it could be shown that an injured organism as a whole once again strives toward a perfect form either by reduction (negative growth) or compensation (surplus growth). Przibram, therefore, took regeneration as an adaptive reaction of animals to their environments; it became a key feature of his physiomorphogenetic program, grounded in his belief that the chemical composition of the body can generate specific bodily forms (Przibram, '10a). Across his programmatic publications, Przibram repeatedly discussed the main themes of morphogenesis. These included the following: (1) The quality of form, or self-differentiation; (2) the issue of which species-specific or phyletic characters the developed organ retains (the problem of specificity); and (3) the problem of sex determination, perhaps, the most difficult of all biological problems (see Morgan, '13). In an experiment on the regeneration of the gonads of a hermaphroditic worm (Criodrilus lacuum Hoffm.), Viktor Janda, a zoologist from Prague and a collaborator at the BVA, removed the anterior part of the annelid and observed that a new anterior end regenerated with ovaries and testes (Janda, '12). Thus, the annelid's primary segments could develop post-larval segments with gonads, which “must be produced from parts of the body that have never produced them before,” as a puzzled Morgan wrote in 1913 (p 168–169). Using experiments with the biotechnical tools invented by Tornier (1897), Przibram also studied the “triples” that appear in the body parts of different animal groups. He discovered that triples are surplus regenerates of fractures and, therefore, a symmetrical repetition of the form of the same side of the body. He concluded that the formation of such fracture triples [Bruchdreifachbildungen] might involve collapsible organic spatial grids. Bateson had earlier argued instead that triples represented inborn mutations, in effect rejecting their regenerative origin (Bateson, 1894). But he later agreed with Przibram's interpretation in terms of injury, emphasizing that the origin of triples differs from simple regeneration (Bateson, '13). But using crabs, Przibram then demonstrated that the development of fracture triples corresponds to Bateson's symmetry rule, suggesting a regenerative process. The formation of triples and typical regeneration, therefore, differ only in the nature of the injury involved (Przibram, '19). J. Exp. Zool.

220 After the war, Przibram's decision to narrow the Department's emphasis to ontogeny and regeneration is well reflected in the research of Paul Weiss, who exemplifies the shift in emphasis from the environment of the germ-plasm to the physiology of morphodynamics. In his experiments on nerve–muscle regeneration, Weiss followed a path that was pioneered, among others, by Harrison. Harrison's results indicated that limb tissues provide pre-neural pathways, later named “stereotaxis (Harrison, '08, p 409).” Combining extirpation with the autophore replantation of amphibian eyes (Koppanyi, '23; Pardo, '06; Uhlenhuth, '12, '13), Weiss succeeded in transplanting limbs in amphibians. He poked a small hole into the axillary area of the animal and slid the proximal part of the upper arm (or upper leg) into the hole. The muscle tension around the opening fixed the transplant onto the host (Weiss, '23). For example, he implanted a denervated limb muscle from the right side into the left side of the back, where a nerve regrew into the muscle. The muscle was then strengthened with both ends to the skeleton, enabling it to carry out isometric contractions alone, but with no perceivable locomotion. Nevertheless, the supernumerary transplanted muscle did move in the same way as the normal control muscle in the right leg. This meant that muscles in the transplanted limb functioned simultaneously and with the same degree of intensity as the homologous muscles in the adjacent normal limb. In Weiss's model, chemical constitution acts as an agent that selects the appropriate pathway to the muscle. Weiss rejected conventional tropism theories and instead invented a concept of specificity that assigns to every nerve its appropriate muscle. The outgrowing nerve reaches its muscle because nerve and muscle resonate with each other, both tuned to a specific mode of oscillation (Weiss, '24). Weiss's resonance theory was later attacked for several reasons, most sharply by his doctoral student Roger Sperry, and the issue was never completely resolved. Physiology. Eugen Steinach's Physiology Department was smaller, and it concentrated on a much narrower range of questions than Przibram's Zoology Department. But these questions had a pervasive and controversial impact on contemporary views of the origin of sex differences and their cellular foundation. Steinach explored the neural and endocrine physiology of anatomical, histological, and behavioral sexual attributes in vertebrates, especially frogs and mammals. He was an early and significant proponent of the idea of an interstitial gland, an hypothesis first proposed by the French anatomists Paul Ancel and Pol Bouin. Until 1900, the interstitial cells were considered connective tissue packed into the testicular spaces containing the seminiferous tubules; gonadal hormones had been proposed, but they were considered the product of the sperm produced in those tubules. In 1903, Ancel and Bouin proposed that the gonad was actually a double organ, in which the interstitial cells of the testis functioned as a separate endocrine structure biologically and functionally distinct from that J. Exp. Zool.

LOGAN AND BRAUCKMANN producing the gametes. Between 1910 and 1930, Steinach's analysis of gonadal histology repeatedly demonstrated that, in mammals, it was indeed the interstitial cells that produced the chemicals promoting the development of masculine and feminine anatomical and behavioral attributes. Harry Benjamin described his eyewitness account of the reaction to Steinach's summary of his work, which was presented in 1926, at the First International Congress of Sexual Research. The account shows that important figures in European microscopic anatomy, who had once challenged the idea that the interstitial cells were secretory, finally acceded to Steinach's evidence showing that the interstitial cells are a secretory “organ” guiding sexual development (Benjamin, '45). In 1910, Steinach proposed a process that he termed “erotization.” The concept referred to the integration of neural and secretory processes that, under the influence of sex hormones (and sometimes acting with social synergy), give rise to the sex drive and to structured sexual behavior. To demonstrate this, he pioneered in mammals the method of infant castration accompanied by gonadal transplants that were executed on very young animals (Steinach, '10). Both same- and crossed-sex transplants showed that the putative hormones could (because the transplants severed all neural connections) act autonomously to flexibly shape the development of masculinity or femininity in either males or in females. Infant males were feminized by transplanted ovaries, and infant females were masculinized by transplanted testes (Steinach, '12, '13). Strikingly, implants of both ovaries and testes in the same individual during infancy produced cyclic hermaphrodites—animals (either males or females) that alternately expressed both male and female sexual attributes (Steinach, '16). Most of this work was completed before the chemical structure of gonadal steroids became known in the late 1920s and 1930s. But after about 1925, as the chemicals' structures were being revealed, Steinach, like others, started working with injections of purified glandular extracts. Through his collaboration with the German pharmaceutical company Schering-Kahlbaum AG, these extracts were rendered more purified and made more therapeutically effective. By 1930, he and his students had replicated experimentally most of the implant findings using injections, confirming that masculine and feminine attributes in anatomy and behavior develop flexibly under the influence of hormones produced by interstitial cells. To integrate hormonal action with other influences also altering sexual motivation, Steinach also proposed retroactive control of the anterior pituitary gland produced by secretions from the gonads, as well as the presence in the brain of neurohormones that, he speculated, could alter both pituitary and gonadal output. His suggestion that gonadal secretions alter the anterior pituitary preceded by 4 years, the more famous announcement made in the United States in 1932 by Carl Moore and Dorothy Price (Moore and Price, '32). In 1928, with the BVA

EXPERIMENTAL ZOOLOGY IN VIENNA scientist Heinrich Kun, Steinach reported the effects of injections of pituitary extracts taken from cattle on male reproductive behavior and anatomy. This was at almost the same time that Bernhard Zondek and Selmar Aschheim in Germany reported the effects of pituitary hormones extracted from the urine of pregnant mammals on females. Steinach even replicated his findings on males with extracts of pregnant females' urine purified using Zondek and Aschheim's procedure. Both teams helped establish the importance of pituitary secretions: in males and females, anterior pituitary extracts stimulated the early development of sexual and reproductive attributes via their effects on gonadal secretions. Steinach and Kun ('28) also showed that pituitary injections restored and maintained normal sexual attributes in sterile young males and in “senile” adult males. Steinach's student Berthold Wiesner had already demonstrated a comparable effect on aging ovaries: A follicular “reenlivening” occurred after sexually mature (castrated) adult females with intact pituitaries were given atrophied ovaries from aging females (see Benjamin, '28). The effects of pituitary injections on reactivating male and female gonads led Steinach to propose that many reproductive insufficiencies, which had been attributed to impaired gonads, might instead result from the insufficiencies of under-functioning pituitaries. Steinach and Kun, therefore, speculated that gonadal secretions also regulated the pituitary gland. Steinach had interpreted his controversial vasoligation procedure as indicating that increased gonadal secretions could rejuvenate old mammals and elderly humans (see Hirshbein, 2000). But Steinach and Kun now proposed that such rejuvenating effects of gonadal steroids could be the result not of the direct action of increased gonadal secretions, but of the reactivation of the pituitary by surgically induced increases in gonadal hormones. They wrote: “One can more easily consider [rejuvenation] as a retroaction [R€ uckwirkung] of the gonads on the hypophysis . . . ('28, p 529).” The regulatory control was, on their view, the result of “the reciprocal interaction of hypophysis and gonads ('28, p 529, our stress).” Steinach's research on neurohormones and on the anterior pituitary was preliminary; his age, the economic situation in Austria, and the fate of some of his students, precluded its mature elaboration at the BVA.6 But his influence was eventually felt through several students and colleagues who were either not Jewish or who had been able to leave Central Europe before the Nazis entered Austria. All of them established scientific careers in endocrinology elsewhere: Alexander Lipsch€ utz in Chile, Berthold Wiesner in England, and Oskar Peczenik in England and then Israel; all had done developmental analyses of hormone action at the BVA as well as research on the anterior pituitary. Further, Przibram's student Eduard Uhlenhuth, who came to the United 6 Josef Schleidt who performed the first pituitary studies in Steinach's laboratory was killed in World War I; Heinrich Kun was killed in the Holocaust.

221 States in 1914 and remained until he retired, is credited with the first demonstration of an anterior pituitary thyrotropic hormone, a pituitary secretion that regulates the thyroid gland (Schwarzenberg and Uhlenhuth, '28; Uhlenhuth and Schwarzenberg, '27).7 Most significant, however, is the work of Steinach's student Walter Hohlweg. Born in Vienna, Hohlweg was trained as a chemist. Before completing his doctorate, however, he worked in Steinach's Physiology Department from 1925 to 1928, as part of Steinach's effort to bring greater chemical expertise to his research program. Hohlweg left Vienna in 1928, when he was hired by the Schering Pharmaceutical Corporation in Berlin. There he continued to use Steinach's physio-behavioral techniques along with increasingly sophisticated chemical methods. After World War II, war damage caused major restructuring at Schering, and Hohlweg left to become head of the Institute for Experimental Endocrinology at Berlin's Charite hospital (Walch, 2010). In 1932, working with Max Dohrn at Schering, Hohlweg published further evidence of the regulatory effect of gonadal secretions on the pituitary, confirming Steinach's interpretation that the two glands act reciprocally. Exploring the so-called “castration cells” that appear in anterior pituitary histology following castration and that increase the gland's secretory effects, Hohlweg and Dohrn ('32) showed that administration of a purified estrogen extract reversed the histological changes produced by castration. This was the same year in which Moore and Price published their similar conclusion. Using very different concepts and procedures, the Hohlweg and Moore teams had both confirmed that gonadal secretions inhibit secretions in the anterior pituitary. The two glands indeed act reciprocally.8 In the same year, Hohlweg and Junkmann ('32) went further to implicate involvement by the brain. They used pituitary implants in kidney to demonstrate that the histological changes induced by castration in the pituitary gland could be reversed by estrogen injections, but only in native pituitaries retaining their connections to brain. Those glands transplanted to kidney failed to react to estrogens. This led the authors to propose the first model of brain–pituitary–gonadal interaction (see Logan, 2015). The model hypothesized a neural “sexual center,” which detected gonadal hormones and inhibited anterior pituitary secretions. Hohlweg and Junkmann confirmed such neural involvement by 7 Uhlenhuth obtained his PhD at the University of Vienna and then worked in Przibram's Department. After 1914, he worked in Alexis Carrel's laboratory at the Rockefeller Institute for Medical Research, and he eventually became Dean of the Medical School at the University of Maryland. 8

The announcements were apparently independent of one another. On their priority dispute, see Simmer and Suß ('93). Interestingly, Moore and Price did not use the word “feedback;“ they wrote of “suppression.“ Hohlweg and Dohrn used a modification of Steinach's term “hemmende R€ uckwirkung (inhibitory retroaction) “ to describe a feedback effect.

J. Exp. Zool.

222 showing that sectioning cranial nerves X and XI in rabbits and then castrating the animals produced a partial or complete “inhibition” of the castration-induced histological changes commonly seen in the anterior pituitary. These two path-breaking papers were the foundation of Hohlweg's long career, one that also included the discovery of the positive effects of estrogens specifically on the secretion of luteinizing hormone in the pituitary, pituitary “rebound” effects that follow sustained injections of high doses of estrogens, which, Hohlweg suggested, result from reduced sensitivity of the brain to estrogens, the isolation (with A. Butenandt and W. Westphal) of the crystalline form of progesterone, the use of progesterone to trigger menstrual bleeding in castrated females, and the development of oral preparations of estradiol and progesterone (see Rohde and Hinz, 2010). Using a range of methodologies, Hohlweg's work reflects the long-ranging impact of the BVA Physiology Department. Steinach's group had combined biochemistry, endocrine physiology, and neural mechanisms to explore the integration of chemistry with anatomy, neurophysiology, and sexual behavior. In so doing, Steinach and his students laid the foundation for significant aspects of a regulatory and functional neuroendocrinology of masculinity, femininity, and fertility. International Impact Przibram's strong connections to the international zoological community and to the emergence of experimental embryology and endocrinology as distinct disciplines led to considerable research transfer to other countries, as BVA scientists traveled to other parts of Europe (Karl von Frisch, Wiesner, Peczenik), South America (Alexander Lipsch€ utz), and North America (Uhlenhuth, Theodor Koppanyi, Weiss). For example, Weiss who started his scientific career as Przibram's doctoral student, moved to the United States in the fall of 1931, when he was awarded a Sterling Fellowship at Yale to work in Harrison's lab. His research program further developed the transplantation and tissue experiments that he had learned in Vienna and at the Kaiser-Wilhelm Institute for Biology in Berlin-Dahlem. In the United States, with funding from the US Office of Scientific Research and Development, he transplanted fragments of the central nervous system using new techniques, such as freeze-dried nerve banks. Further, he discovered that axons facilitate the progression of their own growth (Weiss and Hiscoe, '48). At about the same time, he conceptualized cellular surface contact as a mode of cell recognition, which he baptized “molecular ecology (Weiss, '49, p 147).” This led to experiments on the interactions between cells and tissues as contact agents and to regulatory cell-type specificity as possible catalysts in cell differentiation (Weiss and Taylor, '60). By his retirement in 1965, Weiss had become one of the most influential developmental biologists in the United States, working first at the University of Chicago and then at the Rockefeller Institute in New York. He even motivated the turn J. Exp. Zool.

LOGAN AND BRAUCKMANN toward the disciplinization of developmental biology. For in 1950, as Chairman of the National Research Council Division of Biology and Agriculture, Weiss sent his colleagues a questionnaire about institutionalizing the field of “developmental biology.” His plan was to split classical experimental embryology into two fields: developmental biology and (molecular) cell biology (Brauckmann, 2013). The Fate of the BVA In the 1920s, the BVA was steeped in controversies that surrounded several topics: Steinach's rejuvenation concept, the place of phenotypic plasticity and biological novelty in the natural world, Kammerer's suicide and the status of the inheritance of acquired characteristics, and the social implications of plastic sexuality. All were hotly debated as new scientific specialties emerged in ways fractured by a world war, and all were inflamed by an intensification of Austrian antisemitism within the University of Vienna (Taschwer, 2014) and aggravated by decreasing financial support for the BVA. Further, several findings and the controversies surrounding them were discussed in newspapers in Europe and America. The New York Times, for example, often quoted Kammerer's sometimes sensationalist comments on the inheritance of acquired characteristics as well as descriptions of Koppanyi's (by then at the University of Chicago) work on eye transplantation. Kammerer himself welcomed the attention. But many scientists believed that inviting such press notoriety had no place in science. It was considered unbecoming to a serious scientist, and the excitement that surrounded it stigmatized the science in ways that altered the view of the BVA outside of Vienna. Thomas Hunt Morgan, for example, refused to be on a scientific committee welcoming Kammerer to the United States in 1923, in part, because he believed that “this kind of advertising is the sort of thing for which no man interested in real scientific development” should be responsible.9 Even in Vienna, the powerful scientist-politician, Julius Tandler, disapproved of Kammerer's grandstanding (see Logan, 2013). The Kammerer Scandal. In 1926, Paul Kammerer's, one remaining midwife toad specimen, reflecting animals studied and bred in careful experiments done before 1910 (Kammerer, '09), was discovered to have been enhanced with India ink. Though several key scientists, including Przibram, defended Kammerer, and his data were replicated by several others (Berz and Taschwer, 2010), Kammerer's suicide was taken by many as an admission of guilt. Whether or not he committed the fraud, the scandal and its effect added to concerns about Kammerer's sensationalism to alter the reputation of the BVA, at least in American eyes. A key concept—

9 T. H. Morgan to Paul Maerker-Brandon, American Philosophical Society Library and Archives, Paul Kammerer Papers.

EXPERIMENTAL ZOOLOGY IN VIENNA the inheritance of acquired characteristics, which had once been central to the BVA program—had been discredited. The scandal damaged the international reputation of the BVA. But many other influences also altered the international visibility of the Institution in the late 1920s. The attitude of European biologists toward the BVA is a complex fusion of the disruption caused by war, scientists' diverse personalities, important social biases, as American eugenics and Fascism both influenced the conduct of science, and jealously, as the scientific establishment in Europe began to be challenged by the upstart United States. Even in 1924, for example, a sympathetic Herbert Spencer Jennings of the Johns Hopkins University privately accused the British geneticist William Bateson of making “a rather sorry spectacle” in his criticism of Kammerer's work. On a visit to the BVA in 1910, Bateson had admitted that he just didn't like the idea of the inheritance of acquired characters, and that view strengthened over time. One BVA visitor even referred to the Institute as “Pzribram's sorcery institute.”10 It was as if a fear of adaptive novelty and increasing nationalist jealously (perhaps fueled by war) had contaminated the consideration of BVA science. Though much historical documentation has been lost, we can conclude that for a number of reasons, the integrative approach to adaptive novelty in morphogenesis, physiology, and behavior represented by the two BVA departments most centered on animals, was by the 1930s, no longer relevant to several pioneers of early 20th century experimental biology. Other important changes were also occurring within biology itself: specialization was rampant and organismic syntheses began to be impeded by a method that, necessarily and to great ends, stressed precise analysis over synthesis (Maienschein, 1991). In addition, in the United States and Great Britain, especially, the historical connection between evolution and development was weakened by the growth of the statistical approach to evolution represented by Sewell Wright and others (Provine, 2001). This was less the case in traditional experimental embryology, where developmental perspectives continued to dominate. Harrison, for example, exchanged reprints with Przibram, brought Weiss to his lab, helped Weiss obtain a visa for entry into the United States, and later supported Weiss's search for a tenure-track position at Chicago. Harrison clearly welcomed the BVA emphasis on ontogenetic plasticity reflected in its younger scientists who immigrated to the United States. But this was only one of many fields represented in the promised BVA synthesis of the pre-war period. Internationally, the exploration of phylogeny turned away from an understanding of adaptive morphogenesis to emphasize instead the rise of classical genetics, Mendelian approaches to heredity, and the statistical analyses of population variability. Topics, such as regeneration and morpho10

Quotes are from the HS Jennings Papers and the William Bateson Correspondence, Paul Kammerer Papers. The American Philosophical Society Library and Archives.

223 genesis, were increasingly confined to special disciplines and divorced from evolution, while others, such as the inheritance of acquired characteristics almost disappeared. Even the biologically oriented Richard Goldschmidt would, in 1950, write that consideration of Lamarckian problems like the inheritance of acquired characteristics was one of the “childhood diseases (1950, p 320)” of the young science of genetics. CONCLUSION

The 20th century saw enormous progress in virtually all areas of the biological sciences. The BVA was a part of this for over 30 years. The institute pursued an ambitious research program that covered most disciplines of the biological sciences: developmental physiology, inheritance and variation, plant physiology and botanical microchemistry, protein chemistry, evolution, and neuroendocrinology. For example, as we show, Przibram's research program combined development and evolution, issues of inheritance and breeding, with an experimental analysis of the impact of environmental factors on ontogenesis. What further distinguishes the BVA was the initiative to combine a thoroughgoing experimental approach with a theoretical underpinning and, even more important, a new take on experimental and theoretical biology. It aimed at to clarify how ontogenetic adaptations, conceived chemically and physiologically and made experimentally accessible, could then help reveal phylogenetic adaptations, and the reverse. Ontogeny and phylogeny were once again linked, but in a very different way than that associated with the old idea of biogenesis (Gould, '77). Further, the Institute addressed the problem of regulation with rigorous work on transplantation and regeneration, employing advanced methods and relying on exceptional techniques and equipment. The number of species bred at the BVA was impressive, as was the diversity of the organisms used in experiments to facilitate the integration of comparative and evolutionary perspectives with precise causal control. Evolutionary-oriented comparative physiologists today often address the ironic paradox of “malignant species neglect” (Preuss and Robert, 2014, p 65) in contemporary cellular and molecular research. Despite its tragic fate, therefore, it is reasonable to conclude that the Biologische Versuchsanstalt serves as one of many excellent historical examples of the thoroughgoing compatibility between molecular approaches and organismic, developmental, and ecological zoology.

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