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The History of the Stazione Zoologica Anton Dohrn
An Outline
by Bernardino Fantini
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The creation and the first successes
The foundations of the Stazione Zoologica were laid in March 1872. Anton Dohrn, founder and first director, was born in Stettin, Pomerania, present day Poland, in 1840, into a well-to-do bourgeois family. Anton’s grandfather, Heinrich Dohrn, a merchant in wine and spices, had made a fortune in the sugar industry; Anton’s father, Carl August, could therefore devote himself to his various hobbies, such as travelling, collecting folksongs and insects. Anton, the youngest son, studied zoology and medicine at various German universities (Königsberg, Bonn, Jena and Berlin), without much enthusiasm. His ideals changed in summer 1862 when he arrived at Jena where Ernst Haeckel introduced him to Darwin’s work and theories. Dohrn became a fervent defender of Darwin’s theory of ‘descent with modification’, the theory of evolution by natural selection. He then decided to dedicate his future life to collecting facts and ideas in support of Darwinism, the starting point for a lifelong adventure.
At that time comparative embryology was becoming the cornerstone of morphology and evolution, based on Haeckel’s recapitulation theory : the idea that an organism during its embryonic development passes through the major stages of the evolutionary past of its species. Morphology thus became one of the major ways in which zoologists sought to expand and develop Darwinian theory in the last 30 years of the 19th century. Dohrn chose to become a ‘Darwinian morphologist’: the evolutionist Charles Darwin and the embryologist Carl Ernst von Baer became his scientific ideals, immortalized in the two marble busts located in the Fresco room of the Stazione Zoologica.
While pursuing his university career (PhD in 1865 at Breslau, ‘Habilitation’ in 1868 at Jena) Anton Dohrn worked several times at facilities located on the seashore: at Helgoland with Ernst Haeckel in 1865, at Hamburg in 1866, at Millport in Scotland with David Robertson in 1867 and 1868 and at Messina during the winter of 1868-1869 together with his Russian friend and colleague Nicolai Micloucho-Maclay. In Messina the two friends planned to cover the globe with a network of zoological research stations, analogous to railway stations, where scientists could stop, collect material, make observations and realize experiments, before moving on to the next station.
The fascination for the sea
Naturalists and philosophers have always shared a fascination for marine life. This curiosity had produced, with a mixture of scientific observation and traditional tales, the great Naturalis Historiae of the past, which devoted large attention to animals living in the sea, an inexhaustible source of mirabilia and fantastic monsters, a place of mystery and a permanent source of life and beauty. In German scientific culture of the 19th century, the sea was at the same time the place to look for the most elementary forms of life and the symbol of the endless search for knowledge. This search needed to move out of the secure harbor, venturing in the open sea of doubts, questions, and critics. As Alexander von Humboldt wrote at the end of a long life devoted to travel and research on nature and truth : "He who loves to create in himself a world apart in which to free the spontaneous activity of his soul, he will fulfil the sublime idea of the infinity, as the sea free from any coast".
Marine organisms became the center of interest for philosophers and naturalists who accepted the Naturphilosophie and, after the origins of the first forms of cellular theory, adhered to a ‘protoplasmic theory of life’, looking into the sea depths for an ‘elementary living matter’, the Urschleim, the elementary monad endowed with life. The new generation of biologists in the 1850s and 1860s, well versed in the Naturphilosophie and the new biological doctrine, Darwinism, looked to the sea as a source of knowledge and as a life experience. In the second half of the 19th century, the sea, because of the richness of living forms that inhabit it, in particular the simplest forms of life, produced models, experimental objects, and metaphors for fundamental biological problems: the organizational plan of living systems, embryogenesis, general physiology, evolution, and phylogeny.
The interest in marine organisms for the understanding of the fundamental problems of biology was prompted by Johannes Müller (1801-1858), the father of physiology and theoretical biology in Germany. Müller widely publicized the concept of marine biological research as a means of elucidating fundamental biological concepts. And Johannes Müller was the name of the first boat used by the Stazione Zoologica for collecting marine fauna and flora.
The first laboratory of marine biology
The potential of marine organisms for morphological, physiological and systematic studies, and their economic interest, had already led several scientists and academic institutions to try to establish study facilities near the sea.
The first marine laboratory was created in 1843 at Ostende by P.M. van Beneden, zoologist and parasitologist, professor in Leuven, father of the more well known Edouard, the discover of meiosis. In France, the first laboratory on the sea edge was founded in the 1850s at Concarneau, on the Southern coast of Bretagne, by J.-J. Coste, professor at the Collège de France. This laboratory produced relevant physiological researches. In fact, marine biology laboratories played an important role in France in the formation of a new generation of biologists and physiologists, who became for the first time a real part of the scientific establishment. Another French marine laboratory was created by A. Giard in 1873 at Wimereux and this laboratory, two years later, became part of the École Pratique des Hautes Études. The Roscoff marine biology laboratory, created in 1873 by Henri de Lacaze-Duthiers also played a fundamental role in the institutionalization of marine biology in the last decades of the 19th century, thanks as well to the diffusion of the results realized with the publication of the Archives de zoologie expérimentale. Lacaze-Duthiers founded another French laboratory in 1881 at Banyuls-sur-mer on the Mediterranean coast.
In the United States one can find practically the same initiatives, a good indication of the need for these kind of research institutes. In 1873 Louis Agassiz created the Anderson School of Natural History, on Pekinese Island, and in 1878 Johns Hopkins University founded the Chesapeake Zoological Laboratory, under the direction of W. H. Brooks. Then, in 1888, the Marine Biological Laboratory was founded at Woods Hole and in 1892 the first laboratory on the west coast, the Hopkins Marine Station, opened in California.
However, these were mostly field stations, connected to a university or institute; they were not independent facilities meant to house a host of different sorts of investigations and a wide variety of projects. Furthermore, there were two different types of marine laboratories: the Stazione Zoologica di Napoli, as well as the laboratories in Trieste and Sebastopole, were exclusively devoted to research and advanced training, while French and American institutions were mainly aimed at teaching, although during the summer term, they housed relevant research groups and activities.
Why Naples ?
At Messina Dohrn rented two rooms for the "Stazione Zoologica di Messina" (February 1869), but he quickly realized the technical difficulties of studying marine life without a permanent structure (difficulty in collecting specimens, lack of storage tanks with flowing seawater, lack of a library and lack of technical assistance from well-trained personnel). Faced with such difficulties, Dohrn started to dream about the usefulness for scientists to arrive near the sea and find the "table laid" for work, that is, to find on arrival instruments, laboratory space, services, chemicals, and books available together with records of where and when certain species could be found and useful information on local conditions.
He left, in Messina, his books, equipment, diaries and the portable aquarium he had brought with him from Scotland for those who would come after his departure. In 1870 Dohrn decided that Naples would be a better place for his Station. This choice was due to the great biological richness of the Gulf of Naples and also to the possibility of creating a research institute of outstanding international relevance in a large town that itself had a strong international vocation.
After a visit to the newly opened aquarium in Berlin he reasoned that an aquarium open to the public might earn enough money to pay a permanent assistant to the labs. Naples, with its 500,000 inhabitants, was one of the largest and most attractive cities of Europe and also had a considerable flow of tourists (30,000 a year) that would be potential visitors to the Aquarium.
With a mixture of imagination, will-power, diplomatic dexterity and good luck, and with the friendly support of scientists, artists and musicians, Anton Dohrn overcame doubts, ignorance, and misunderstandings and persuaded the city authorities to give him, free-of-charge, a plot of land at the sea edge, in the beautiful Royal Park (today, the Villa Comunale). For his part he promised to build a Stazione Zoologica at his own expense.
Dohrn knew exactly what he wanted and how he wanted it done — the plans for the buildings are his. The foundations of the Stazione Zoologica were laid in March 1872, and by September 1873, the building was finished. Two-thirds of the building costs came out of Anton Dohrn’s and his father’s pockets, the remaining third was provided by loans from friends. After the first building - today’s central part - a second building, connected with the first by a bridge, was added in 1885-1888, the courtyard and western part in 1905. Fifty years later the Library was inserted between the first and second building.
The core building contained pumps, machines, store rooms and seawater tanks in the cellar, the public aquarium in the basement, a large laboratory for about 12 scientists and the fresco room housing the library on the first floor, and 12 smaller labs and living quarters for the custodians and assistants on the second floor.
The first scientists arrived in September 1873: 2 from Germany, 3 from the U.K., 2 from Russia, 2 from Italy, and 1 from the Netherlands.
The public aquarium, which covers 527 square meters, was opened on 26 January 1874. It is unique because it has changed very little since its creation and it is the oldest 19th century aquarium still functioning as well as the only one exclusively dedicated to Mediterranean fauna. It was built under the supervision of William Alford Lloyd, an English engineer who had helped to build the public aquariums of Hamburg (1868) and London (1871).
The official inauguration of the Stazione Zoologica took place on April 14th, 1875 and only in December of the same year the contract between the City of Naples, represented by the Mayor Senatore Antonio Winspeare, and Anton Dohrn was signed.
Anton Dohrn’s Dream
The Stazione Zoologica was the product of a dream, of a visionary project. "To create, to organize, to develop – this is my need, this is even my passion" (Anton Dohrn to Marie Dohrn, August 1883). Dohrn wanted "to make a work of art out of life itself … I found chaos before me and have created out of that both a practical organism (the Stazione), and a theoretical one, the "Urgeschichte der Wirbelthiere. Each step on the path of these two things I have envisioned beforehand, as an artist sees first the complete work of art and then starts to create its parts" (Anton Dohrn to Marie Dohrn, August 1, 1886).
The Stazione Zoologica served as an example for many other marine stations and institutions: the Marine Biological Laboratories of Woods Hole, Misaki, Plymouth, the Kaiser Wilhelm Institutes (later Max Planck Institutes) and the Rockefeller Institute for Medical Research.
The Stazione Zoologica di Napoli fairly soon acquired a special status in the panorama of scientific institutions, because of its peculiar institutional nature, its potentiality for advanced biological and marine research, and its pervasive and involving cultural atmosphere. It was an international institution founded by a German, managed as a private and familial enterprise and organized according to the German academic tradition. But it was located in Italy and allowed any country to participate in its scientific life and provide for its financial support.
One of the features that contributed to its success was the lightness and flexibility of its structure. Because of the high costs and the technical difficulties implied, for marine biology, it was particularly apt what Anton Dohrn wrote in 1872 for biology in general: "Concerning its structures as a science, Zoology compares to astronomy and Mechanics as Garibaldi’s volunteers are in relation to a regular army corps. The former cannot realize great strategic moves, in which very little depends on change and individual initiative, but everything comes from the well coordinated and calculated action of many. In the same way, zoology, in its present condition is unable to solve the big questions gradually and according to a pre-established plan. It lacks the organization which is necessary for that".
The idea of a small scientific structure, flexible and agile, but full of initiatives and bravery, as a Garibaldean corps, constitutes a constant character of the Stazione, from its origins to today.
The international character of the Institute (a novelty at that time) was secured with the help and support of influential scientists such as Darwin himself (see "Darwin-Dohrn Correspondence", Naples, 1982), Michael Foster, Louis and Alexander Agassiz, Carl Ernst von Baer, Rudolf Virchow, Emil du Bois-Reymond, and, of course, several Italian zoologists, the most prominent being Paolo Panceri, the teacher of outstanding Italian zoologists of the end of the 19th century (Carlo Emery, Antonio Della Valle). Panceri unfortunately died in 1877 at age 44; according to Dohrn himself, he was the first to help him in Naples and the first to understand what the Stazione needed to be, as did his successor Salvatore Trinchese, the teacher of Francesco Saverio Monticelli, Federico Raffaele and Giuseppe Jatta. Notwithstanding his nationality and cultural background, Dohrn found a great source of strength, encouragement and help for his dream of a marine laboratory in the British natural history tradition, because his project was really at the focus of biological research. In 1870, at the annual meeting of the British Association in Liverpool, a committee was formed ‘for the purpose of promoting the foundation of zoological stations in different parts of the world.’ Indeed, it was this committee, through its many reports as well as notes and articles regularly published in Nature, that gave such widespread publicity in the English-speaking world to Dohrn’s Stazione Zoologica.
In order to promote the international status of the Stazione and to guarantee its economic and hence political independence and freedom of research, Dohrn introduced a series of innovative measures to finance his project, first of all the rental of work and research space (‘table system’): for an annual fee the contract partner (universities, governments, scientific institutions, private foundations, even individuals) could send one scientist to Naples for one year where he or she would find available all that was required to conduct research (lab space, animal supply, chemicals, an exceptional library and expert help from the staff), ‘no strings attached,’ in the sense that investigators were completely free to pursue their own projects and ideas.
This so-called ‘research table system’ or ‘Bench system’ worked extremely well. By 1890, 36 tables were rented annually by 15 different countries, and when Anton Dohrn died in 1909 more than 2,200 scientists from Europe and the United States had worked at Naples and more than 50 tables per year had been rented out. It was in fact at Naples that international scientific collaboration in the modern sense was invented, based on quick and free communication of ideas, methods, techniques, instruments, and on exchanges and personal contacts between scientists of different cultural traditions.
In order to diffuse the results of the scientific work at the Stazione and at the same time to secure additional income, Dohrn launched three editorial initiatives. The first was a scientific journal, Mittheilungen aus der Zoologischen Station zu Neapel (1879-1915), continued as Pubblicazioni della Stazione Zoologica di Napoli (1924-1978), and later as series I Marine Ecology (1980- ) and series II History and Philosophy of the Life Sciences (1979- ). The second was Zoologischer Jahresbericht, a reference journal (1880-1915) and the third the outstanding series of monographs Fauna e Flora del Golfo di Napoli (Fauna and Flora of the Gulf of Naples, 1880-1982). Through its publications the Stazione greatly contributed to the knowledge of marine fauna and flora. Dohrn hired two Neapolitan artists, Comingio Merculiano and Vincenzo Serino, for the illustrations which soon became famous for their naturalness and beauty.
Dohrn also started a specimen supply program as another source of income. Thanks to the inventiveness and skills of another Neapolitan, the preparator Salvatore Lo Bianco (1860-1910) who entered the service of the Station at the age of 14, the methods for preserving marine organisms were improved to such a level that the Stazione Zoologica soon became known for the beauty and perfection of its collections of preserved marine animals. Specimens and collections of preserved animals were sold to museums, universities, schools and private individuals. Some of Lo Bianco’s specimens can still be admired in the Stazione’s Aquarium and Special Collections.
Lo Bianco also became a systematicist in his own right and many guests of the Stazione acknowledged his relevant contribution to their researches. According to Grassi (1911) ‘one can say that the world renown of the Stazione Zoologica di Napoli is due in large part to Lo Bianco’s activity’.
Anton Dohrn was convinced that the availability of all major published sources in addition to good working facilities was a necessity for advanced research. He donated his own large library to the Stazione Zoologica and requested donations from scientific publishers, academies and scientists such as Darwin, Huxley, and Virchow. The Stazione’s journal Mittheilungen served also for exchange purposes, while literature reviewed in the reference journal Zoologischer Jahresbericht was usually added to the library. Furthermore, visiting scholars felt morally obliged to send to the Station their own publications, as a sign of gratitude to the Institution. This was the source for the outstanding reprint collection catalogued and bound according to subject, discontinued only in the late 1960s because of a lack of space and personnel and due to the emergence of new technologies for information retrieval. Altogether the Naples Station’s biological reference collection became an unrivalled source for bibliographic work and in fact scientists often went to Naples just to have access to such a wonderful library, still unparalleled in Europe even today.
The Stazione also offered the best scientific equipment available, acquired through donations or at special low prices. The latest Zeiss instruments were always tested and kept available at Naples. Ernst Abbe (1840-1905) of the Zeiss factory, one of Dohrn’s few close friends, allowed the Station to purchase sets of Zeiss microscopes and other optical instruments at a significant discount; in return, workers at the Station suggested ways in which the equipment could be improved and Zeiss was brought to the attention of the international scientific community. Microtomes, staining and section cutting methods were also collected, tested and improved. Assistants and guests collaborated in improving section cutting and staining methods, thus maintaining the high level of technical services offered by the Station.
E.B. Wilson, the first American biologist to work officially at the Naples Station, that is at a properly subscribed table, believed that ‘the Station has now become practically the headquarters from which most of the leading European laboratories derive their best methods, and where, indeed, much of their most telling work is done’ (Wilson, 9 March 1883). Two things especially struck him as characteristic of this laboratory. The first was the perfection of the technical methods of research.
It is now almost proverbial for zoologists to say : "For methods go to Naples" and in the same breath is usually added "A good method is half the battle." Certain it is that many of the best modes of work now used at Leipzig, Cambridge and elsewhere have originated here. The secret of this is simply that fifteen or twenty zoologists are usually at work, who come from laboratories in all parts of the world and bring their experience to a focus here. They are all experimenting and comparing results and new methods can thus be very thoroughly tested …" [Wilson, 13 April 1883].
As Charles Otis Whitman (1883) aptly summarized it in his article on ‘The advantages of study at the Naples Stazione Zoologica,’ written after he had worked at Naples in 1881-82: ‘[The Station is] a sort of international depot for the reception of discoveries and improvements made elsewhere. The heterogeneous material thus obtained is sifted, systematized, tested, further elaborated and refined and redistributed.’
The year round animal supply, one of the advantages of the Naples Station, was guaranteed by an efficient fishing-fleet and well-trained fishermen, who knew perfectly well the local marine life forms. In 1877 the Berlin Academy of Sciences and the Prussian Ministry of Education provided funds for the ‘Johannes Müller,’ a 24 t, 17 m steamer which served for both collecting and excursion trips. Three other ships were added : the ‘Frank Balfour’ (1882-1914), steam-launch, 9 m; the ‘Salvatore Lo Bianco’ (1913-1918), motorboat, 12 m; and the ‘Anton Dohrn’ (1914-1917), covered steamboat, 27 m.
Lines of research in the first decades
The Stazione Zoologica, apart from Dohrn’s own project, did not have an in-house research project. The structure of the Institute reflected the main interests of the visiting scientists. However, according to Theodor Boveri (1910), Dohrn had ‘an unusually sure eye for the significance of the different sections of our science, and for the way in which they interrelate and complement each other.’ In such a way he was able to create a structure perfectly in tune with the main scientific problems of the time and in such a way he was able to bring to Naples the best researchers. Embryology, comparative anatomy, systematic zoology and botany - including life history, behavior and ecology - were the prevailing fields of investigation during the first decades of the Stazione’s activity.
Biology was searching for general laws: evolution by natural selection; recapitulation principle and recapitulation of the phylogenetic steps in embryonic development. The study of the morphology of the embryo during its development was the main tool for the understanding of the natural order, that is phylogeny. The synthesis between comparative anatomy, morphology, embryology and phylogeny characterizes life sciences at the time of the creation of the Stazione Zoologica. As Anton Dohrn himself put it in 1872 "In the same way as linguistics reconstructs original languages, the zoologist should be able to outline a comprehensive picture of an animal group from a large amount of embryological data … to identify the ancestor of the whole group".
As a consequence, Zoology, that is Morphology, was the first department to be created at the newly founded Stazione Zoologica in 1873, and the most important during the first twenty years. In this department worked the following: Nikolaus Kleinenberg (1873-1876); Hugo Eisig (1873-1920); Paul Mayer (1878-1913); Wilhelm Giesbrecht (1881-1913); Giuseppe Jatta (1886-1893); Pio Mingazzini (1888-1890); and Reinhard Gast (1899-1925). Marine botany was created in 1876 under the direction of Paul Falkenberg (1876-1878) and then Gottfried Berthold (1879-1881).
Although there was no room initially in the main building, Physiology was added in 1882 by renting a small building near the Station. Carl Schoenlein was the leader of this department between 1892 and 1899. The space problem was alleviated by adding two new sections to the main station, one in 1885-1888, and another in 1903-1906. A department of Bacteriology was also added in 1887, while Physiology could expand to include both Comparative Physiology and Physiological Chemistry.
Physiology
Cytophysiological studies of the embryo and the physiology of muscular and nervous functions in marine organisms cover a large part of the scientific history of the Station.
From 1892 to 1902, Jacob Johann von Uexküll was head of the Physiology department, and, for ten years, he carried out original and ingenious experiments on the nervous and muscular systems of marine animals. Uexküll was a pioneer of modern behavioral biology, investigating the environmental relations, the ‘sphere of function’ as he called it, which connect the individual to its environment. Among the other scientists that worked in the department of Physiology one should mention Richard Burian (in Naples during the years 1903-1920) and Martin Henze (from 1903 to 1921, head of the chemistry department).
Plant physiology was also represented in Naples, thanks to the research done by Friedrich Tobler ( 1879-1957) on filamentous algae and by Hans Winkler (1877-1944) on greenalga Bryopsis.
Two leading scientists working in the Physiology department at the end of the 19th century and the beginning of the 20th were Silvestro Baglioni and Filippo Bottazzi, who, as human physiologists themselves, were well aware of the great importance of the expanding new disciplines of comparative physiology and physiological chemistry. From 1905 to 1909, Baglioni worked mostly on the sensory organs of cephalopods and fishes and Bottazzi carried out his well-known experiments on osmoregulation in marine animals. This line of research originated from the concept of the milieu intérieur, formulated years before in France by Claude Bernard.
At the end of the 19th century the concept of internal environment was at the center of a large research program, whose starting point was Léon Frédéricq’s observation, made in 1882, that the body fluids of marine animals have a salt concentration practically identical to the sea in which they live. Frédéricq concluded that the milieu intérieur of these animals did not have the same level of independence as that of higher organisms. Along the same lines, at the beginning of the 20th century Filippo Bottazzi studied the influence of the chemical and physical constitution of the external environment on the internal environment, confirming Frédéricq’s hypothesis and classifying marine animals according to their independence from the external environment. Bottazzi distinguished aquatic animals as respectively homeosmotic and pecilosmotic, a distinction which is maintained today with the modern, although not necessarily more transparent terms, osmoconformers and osmoregulators.
The study of internal fluids of marine forms gave a new dimension to the concept of milieu intérieur, placing it in the context of an evolutionary theory and confirming the much older hypothesis, present more as mythology than as a scientific hypothesis, that the origin of life was in the sea. For this reason the chemical composition of fluids of marine animals is very similar to the sea. Animals had to acquire the ability to regulate autonomously their milieu intérieur before invading the terrestrial environment, bringing their own environment with them. In such a way, the Bernardian metaphor became a scientific fact: the milieu intérieur is simply the primordial environment made internal, included in the biological organization.
In 1886 Dohrn accepted, as a guest of the Stazione Zoologica, a young Norwegian zoologist, Fridtjof Nansen, the future Nobel Peace Prize Winner, at a time when neither Norway, nor any of the other Scandinavian countries had reached financial agreements with the Stazione Zoologica. In such a way he introduced a new field of interest, that would later become a ‘classic,’ one of the distinctive research projects of the Stazione. Nansen was in fact keenly interested in a problem that at the time engaged both physiologists and histologists in heated debates : the relationship between ganglion cells and nerve fibers, the nature of the nervous impulse and the cellular basis of the functioning of the brain. Nansen was one of the first scientists to recognize the great potential of the new staining method introduced and developed by Camillo Golgi at Pavia.
Within the history of Neurohistology at the end of the 19th century a relevant role was played by Stephan von Apáthy, a Hungarian neurohistologist from Koloszvar who, for three years (1886-1889), occupied the Hungarian ‘research table’ at Naples. His major breakthrough was the development of new staining methods, involving gold-chloride, formic acid, methylen blue, and hematein, which permitted the observation of ‘Fibrillengitter’ (neurofibrils) : ganglion cells and nerve fibers were shown to contain fibrils. Apathy’s preparations seemed to indicate that these fibrils formed a continuous network throughout the nervous system. The fibrils were seen to cross from one cell process to the next at their contact points.
In 1896, Albrecht Bethe, a pupil of Goltz in Strasbourg, later professor of physiology in Frankfurt, came to Naples to study the nervous system of the shore crab, Carcinus. His goal was a complete neurohistological and physiological description of the neurons in what he then considered to be a ‘simple nervous system.’ His encounter with Apáthy forced him to completely revise his ideas about the structure and function of nervous systems. As a consequence of his ‘conversion,’ Bethe looked at the nervous system as a syncytium. In an experiment that henceforth would be known in the literature and in physiology textbooks as the ‘Bethe experiment,’ he showed that ganglion cells are either unnecessary for these reflexes to occur, or the reflex arcs do not go through the ganglion cells. As he was convinced that it was the fibrils that serve as the connecting elements, he concluded that the elementary fibrils of which the composite fibrils are composed, the ‘Primitivfibrillen,’ or neurofibrils to use a modern term, are the true conducting structures of the nervous system, confirming Apáthy’s theory.
Another of the earliest contributions to the understanding of fundamental neural processes made in Naples was the discovery by Friedrich Fröhlich, in 1914, of the electroretinogram of cephalopods.
Bacteriology
In 1887, during a visit to the Stazione by Robert Koch, Anton Dohrn became convinced of the need to create a bacteriological laboratory, for three main reasons. The first was the need to include the ‘microscopical forms’ in the realm of zoological or botanical work. The second was the possibility of contributing to the scientific knowledge of bacteria, and of the specific causes of many infectious diseases, knowledge necessary to fight the cholera epidemics recurrent in the ports of Southern Italy. The third reason was as a service to public authorities for the bacteriological control of waters and marine food. The General Direction of Public Health, at the time in the Ministry of Internal Affairs, and the Municipality of Bari provided financial support for two research tables devoted to Bacteriology. Walter Krause, the co-discoverer of the dysenteric bacillus (1900), was in charge of this department and two Italian bacteriologists, Francesco Sanfelice and Sergio Panzini, worked there for a while. However, a few years later, Naples University and other Italian universities created their own bacteriological professorships and the bacteriological research was discontinued at the Stazione.
Art and culture in a scientific institute
The importance of the Stazione Zoologica goes far beyond purely scientific aspects or practical interests. It is also famous for its humanistic values and for its cultural climate: the ‘creative atmosphere’ of the Naples Stazione Zoologica and the cross fertilization between different research and cultural traditions; international contact between visiting scientists and a ‘permanent congress’ that lasted several months instead of a few days. For many scientists the ‘Naples experience’ was a good mix of new research, human experience, acquisition of new methods and new cultural experiences.
The Stazione Zoologica is the only scientific institution at which, from the very beginning, science, music and art were integral components of a unique project, the two complementary halves of an unique dream. Even at the level of the architecture, the music space, the ‘fresco room,’ with its statues, frescos, and allegories, was symmetrical to the laboratory, with its organisms, aquariums, microtomes and microscopes. The two halves formed a coherent and organized whole, defining together the ‘soul,’ the essence of the Stazione.
From the beginning Dohrn wanted to include the arts as an integral part of the home he was creating for science. The large room facing the sea and Capri was intended for the arts, music in particular. During the summer and fall of 1873, while the building was still under construction, the German painter Hans von Marées and the sculptor and architect Adolf Hildebrand decorated the room with a cycle of wall-paintings, depicting scenes from Mediterranean life: fishermen, Dohrn and his friends relaxing after a hard day’s work and orange groves with children, men and women. Space was scarce from the beginning and Dohrn had to use the room for the Library.
Art and music were essential parts of the life of the cultural elite of the 19th century, aspects of intellectual and social status, and Dohrn, like a Renaissance prince, wanted to have his ‘musical laboratory.’ At the same time, the search for form and beauty created a common background for both science and the arts, activities that share some fundamental aspects in their endless search for truth. As Dohrn himself wrote to E.B. Wilson in 1900: "Phylogeny is a subtle thing, it wants not only the analytic powers of the ‘Forscher’, of the researcher, but also the constructive imagination of the ‘Künstler’, of the artist – and both must balance each other, which they rarely do – otherwise the thing does not succeed".
The ‘golden era’ of experimental embryology (1890-1914)
Anton Dohrn did not want to interfere with the work and the projects of the scientists working at the Stazione, considering freedom of research a prerequisite for creative work. In the years 1890-1914 the SZN became the focal point for the new research direction of experimental embryology. The most important biologists of the day spent long periods of research in Naples, and the whole history of experimental biology at the dawn of the 20th century cannot be understood without reference to the individual scientific contributions made by scientists working in a collective, international and interdisciplinary environment at the Stazione Zoologica, the research institute where, in the words in Hans Driesch in 1909, ‘all basic work in modern zoology has been done’.
Created during a period when morphological studies predominated and on a clear, ideal impulsion from Haeckelian biology, the Stazione Zoologica was paradoxically one of the strongholds of the revolt against Haeckelian, phylogenetic embryology. In fact, it was in Naples that the advocates of the new experimental approach to the problems of development – Entwicklungsmechanik and Entwicklungsphysiologie – made some of their most important discoveries, thus beginning a new era in the study of development.
The Entwicklungsmechanik
Developmental mechanics or causal morphology of organisms is defined as the doctrine of the causes of organic forms, and hence the doctrine of the causes of the origin, maintenance and evolution of these forms.
The causal reasoning in biological sciences and medicine, the theoretical basis of the new experimental biology was based on the separation between intuition and demonstration. Wilhelm Roux, in the Introduction to the first issue of his Archiv, wrote: "The causal method of investigation is experiment. Certainty in causal deduction can only come from experiment, either from ‘artificial’ or from ‘nature’s experiment’, such as variation, monstrosity, or other pathological phenomena".
Developmental mechanics must seek to utilize all the tools of causal investigation of organisms, it must look for causes and modi operandi.
Richard Hertwig (1850-1937) had observed in 1875 the entry of the spermatozoon into the egg of a sea urchin and the fusion of the two nuclei, thanks to the egg’s transparency. Hertwig’s discovery produced the ‘right tool for the job’ of investigating the factors involved in embryonic development. The sea urchin egg, easy to obtain, easy to keep in the laboratory and fully transparent, became the favorite research material in experimental studies of development.
With these new tools and with the new experimental methodology, Roux conducted, in 1888, the famous ‘injury experiment’ : using a fine needle, Roux destroyed the nucleus of one of the two cells formed after the first segmentation of the zygote. He observed that the undamaged part continued its own development and produced half an embryo. Roux concluded that cellular division is qualitative, based on the mechanical distribution of the ‘determinants.’ Each cell of the early embryo has a predetermined fate, independent from the presence of other cells. These results seemed to confirm the observations made by August Weismann (1834-1914) at the Stazione Zoologica in 1881-2, following the origin of sexual cells over several generations in Hydromedusae. Weismann observed that in embryogenesis a single cell produces the whole germinal tissue of the adult animal. From those observations, he drew the conclusion that in ontogenesis the tissues that produce sexual cells (the germinal plasm) remain completely separated from the other tissues of the body (the somatic plasm).
Hans Driesch
Driesch was certainly one of the most interesting and influential personalities at the Naples Station beginning with his first visit in 1891. For a number of scientists who worked at the Station his influence was a decisive factor in their scientific life. Also, it was largely due to Driesch that researchers recognized the great advantage of the sea urchin egg. Indeed, it became a widespread belief that ‘what is true for the sea urchin must be true for all animals.’
In this context, another relevant technical advance which came to play an important role in future research was made by Kurt Herbst (1866-1946), who worked for many years in Naples, together with Driesch. In 1891 he observed that the blastomeres of the cleaving sea urchin egg spontaneously separate from each other after a brief exposure to calcium-free sea water. This produced an elegant method for isolating undamaged blastomeres and for following their development independently. At the same time, these experiments paved the way for one of the most fascinating areas of research in embryology, that of cell interactions.
Using Herbst’s technique, Driesch succeeded in separating the first two blastomeres and showing that a whole embryo could arise from each one of them. This observation, which at first appeared to contradict Roux’s results on the amphibian egg, began the long controversy on the mosaic versus regulatory organization of the egg. The aggressive method used by Driesch to separate the blastomeres by shaking the eggs (the embryologists who used Driesch’s method were derisively called ‘egg-shakers’) was later superseded by the highly sophisticated microsurgical technique devised by Sven Hörstadius in the early thirties.
In another classic experiment, Driesch changed the orientation of cellular division, squashing a sea urchin egg between two slides, showing that cells doomed to become ectoderm can also become endoderm. Later Driesch was also able to fuse two eggs together, obtaining a normal but larger embryo. Driesch’s conclusion was that the fertilized egg was a ‘harmonious-equipotential system,’ able to react to different conditions of development. Each cell of the embryo has a ‘prospective potency,’ that is what it can become, and an ‘actual potency,’ what it really becomes in a given condition of development. The harmony of the development is the result of the interaction between nucleus and cytoplasm; the nucleus guarantees the ‘transport’ of the totality of the ‘prospective potency,’ whereas the cytoplasm produces the specific effects that realize the ‘actual potency’ of each single cell.
Driesch postulated a non-material causal agent that initiates, directs and controls developmental processes. This agent controls morphogenesis, that is ‘the transformation of the possibilities into the wholeness of an actuality.’ This integrated directing, shaping, and developmental force of the embryo is called by Driesch Entelechie, recognizing that in a living organism ‘something is at work that is of non-physico-chemical character.’ This ‘something’ can only be recognized by its effects, the production of a ‘harmonious-equipotential system.’
In a letter to Reinhard Dohrn of 1911 Driesch recognizes his debt to the Stazione Zoologica: ‘my whole scientific personality is rooted in Naples … this could be only be achieved on the basis of your father’s great creation".
The problem of the cellular determination during embryogenesis is also discussed by the American embryologist and cytologist E.B. Wilson, who first entered this research field and the debate between Roux and Driesch during a period of research in Europe in 1891-92, initially in Monaco with Theodor Boveri, where he ‘discovered’ the centrality of the nucleus in development, and then at the Stazione Zoologica di Napoli, where he began a collaboration and established a friendship with Hans Driesch. In a classic paper published in 1892, which summarizes the results of his research on the annelid Nereis carried out in 1885-1890, Wilson exposed a theory of the interaction between hereditary characters and environmental conditions during development. These studies showed the plasticity of the ontogenetic processes, in contrast to the deterministic theory of Weismann and Roux, because, as shown by Driesch, the modification of the environment can change the pathways of morphogenesis, producing the same results through different pathways. However, in contrast to Driesch, Wilson did not consider the fertilized egg as ‘totipotent,’ as the number of possible choices is rigidly limited by heredity, which produces a ‘prelocalization’ of ‘formative substances’ in the cytoplasm even before segmentation. Each cell gets the same number and type of hereditary factors (the chromosomes), but differentiation is ‘modulated’ by the cytoplasm. As a consequence, development is not preformed but epigenetic. As suggested by another American embryologist, E.G. Conklin, embryogenesis is the result of two interacting factors: the heredity of an ‘ancestral tendency’ and an adaptive, actual tendency. The aim of the research on ‘cell lineage’ was to evaluate the relative role of these two factors.
Jacques Loeb
Another scientist who spent long periods of research and creation in Naples was Jacques Loeb, but his philosophical tendency was opposed to that of Driesch. If the latter accepted a vitalistic, Aristotelian biological philosophy, Loeb adopted a rigorous materialism, proposing a comprehensive theory of animal reactivity based exclusively on physico-chemical forces. In Naples Loeb studied regeneration, the possibility of controlling and regulating morphogenesis by ‘external means,’ that is physical and chemical agents (light, salts, acids, etc.). From isolated fragments of planaria Loeb was able to produce animals with two heads. On the basis of his experiments with the hydrozoon Tubularia he explained growth and regeneration with the accumulation of substances in certain regions of the embryo (growth factors). According to Loeb this was clearly in contrast to the importance of evolutionary theory in morphogenetic processes, and in favor of a purely mechanical explanation : ‘any theory of life must be based on our knowledge of the physico-chemical constitution of living matter and neither Darwin nor Lamarck was concerned with this’ (1916).
Loeb suggested a chemical explanation of life and his vision played a relevant role in biochemistry, which became the leading biological discipline in the first half of the 20th century.
In Naples Loeb also realized artificial fertilization, showing that the sea urchin egg can begin development after being exposed to an acid or to an increase in osmotic pressure. This experimental ‘chemical parthenogenesis’ had a strong effect on the scientific communities and on public opinion. Someone even suggested that women should avoid sea baths because of the risk of ‘chemical parthenogenesis’. Loeb successfully repeated this experiment with frogs, becoming the ‘father of the fatherless frogs.’ According to him, the proof of replacement of the mysterious ‘vital agent’ with a purely physico-chemical agent (sea-water concentration), ‘liberated the field of fertilization from vitalistic mysticism’ (1912).
Theodor Boveri
In 1889 the German biologist Theodor Boveri began his classic experiments in Naples on the hybridization of different sea urchin species, in order to establish whether the nucleus, the protoplasm, or both determine inheritance and development. Boveri fertilized enucleated egg fragments of one species of sea urchin with sperm from a different species, obtaining larvae that exclusively showed the characteristics of the paternal species, whereas using intact fertilised eggs produced larvae with characteristics from both parents. According to Boveri (1890) this experiment proved that the nucleus had a dominant role in inheritance.
Using a method of ‘double fertilization’ (polyspermic eggs), Boveri obtained ‘hybrid merogones,’ in which a tetrapolar spindle was formed. The separation of the different blastomeres was followed by an irregular distribution of chromosomes that produced abnormal development of the embryo. When the first four blastomeres were taken apart, at variance with normal embryos in which a larva is obtained from each blastomere, in the case of polispermic eggs some of the blastomeres developed normally, whereas others did not develop at all or became arrested during cleavage. Since Boveri could not detect any correlation between the number of chromosomes and the destiny of each blastomere, he concluded that the quality of the chromosome rather than number is the determining factor. From this he concluded that chromosomes are ‘individual’ and possess different qualities : ‘only a precise combination of chromosomes, probably only the totality of those contained within each pronucleus, represents the entire nature of the form of the organism.’ The embryonic development is therefore considered as the unfolding of the ‘qualities of the nuclei.’ Discussing his findings in relation to the newly rediscovered laws of Mendel, Boveri stated that individual chromosomes were the bearers of Mendelian hereditary factors, the first insight into the relation between genetics and cytology, the ‘chromosome theory of heredity.’ He also suggested that these ‘multipolar mitoses’ could be the causal factors in tumor formation (1903).
In Theodor Boveri’s hands the sea urchin egg also proved to be excellent material for the study of the nuclear-cytoplasmic interactions in development by his student Fritz Baltzer and is still widely used (Baltzer, 1967).
Otto Warburg
From 1908 to 1914, Otto Warburg spent several periods at the Station, where he carried out his first major independent work on the oxygen consumption which occurs when a sea urchin egg begins to develop after fertilization. In 1908 Otto Warburg, pushed by Loeb to study biological oxidation, began his study of metabolic changes during cell division by determining oxygen consumption. He made the classic discovery that upon fertilization the rate of respiration increases as much as six-fold. In 1909 he also discovered that iron is essential for the development of the larval state, inaugurating a new research program on cellular respiration, research that later won him the 1931 Nobel prize for the discovery of the cytochrome oxydase.
These results provided the basis for his future work on the metabolism of tumor cells, and are described in three of his very first papers (in Hoppe Seyler’s Zeitschrift für Physiologische Chemie) completed before obtaining his M.D. degree in 1911 from Heidelberg.
Otto Warburg’s work has a special prominence. His discoveries of the change in the respiration of the sea urchin egg as a result of fertilization inspired a new field of ‘chemical embryology,’ a line of research pursued at the Stazione Zoologica primarily by John Runnström and his school (Warburg, 1910). Runnström was indeed a frequent visitor to the Station almost up until his death. Furthermore, the ‘Warburg apparatus’ became the tool (and the nightmare) for two generations of cell physiologists and biochemists. The theory and practice of manometry was perfected by Warburg later, in 1920, and provided the key techniques for his later discoveries.
Thomas Hunt Morgan
Thomas Hunt Morgan (1866-1945) worked as an embryologist at the Stazione Zoologica around the turn of the century before devoting himself to genetics, to the creation of the Drosophila Group and to the development of ‘chromosome genetics’. At Loeb’s suggestion he had started working on regeneration, but later considered that this problem was too complex to be solved rapidly and thus decided to switch to inheritance, as a more suitable experimental problem, before returning to embryology at the end of his career. At Naples, where Morgan became friends with Driesch, his attention turned directly towards Driesch’s work on fragmentation and partial embryos and their impact on interpretations of development. In the heated debates on preformation and epigenesis, on Weismann and Roux’s mosaic or on Driesch’s regulative views of development, Morgan himself maintained a moderate position, sympathetic to Driesch but closer to the idea of an ‘organic continuity to explain development, as proposed by American embryologists. This tendency included Charles Otis Whitman and Charles Manning Child (1869-1954), who, in 1911, studied regeneration in Naples and produced the theory of the axial gradient, according to which morphogenesis is based quantitatively on a gradient of differentiation distributed along the axis of the embryo.
Through two World Wars. Crises and new beginnings (1915-1950)
World War I
When Italy entered into the First World War, Reinhard Dohrn and the German assistants had to leave Naples. Dohrn charged Federico Raffaele, a professor at Naples University who had been an assistant at the Stazione, with direction and established himself in Zurich, as a guest of the Zoological Museum, where the editorial office of the journals was also transferred. The Stazione, the private property of a German citizen, was placed under national control and its direction was assigned to Francesco Saverio Monticelli (1915-1924). In 1916 the Stazione was solemnly inaugurated as an ‘Italian Institute,’ under the responsibility of a national committee. The covered steamboat ‘Anton Dohrn’ was commandeered in 1917 and converted into a warship with the name ‘Salvatore Lo Bianco.’
At the end of the war, after a period of uncertainty marked also by nationalist demonstrations, Benedetto Croce, the Minister of Public Instruction, in a speech to the Senate on December 9th 1920, replied vehemently to the charges against Reinhard Dohrn and suggested placing the Stazione Zoologica again under the responsibility of the Dohrn family, the only way, according to the Neapolitan philosopher, to guarantee the institution its scientific links and its functionality. In the same session, the senator Battista Grassi, a professor of comparative anatomy and Rome University who had contributed a monograph on the Chetognats to the series ‘Flora und Fauna’ and who had spent several research periods at the Stazione, also spoke in favor of the solution proposed by Croce.
In October 1923 its legal status was redefined and the Stazione Zoologica became an ‘Ente Morale’ (a semi-private institution), under the surveillance of the Minister of Public Instruction, directed by an administrative board which was chaired by the mayor of Naples. Rinaldo Dohrn was designated ‘managing director and administrator.’ The relative acts would be given final approval in 1933.
The new institutional structures maintained the ‘international status’ of the Stazione, fostering the contacts among scientists of different countries and encouraging their presence in Naples, thanks to total freedom of research and the assistance of a highly effective technical structure, which created the best working conditions in terms of material and laboratory supply and in terms of cultural atmosphere. In this context the contradictory but meaningful concept of ‘Italian internationality’ was also introduced. The main aim of the Stazione Zoologica however, as articulated by Giuseppe Colosi in 1930, remained ‘to give hospitality to the scientists and assist them in their own researches, largely by offering tools and materials.’ The only specific and autonomous objective of the institute was the completion of the large systematic series on the Flora and Fauna of the Gulf of Naples.
These politics soon bore new fruit. Ignoring political and economic differences, many governments as well public and private institutions started to rent research tables again. One can underline two important financial contributions from the Rockefeller Foundation in 1924-1929 (respectively 10000 and 7000 dollars) and the renting of research tables by the newly born Soviet Union. The demographics of scientists at the Stazione was similar to that of the years before the war: roughly one third Italians, one third Germans, and one third from other countries, in particular the United Kingdom, Austria, Sweden, Switzerland, and the USA. Additionally the departmental structure remained practically the same. At the head of the zoological department one finds Umberto Pierantoni (1915-1924), later director of the Zoological Institute of Naples University, Marco Fedele (1924-1930) and Giuseppe Montalenti (1939-1944). The department of physiology was directed, successively, by Filippo Bottazzi (1915-1925), Enrico Sereni (1926-1931), and Luigi Califano (1931-1935).
Lines of research in the interwar period.
The scientific activities at the Stazione in the interwar period indicate a great continuity with the previous period. Systematics remained a priority and in this field one should mention Paolo Enriques’ discovery, during the course of a long inquiry unfinished due to the death of the researcher, of a new species of plantonic radiolars, a collective work on plankton and benthonic life in the Gulf of Naples, and the completion of the great monograph on teleosteans, initiated by Salvatore Lo Bianco in 1910 and finished in the 1930s by Umberto D’Ancona, Leonardo Sanzo, Silvio Ranzi, and Giuseppe Montalenti among others. Also worth mentioning, for their zoological value, are the beautiful anatomical specimens prepared after WWI by F.S. Monticelli.
The aggregation points of research in those decades are physiology of the embryo, developmental mechanics, regeneration, the analysis of the biochemical components of marine organisms and their variation during embryogenesis, in particular biochemical gradients, bioluminescence and photogenesis in marine organisms, and hereditary symbiosis. At the same time, the richness of the flora and fauna of the Mediterranean, and the great diversity of superficial and abyssal forms favored the development of a new ecological approach, centred on alimentary chains, and on organism complexes in relation to the biological and physico-chemical environment. Among the Italian scientists working at Naples during the interwar years, one should recall Silvio Ranzi, who spent fourteen years in Naples, often returning later to study developmental mechanics of cephalopods, axial gradients and the biochemical aspects of development in several species, a field of research also practised by Federico Raffaele, together with the morphology of the circulatory system of fishes. It is worth mentioning the research carried out by foreign guests such as Fritz Baltzer (Berne) on sexual determination in Bonellia, Kasimierz Bialaszewicz (Warsaw) on the chemical determination of the egg, Rhoda Erdmann (Berlin) on the in vitro culture of animal tissues, Richard Goldschmidt (Berlin) on the genetical basis of embryology, Ross G. Harrison, who studied, in collaboration with Pasquale Pasquini (Rome), the experimental problem of embryonic determination; Curt Herbst (Heidelberg) again on determination of sex, Ernest Everett Just (Washington) on fertilization; and finally John Runnström (Stockholm) on experimental study of development.
And, as in the previous periods, the Stazione was the ideal place for the experimentation of new technologies applied to research, such as the use of cinema for the observation of experimental embryology by Friedrich Kopsch (Berlin), the first experience of scientific cinematography by the Istituto Luce and the research supported by Kodak on gels of animal origin for film.
The embryonic organiser and biochemical embryology
Once again, the main scientific thrust at the Station came from experimental embryology. The turning point was the publication of the classic paper by Hans Spemann and Hilde Mangold in 1924, that would come to represent for the following two decades the equivalent of the classic experiments by Roux and Driesch. In a series of experiments initiated in the first years of the 20th century, Spemann had shown that the formation of the lens of the eye in the ectoderm of amphibians was induced by the optical cup, a part of the cerebral tissue. In the following years, there was a long series of experiments, many realized in Naples, where Spemann worked quite often, always accompanied by a substantial group of assistants.
Spemann observed that the region of the blastopore was determined earlier than more distant regions. Furthermore, there appeared to be an additional ‘organizing action’ of the dorsal lip region when transplanted to another embryo. These suggestive findings set the stage for the definitive experiments by Spemann and Mangold (1924) which proved that a piece of the dorsal lip of the blastopore, in the process of gastrulation, exerts an organizational effect upon surrounding tissues in such a manner that it causes a secondary embryo to be formed if implanted in an indifferent place on another embryo. The grafted piece of the blastopore was therefore designated as an ‘organizer.’
The discovery of the ‘organizer’ and the revelation that it was not species specific produced twenty years of intensive and largely fruitless research on its chemical nature, a biochemical version of the classical ‘developmental mechanics’. That gave rise to the new research program of ‘chemical embryology’, aimed at the chemical characterization of the ‘organizing or morphogenetic substances’. This program used the already established biochemical and physiological techniques and also new cytochemical methods to study the distribution of the different chemical species in the fertilized egg, in segmentation and differentiation. The main goal was to find the biochemical basis of the organizer and to explain its morphogenetic action in physico-chemical terms.
Chemical embryology flourished in the 1930s and ‘40s, finding a first synthesis in Biochemistry and Morphogenesis, published by Joseph Needham in 1942 and in the first book by Jean Brachet, Embryologie chimique, published in 1944. Both books were centered on the hope of finding a unity between biochemistry and morphogenesis through a chemical explanation of morphological discontinuities during development and differentiation.
This search parallels the more general program of biochemistry to find a unified explanation of the fundamental aspects of living systems: function, form, development, and heredity. As in the previous period, a fundamental dualism existed between a chemical explanation (upward) and a morphological explanation (downward) of a biological function. The first attributes to chemical agents the causative role in morphogenesis (organizer, chemical gradient, etc.) whereas the second gives priority to the morphological aspects (cell interactions, spatial distribution, etc.). The central problem remained the chemical nature and role of ‘organizers.’ In this context, an important result was obtained in 1939 when Albert Tyler demonstrated that the metabolic processes in the egg and its growth are distinct and relatively independent from morphogenesis and differentiation. Enhanced glycogen breakdown seemed to correlate to morphogenetic movements but not to neurogenic induction and differentiation. In such a way the problem of respiratory metabolism was separated from the problem of morphogenetic stimuli. Respiratory processes are essential for the life and growth of the embryo but they are not the cause of morphogenesis.
Systemic biology and ecology
Another relevant theoretical transformation in biology took place in the 1930s. As a consequence of the origins of the synthetic theory of evolution, which shifted interest towards population level, much attention was devoted to the systemic aspects and to the observation of complex systems in nature. Until that period attention of the marine biologists was mainly devoted to the description of new species and to the experimental study of physiological functions and embryological development. During the 1930s a new interest in the ecological relationships between species and on the physico-chemical conditions that shape the behavior of large animals and vegetal communities emerged. This is the origin of the ‘new ecology’ as a discipline and of a new sensitivity towards the conservation of the biosphere.
In Naples, in this new context, Raffaele Issel studied the quantitative and qualitative distribution of plankton, its daily and seasonal variations, whereas Paul Buchner published his original observations on symbiosis and its ecological role.
Biochemical and physiological research
In the interwar period biochemical and physiological research continued to play a relevant role in the scientific life of the Stazione. During this period both Hammerling (1934) and Schulze (1939) pursued studies of Acetabularia species present in the local waters. These studies were part of the pioneering work developed in their native Germany, which established Acetabularia as an important physiological model and tool in the studies of cell biology. Albrecht Kossel and The Svedberg, the founders of nuclei acids chemistry, spent long periods of research at the Stazione, as did Jean Brachet, who at that time developed the cytochemical methods for localizing the nuclei acid in the cell and in the embryo. In the physiological domain, Albrecht Bethe (Frankfurt) continued his research on muscle physiology and morphology, and Otto Meyerhoff, a future Nobel prize winner, studied muscular metabolism, concentrating on the chemistry of the stimulation of the muscular fiber. Within the same physiological framework, Karl Heider studied the nervous system of annelids, Johann Jacob von Uexküll (Hamburg) carried on with his research on the experimental psychology in echinoderms and Alfred Kühn (Göttingen), in 1927-1929, developed the same field using cephalopods as experimental material.
Altogether, during the 1930s and 40s the effort to offer a chemical explanation of the nervous system and behavior gained increasing relevance. In 1935, Zénon-Marcel Bacq and Francesco Paolo Mazza demonstrated the presence of acetylcholine in the optical ganglia of the Octopus, and identified the substance chemically. Earlier, in Germany, acetylcholine had been identified as the Vagusstoff by Otto Loewi. The work of Bacq and Mazza is a keystone in the history of chemical nervous transmission, and provided the first direct demonstration of its existence in nervous tissue. Bacq, a pharmacologist from Belgium working with Francesco Ghiretti, employed cephalopods to advance endocrinology, using adrenaline-like agents. It should be mentioned that Ernst Florey started his research career in the laboratory of Bacq and ‘Ghiro’, as Ghiretti was called.
Jerzy C. Alexandrovicz was a professor of ophthalmology and, from 1937, Under-secretary of State in the Polish Ministry of Education. Alexandrowicz’s meticulous neuro-anatomical studies of crustaceans and cephalopods using methylene-blue staining techniques have earned him a special place in neurobiology. Perhaps his most important discovery, rivaled only by John Z. Young’s discovery of the giant axons of squid, were the stretch receptor organs of crustaceans. The crustacean stretch receptor preparation thus became an important tool in the investigation of inhibitory synaptic transmission.
The Stazione Zoologica has been instrumental in yet another important advancement in the field of neurobiology: the discovery of what has become known as ‘neurosecretion,’ the elaboration and secretion of hormones by nerve cells. In 1928 Ernst Scharrer, assistant under Karl von Frisch at the Zoological Institutes of the University of Munich, was granted the use of a research table at the Stazione Zoologica at Naples. As he stated in a letter to Reinhard Dohrn, he wanted to fix the brain of many species of fish, and, ‘if possible to investigate, with the aid of methylene blue staining, the innervation of the epiphysis.’ He discovered that certain neurons in the midbrain show evidence of secretion, confirming earlier findings of Carl Speidel (1919). With material from Naples, Scharrer continued his studies on these ‘neuroglandular’ cells of fishes and extended these studies to higher vertebrates where the same cell type was found.
What may be called the modern epoch of cephalopod research at Naples was begun by Enrico Sereni, who conducted many experiments on the chromatophores and salivary hormonal secretion, summarized in a long article of 1930. Sereni, head of the physiology department, was a brilliant physiologist and produced ingenious experiments on nervous and humoral correlations of the activity of the chromatophores, a subject studied at Naples also by Nikolai K. Koltzoff (Moscow) between 1924 and 1927.
Sereni also worked on the peripheral nervous system in cephalopods, in collaboration with the young J.Z. Young, who founded another line of research, followed for many years almost uninterrupted at the Station. Working with Enrico Sereni on the Octopus, Young discovered by chance the giant axon and this became a fundamental tool in exploring nerve conduction. Until Sereni’s early death he and Young were collaborating on a study of regeneration of the stellar nerves, which was completed by Young. In the course of this study, Young was attracted to a small orange spot at the hind end of the stellate ganglion of Eledone. He cut sections of it and proved that this was a hollow vesicle into which passed a number of projections, apparently of nerve cells. After discussion with Sereni it was named the epistellar body (Young, 1929).
Great developments have flowed from the study of the epistellar body. Having found it in octopods, Young naturally also made sections of the stellate ganglion of decapods. No epistellar body was there but he found instead the giant nerve fibers that became another ‘right tool for the job’. The sections of the brains and giant fibers of squids and cuttlefishes that were made by Young at the Stazione in 1936 showed the possibility of studying higher nervous activities, such as learning and memory.
World War II
During the war years, the Stazione remained virtually closed. A few scientists and technicians remained at the Institute in order to assure functionality of the installations, primarily the aquarium. Giuseppe Montalenti, with the help of a few technicians, among them Alberto Pannone, Giuseppe Della Morte, Giacomo Fiorillo, and Vincenzo Serino, was able to avoid military occupation of the buildings, and the destruction and displacement of the instruments. The library also escaped destruction and heavy losses, because it was transferred to a small village inland, Pontelandolfo, in October 1943.
When the German army left, the Villa Comunale was occupied by units from the American army. Personal passes were granted to the staff of the Stazione Zoologica in order to reach the Institute. Due to the cooperative spirit of the American officers of the occupying units, the life of the laboratory was allowed to continue in the midst of military business, almost without interference, only a few rooms of the building being occupied.
The Allied Military Government granted substantial financial help, in anticipation of funds from the Italian Government, in order to pay salaries and to cover current expenses. Already, in the first months of 1944, the Royal Society of London granted an extraordinary contribution of £1000, important help and a symbolic gesture which marked the scientific relaunching of the institute and highly improved its credibility with the Allied headquarters.
The Aquarium was opened to Allied troops as early as December 10th 1943, and the income derived therefrom was added to the regular funds. The Villa was left by the occupying units on May 1st 1944, and the activity of the Stazione Zoologica returned almost to normal under the leadership of the director Rinaldo Dohrn, who returned from Sorrento, where he had found accommodation after the loss of his house, assisted by the constant cooperation of the Allied Military Authorities. And when the Library was taken back to Naples and put in order in the early summer of 1944 this was the signal for a new start for the prestigious institution.
After the liberation of Rome on June 4th 1944, contacts were established with the Italian Ministry of Public Instruction, and the Italian National Research Council. Both bodies very consistently helped the Stazione Zoologica. The latter founded there a ‘Center for Biological Studies’ under the directorship of Giuseppe Reverberi, for the purpose of granting fellowships to biological students. Postgraduate students were admitted to the Stazione for the first time in 1947.
At the end of the war Switzerland, Sweden, England and the USA renewed some of the ‘working tables’ they had rented before the war, thus beginning again the international cooperation without concern for ideologies and nationalism, the most outstanding feature of the Stazione Zoologica. From May 1945 international scientific activities started again and at the end of that year the occupied research benches already numbered 15, increasing to 31 in the following year and stabilising at an average of 40 between 1947 and 1953.
In the summer of 1946, the president of the National Research Council, Ross Granville Harrison set up a committee, chaired by Ernst Scharrer, to aid the Stazione Zoologica. The committee, which included as members E.G. Conklin, Mrs. E.B. Harvey, R.G. Harrison, S. Hecht, L.H. Kleinholz, A.R. Moore, and H.H. Plough, met for the first time on August 14th,1946 at Woods Hole to decide on a program-in-aid to assist in the re-establishment of the Stazione Zoologica at Naples as an international center of biological research. The program envisioned the establishment of additional American research tables, a funding-drive to improve and enlarge the library, and a shipment of food for the ‘Mensa,’ the institute cafeteria. Further plans concerned the modernization of the laboratory facilities and research equipment. The activities of this committee led to contributions by the Rockefeller Foundation and UNESCO. The committee advised the Stazione Zoologica in matters of library acquisitions and the purchase of research equipment.
After 1950 the newly established National Science Foundation, Lilly Endowment, Inc., and the Rockefeller Foundation enormously increased American engagement with the affairs of the Stazione Zoologica.
The scientific prestige of the Stazione and its location made it an ideal place for scientific conferences that, in the decade after the end of the war, marked the renewal and continuing development of scientific research. This was also an opportunity for Italian scientists to enter the international network and get in touch with new research projects and a new generation of biologists. The Stazione hosted important symposia on embryology and genetics, on mutagens, and on neurosecretion. In 1951 a meeting on the application of X-rays to the study of biological problems took place, focusing primarily on the submicroscopic structure of the protoplasm. During this symposium Wilkins reported on the new direction of biophysical research in John Randall’s laboratory in England. Wilkins went to Naples because he wanted to study, in collaboration with Bruno Battaglia, the sperm of the Sepia in order to investigate with X-rays if the genes were arranged regularly along the head of the sperm ion crystalline array. At the conference, Wilkins showed an X-ray diffraction pattern of crystalline DNA and that stimulated J.D. Watson, who was also present, to begin his work on nucleic acids that brought him in collaboration with Francis Crick to the discovery of the double helix.
On the initiative of Ernst Scharrer and Wolfgang Bargmann, the Stazione Zoologica hosted, in 1953, the first International Symposium on Neurosecretion, the fourth international symposium held at the Stazione Zoologica after the end of the war. This event has a special place in the history of biology because it was this symposium that established the concept that neurons produce hormones and that neurosecretion is an essential feature of the chemical control of animal development and function.
A small institute in the era of the big science (1950-1967). The end of the ‘Dohrn era’. The ‘commissario’.
After WWII science, and in particular biological and medical sciences, evolved at an ever increasing pace and on an ever expanding institutional and economic scale. The number of scientists began to increase exponentially, and the costs of apparatus and materials increased rapidly. The era of romantic science had passed. The ideals of science changed from an individual enterprise linked to individual dreams and projects to a full-scale collective effort. When, in 1954, Peter Dohrn took over the directorship from his father Reinhard, the Stazione Zoologica was confronted with a difficult choice: either to invest heavily and try to cope with expanding costs and management difficulties of the new ‘big science’ or to remain a small appendix of other institutions, a marine facility for research groups established elsewhere.
The Stazione chose the first option, launching ambitious projects for the renewal of its technical structure, buying new expensive laboratory equipment in order to satisfy the needs of a new kind of biological research. A new library was built, in a new five floored structure located between the two old buildings. Two new ships were also made available (‘Federico Raffaele’ (1955), motorboat, 10 m; ‘Rinaldo Dohrn (1959), motorboat, 14 m).
The Stazione remained a reference at the international level, and the ‘Naples experience’ continued to be considered a necessary step in the training of a biologist hoping to specialize in embryology, cytology, or marine biology. This was particularly true for Italian science, as the Stazione was the best way to get experience with advanced biological research. The international character of the institution was maintained thanks to several collaborations, first of all with Woods Hole MBL and the British marine biology laboratories. In the mid 1950s the number of rented research tables again surpassed the figure of 50 per year, increasing to 86 and 88 in the years 1958 and 1959.
However the traditional ‘bench system’ produced inevitable fragmentation and lack of continuity among the research programs and an expensive dispersal of experimental apparatus. Each guest scientist arrived at the Stazione with his or her own research project and needed the best conditions for realizing it, therefore demanding special conditions and tools. As such, it was difficult to afford the increasing costs, which often could not be shared with other projects and distributed over a longer period of time. That produced internal conflicts, a difficult financial crisis and the need for a modern status for the scientific and technical personnel of the Stazione.
The Stazione Zoologica needed a new administrative and institutional structure, in order to obtain a solid and permanent financial basis. For almost a century the Italian government had continued to recognize the unique character of the Stazione, a private enterprise in an international context. But in the new social context of biological sciences, this was not possible any more. In 1967 the Italian government named a ‘Commissario Straordinario’, charged with administration and direction, a post successively held by Mario Pantaleo (1967-1970), Guido Bacci (1970-1975) and Alessandro Barlaam (1975-1976). A Scientific Committee was also created, composed by Guido Bacci, Giuseppe Reverberi and Aldo Spirito. The Italian government took on more and more the burden of financially supporting the ‘old lady,’ a prestigious institution in need of new blood, and the Stazione became part of the Italian national science administration, but at the same time retained very large autonomy.
In the two decades following WWII the structure of the Stazione demonstrated continuity with the previous tradition. The zoological department was directed by Alberto Monroy (1944-1949), who switched in 1949 to the biochemistry department, Guido Bacci (1949-1955); Pierre Tardent (1956-1961), Andrew Packard (1959-69) and Rainer Martin (1963-1973). The physiology department was directed by Francesco Ghiretti (1955-1961).
Lines of research in the postwar period, 1945-1967
Embryology
Following World War II the ‘quest for the organizer’ continued, turning for a time to the nucleic acids, in particular ribonucleic acids, as effectors. However, the new generation of scientists that emerged after WWII was cut off from the research traditions of the 1930s. The problem that had fascinated embryologists of the ‘classic era’ did not stimulate young scientists and the shock produced by Johannes Holtfreter’s observation on the autodifferentiation of the ectoderm made the problem of chemical induction an unattractive field.
The monumental accumulation of information at both the morphological and the biochemical levels on the patterns of development in animals and plants, as well as on cell and tissue interactions, seemed to be useless and embryology appeared to be in decline. Embryologists started to look at other ways of explaining differentiation, considering it as a complex chain of events rather that as a single process (induction).
Before the origins of molecular biology in the 1950s, the relation between the nucleus and the cytoplasm, between heredity and embryogenesis, was interpreted biochemically. The chemical heterogeneity of the cytoplasm, in particular its proteic composition and the quantitative gradients, was explained as a result of variable synthesis of proteins along the gradients. Between 1945 and 1953 the theory of plasmagenes or ‘self-reproducing units’ (genes, virus, cytoplasmic microstructures) seemed to provide a unified explanation of the self-replicating phenomena in microbiology, genetics, biochemistry and medicine, thanks to a generalized replicative property attributed to different chemical and morphological entities, composed mainly of proteins and nucleic acids. This was particularly true for embryologists, in search of a causative explanation of the nucleo-cytoplasmic relationships during differentiation (Brachet, Montalenti). The most accepted view considered nucleic acids as a necessary part of self-replicating structures (ribonucleoproteic granules), ‘organized cytoplasmic particles,’ analogues to the genes, plasmagenes and viruses, and responsible for differentiation and cell metabolism. These self-reproducing particles were supposed to emanate from the nucleus but retained their replicative autonomy in the cytoplasm. Particular nucleoproteic granules, produced by the nucleus and then distributed along the morphogenetic gradient, able to self-reproduce, would be the seat of protein synthesis and hence differentiation. Notwithstanding the hypothetical nature of such ‘plasmagenes,’ the idea of a ‘chemical migration’ of particular molecular species from the nucleus to the cytoplasm remained a guide for research, linking together genetics, cytology and embryology.
The phrase ‘developmental biology’ became popular in the late 1950s when embryology was in a transition period, characterized by a revival of interest in the problems of differentiation and embryonic development, which were again considered to be at the frontiers of biology. The interest started to switch from chemical organizers to subcellular and cell-cell interactions, the exchange of ‘information’ between nucleus and cytoplasm and the genetic control of morphogenesis. The attention of chemical and molecular embryologists centered on cell surface interactions in fertilization and morphogenesis, on the exchange of effectors via cytoplasmic bridges, specialized junctions or other mechanisms, particular cell microstructures (microtubules), and on signals generated at the membrane, transmitted via specific receptor sites and intracellular mediators like calcium and cyclic AMP to specific genes.
The enthusiasm generated by the discoveries in the field of molecular biology, especially after the proposal by François Jacob and Jacques Monod in 1958-1961 of the ‘operon model’ as a molecular explanation of the control of genetic expression, produced high hopes of finding the solution to the problems of differentiation thanks to extensive knowledge available on the genetic regulation in microorganisms. However, the complexity of the eukariotic cell posed new problems and molecular embryologists and developmental biologists had to recognize that ‘the egg is not a glorified Escherichia coli’ (Monroy, 1970).
The creation of a new research project, molecular developmental biology, was the result of this new conscience. The main change arose in the definition of the experimental problems to be solved in the laboratory. The traditional problems of development (e.g. polarity, gradients, induction, determination, etc.) had to be reinterpreted in the light of the new explanatory model based on gene expression and its temporal control during embryogenesis. The central problem in molecular embryology consisted in the understanding of the molecular and cellular mechanisms by which expression of the genome results in the formation of a differentiated embryo. This problem needed the combination of biochemical cellular and genetic methods. The experimental design was aimed at investigating the differential expression of the gene and the nucleo-cytoplasmic interactions in the course of development. Nuclear transplantation, already established in classical embryology, provided a powerful tool in analyzing the processes by which cytoplasmic factors and intercellular interactions regulate gene expression. Finally, the new techniques of molecular biology allowed the isolation and the characterization of the genetic determinants, markers and mutants of the successive developmental events. New animal models had to be found for these kind of experiments, as organisms such as the sea urchin, which for many decades had been the favorite experimental tool for embryologists, were completely unknown from a genetic point of view. Amphibians became the experimental jewel of developmental biology, replacing marine organisms. Only in the following years, could new marine animal models be produced to study the genetics of development. The new generation of biologists working at the Stazione Zoologica faced new technical and scientific demands.
Embryologists contributed relevant concepts and information to the new field of molecular developmental biology. A. Tyler, T. Hultin, J. Brachet and A. Monroy showed that the control of the genetic program seems to be already predetermined in the non-fertilized egg, as the ribosomes, the seat of protein synthesis, are accumulating already during oogenesis, the expression of the ‘messengers’ they contain being blocked by some proteins, eliminated at the moment of fertilization. This new theoretical framework seemed to explain the already old observations by J. Runnström and S.O. Hörstadius, that fertilization induced the formation of proteolytic enzymes. The study of fertilization also suggested new lines of research in the 1950s and ’60s. Monroy’s experiments in Naples and Palermo showed that DNA and protein synthesis starts just a few minutes after the penetration by the sperm of the egg, as showed by the incorporation of radioactive residues in the proteins and ribonucleic structures at the beginning of development. The traditional concept of gradient, proposed in the interwar period by Child and by the Swedish-Neapolitan school of Runnström and Hörstadius, was equally integrated into the new framework. In particular, Hörstadius showed in the sea urchin that at the stage of 16 cells, one can easily recognize in the sea urchin an animal pole and a vegetal pole, as the egg is composed of three different kinds of cells: eight of medium size at the animal pole whereas the vegetal pole is composed of four big cells and four small cells (micromeres). The isolated animal pole cannot produce a gastrula (invagination), unless one transplants into it the micromeres. A greater number of micromeres needs to be added to a fragment from the animal pole, in order for normal development to occur, than for an animal fragment taken from more towards the vegetal pole. These experiments demonstrated the existence of two quantitative interacting gradients going in opposite directions between the animal and vegetal poles. The existence of quantitative gradients leaves open a problem already perceived by Hans Spemann in 1938, that is the difficulty of understanding how qualitative differences may arise from quantitative ones in the different regions: the gradient must be considered as continuous, whereas the series of formations whose differentiation would be determined by that gradient is discontinuous. This problem would be solved after the fusion of embryology and genetics and the origin of ‘developmental biology,’ when it was proven that the quantitative gradient in concentration parallels a qualitative difference in the content of information of the genetic program.
Neurobiology
The incredibly large size of the ‘giant synapses’ between second and third-order giant axons made possible important developments in neurobiology. Outstanding achievements included the placing of an internal electrode and the emptying and refilling of the axon by A.F. Hodgkin and A.L. Huxley. These investigations provided the data that enabled them to deduce the equations of the ionic exchanges involved. After the pioneering research of Otto Loewi (Nobel prize winner in 1936) which showed that the synaptic transmission was mediated by acetylcholine, Bernard Katz and Riccardo Miledi from the University College in London came to the Stazione Zoologica in 1966 to investigate the relationship between calcium and transmitter release. Thanks to the special usefulness of the fibers that allow monitoring by electrodes on both sides of the membrane, Miledi and Katz were able to study, at Naples, the two sides of the synapse. Their experiments have become classics: they proved that extracellular calcium is essential for transmitter release to occur and that calcium ions enter the nerve terminal when this becomes depolarized by the incoming presynaptic action potential.
Numerous researchers have used this property for studies of membrane transport at Woods Hole, Plymouth, and Naples.
Octopus once again provided many opportunities for studying learning and memory mechanisms, and there was the apparently inexhaustible year-round availability of animals in Naples. The Posillipo fishermen were able to bring in 20 or more octopuses a day in excellent condition and the Stazione generously provided space for specially created tanks, built in a special space with funds from the British Science Research Council. This made possible, at the Stazione Zoologica, the establishment of a large ‘cephalopod facility’ which, on its best days, included more than two hundred tanks in which as many octopuses could be individually housed, maintained and observed. Together with several collaborators, especially Bryan Boycott, Martin Wells, and John Messenger, Young mapped the neuronal circuits of the octopus brain and through ingenious training experiments he explored the learning ability of these animals. These studies led to new concepts of the neuronal mechanisms underlying memory.
Botany
From the very beginning, botanical research had been an important component of the research conducted at the Stazione Zoologica. The first researchers at the Naples Institute were German guests who proceeded with descriptive, ecological and taxonomic studies and provided a foundation for later physiological, cytological, life cycle, and biochemical studies of marine flora, conducted at the Stazione. After having been appointed to the marine botany department in 1912, Georg Funk realized several periods of research in Naples, producing three major publications on Mediterranean algae and contributing ecological work, giving the botanical research at Naples a continuity which extended into the late 1950’s. The lasting impact of marine botany has been the contribution of an ecological dimension from which other studies grew and found support. Funk’s monographs (1927, 1955) with extensive description of algae associations, reproductive cycles in nature and distribution, remained a benchmark for algae research in the Mediterranean. Funk’s observations also served as a basis for quantifying changes in natural populations in the Gulf of Naples where increased urbanization and industrial development was strongly affecting coastal waters. An important aspect of his research was the sustained effort over a 47-year period of research at the Stazione to establish a strong ecological perspective in the study of marine algae.
However, it was not until the early 1950’s that botanical studies resumed full activity in research, progressing with ecological studies of algae distribution, algae cultivation, and work dealing with various aspects of Acetabularia metabolism.
From 1960 onwards, the marine botany laboratory assumed a different direction, returning to a physiological-ecological orientation.
In the late 1950’s, a marked change occurred in the marine botany department with the permanent assignment of Dr. Kurt Beth, of the Max Planck Institute, as head of the algae laboratory in Naples, who remained there from 1956 to 1971. With a stronger emphasis on ecology during the decade between 1960 and 1970, botanists began to discuss scientific questions and concerns regarding the marine environment. The Benthic Ecology and Biological Oceanography laboratories were formed to foster ecological studies in the broad sense, including research on the physical-chemical factors as well as plant and animal communities, in particular the benthic ecosystems.
Laying the foundations for a new development. Alberto Monroy. The transition to a public research institute. (1967-1987)
The new organizational scheme of the Stazione Zoologica, established in 1967, took some time to be applied, but finally, in 1976, a new scientific director was appointed, Prof. Alberto Monroy, who was given the difficult task of re-establishing the former prestige through new international ties, grounded on a solid internal scientific program. Alberto Monroy, at the time the director of the Laboratory of Molecular Embryology established in 1969 by the National Science Council at Arco Felice, was a well know and respected scientist. He knew perfectly well where biological sciences were moving, was able to clearly set scientific priorities, and he was conscious of the need to secure modern laboratory facilities, to be staffed with experienced scientific and technical personnel.
His task as director became to breathe new life into the ‘old lady,’ as the traditional way of functioning was a product of its time and was unable to cope with the new demands in scientific research. The number of scientific guests from countries outside Italy was only 48 in 1978, reduced from 130 in 1960. This was due to the fact that administrative and financial difficulties made it difficult to work effectively in Naples. Moreover, the new technical facilities in preservation and transport of biological material made it possible for scientists to have perfectly good laboratory facilities at their home institutions. The ‘scientific tourism’, that was an essential and progressive character of the biological sciences in the first half of the 20th century, became less essential and sometimes useless. New ways of increasing international collaboration were needed. The richness of the Mediterranean Sea continued to provide superior facilities for collecting and holding marine organisms and the Stazione Zoologica continued to offer some of the best conditions for scientific research : a unique animal supply system, a marvelous library, modern research facilities, and, last but not least, friendly support from the direction and the technical staff. Even in the era of molecular biology the variety of the organ systems of marine organisms continued to offer a greater diversity of form and function, necessary for the understanding of the fundamental properties of living systems. Many relevant 20th century research projects which characterized the biological revolution of the 1950s and 1960s were in fact grounded on marine organisms: vision, memory, chemical transmission, protein structure and function, hormones and chemical messengers, and genetic control of embryogenesis.
This was the major challenge at the end of the 1970s and at the beginning of the ‘80s. The ensemble of scientific problems that had characterized the life of the Stazione for a century (fertilization and development, induction and morphogenesis, systematic and evolution, memory and nervous transmission, botany and ecology) had to be reinterpreted in terms of molecular biology and developmental genetics thanks to new conceptual tools (information, program, code, gene regulation and expression) and new experimental apparatus (electronic microscopy, electrophoresis and ultracentrifugation, macromolecular chemistry, and then molecular engineering). What was needed was a new framework for international co-operation, an effort to bring the frontiers of research into the scientific life of the Institute, increasing the participation of local scientists in international scientific joint-programs. During the late 1960s the Stazione Zoologica underwent a substantial structural change. By that time the ‘table system’, which had been the structure guaranteeing ‘internationality’ for almost 100 years, was discontinued and the focus on guest research changed to intramural staff research. But, only the research focus changed, not the international nature. If until 1968 the internationalism of the Stazione Zoologica had been based on the table-system, this character was guaranteed during the following years through joint programs and collaborations with foreign colleagues and institutions. The leading scientific institutions continued to support the Stazione Zoologica in the new context. The Deutsche Forschungsgemeinschaft assured its share of support even beyond the obligatory rental of research laboratory space.
The structure of the Stazione was largely reorganized and the scientific and technical structure showed a great adaptability to the new conditions. What in the first century of its life had been a duty in order to cope with the demands of scientific guests, now became a way to build up new scientific projects and increase international co-operation.
The traditional departments remained, as the main scientific focus at the Stazione in the 1970’s and early ’80’s remained zoology, biochemistry and developmental biology. Others departments were added in order to respond to new scientific and technical demands. Ecology became a focal point for the new development, as a scientific discipline, and the Station remained a focal point for any program of Mediterranean ecology and this competence was later used in other ecological contexts, including the Antarctic.
The department of Ecology, headed by Gioacchino Bonaduce (1970-1971), Francesco Cinelli (1971-1981) and Eugenio Fresi (1981-1982), was located in the summer home of the Dohrn family, the ‘Villa Acquario’ on the island of Ischia, that had been transformed into a laboratory, also thanks to the financial support of German foundations. The philosophy underlying this new ‘ecology group’ was that the expertise of specialists from different areas should be applied to broad integrated research programs designed to study the various physical, chemical and biological processes within the Gulf of Naples, and to investigate how these processes interacted with one another.
The other departments during this period, including the more traditional ones, maintained their central role: Zoology (Rainer Martin, 1973-1975; Gian Carlo Carrada, 1975-1982), Biochemistry (Luisa Tosi, 1972-1978; Giovanna Nardi, 1978-1982), Marine Botany (Kurt Beth, 1973-1982), Cell and Development Biology (Alberto Monroy, 1976-1982), Neurobiology (Rainer Martin, 1975-1981; Amedeo De Santis, 1981-1982), Ultrastructural Biology (Ermino Muzii, 1967-1971).
As director, Alberto Monroy was first able to achieve fundamental results, avoiding the risk of transforming the Stazione into an ecological monitoring or administrative center, and pushing even more the need to develop basic science. The scientific life of the Stazione became increasingly lively, in a context of permanent scientific collaboration. In his own research field, embryology, he started a new research project, directly linked to new trends in the discipline : the study of the molecular biochemical mechanisms of fertilization and development, in collaboration with his previous laboratory of molecular embryology in Arco Felice, and in particular with Jean Brachet.
Alberto Monroy was also conscious of another great tradition of the Stazione, its cultural atmosphere. He therefore placed new emphasis on the organization of public lectures, symposia and summer courses devoted to the history and philosophy of the life sciences.
However, Alberto Monroy remained fundamentally a ‘laboratory man,’ a creative scientist not always conscious of administrative problems. It was difficult to change the objectives and the functioning of the whole institute and to bring in new blood in the different research projects. The growing financial and institutional constraints became a major difficulty for the scientific life of the Stazione. A new institutional and financial leap was needed.
In 1981 Sebastiano Genovese was appointed director, seconded by a board of Administration and a Scientific Advisory board; he remained in the post until his tragic death in 1982.
A new development. From the ‘permanent congress’ to an autonomous research institute. Gaetano Salvatore (1987-1997).
In 1982 the Italian Parliament approved a new special law for the Stazione Zoologica and the government assured an increased annual budget. This law juridically recognized the Stazione Zoologica as a ‘Special Scientific Institution’ of public interest, under the supervision and control of the Ministero della Ricerca Scientifica e Tecnologica, which meant that the Stazione had to observe the rules laid down for public organizations governing personnel and the expenditure of funds. However, the Stazione Zoologica maintained complete autonomy concerning all other decisions related to the running of the Institute and scientific policy. That same year (1982) the institute was renamed ‘Stazione Zoologica Anton Dohrn’ after its founder, a gesture which should be considered not so much as in recognition of Dohrn’s achievements, but rather, or also, as an acknowledgement of his aim of furthering the frontiers of knowledge in a research laboratory free from national, philosophical or disciplinary limitations. Ten years later the Stazione Zoologica was declared an ‘Istituto di ricerca non strumentale’ (basic research institute), thereby according the institute an even larger degree of autonomy.
The management of the Stazione Zoologica was entrusted to the President, the Director and the Administrative Council, the latter composed of the President, the Director and six other members. An International Scientific Advisory Board was created as the consulting body for the scientific and cultural policy of the Stazione. The expertise and direct involvement of the members of this Advisory Board was crucial and instrumental in helping to re-establish the Stazione on the international research scene.
In 1984 a new government commissioner was appointed (Luigi Frunzio). Finally, in 1987, Gaetano Salvatore was named as President, his responsibilities complemented and supported by those of the General Director : Antonio Miralto (1984-1990), and Lucio Cariello (1991- ). A new period of concrete dreams and visionary projects began and the new-born organism reached its full maturity.
Because of his scientific status, his international links and management abilities, Gaetano (Nino for his friends) Salvatore was able to secure scientific and financial support from national and international scientific organizations. The Scientific Advisory Board, composed of experts from disciplines of concern to the Stazione Zoologica, was established in order to maintain the highest possible scientific standards, to strengthen the international character of the Stazione, and to program the scientific development and broaden its research activities, and to stimulate even further active cooperation with the Italian and foreign scientific communities.
The internal structural organization was remodeled, in such a way as to combine two aims that had for a long time been considered contradictory: emphasis was placed, on the one hand, on the Stazione’s in-house scientific activity, conducted by the permanent scientific staff and, on the other, on national and international co-operation, formalized through conventions, research contracts, and allocation of research space to public and private research institutes.
Particular emphasis was placed on the development of new laboratories in the new fields of biology, such as Molecular Biology and Biotechnologies, and in the promotion of and participation in national and international projects aimed at solving problems related to safeguarding the marine environment and the rational exploitation of marine resources.
The training programs, that for a long time had remained a secondary aspect of the Stazione’s activities, became a priority. Regular, specialized courses, meetings and workshops were organized and promoted, together with the specialized postdoctoral training of Italian and foreign scientific and technical staff.
The cultural activities of the Stazione were also further developed, through the organisation of cultural events, such as public lectures, concerts which took place in the Fresco room given by a specialized chamber ensemble, ‘I Musici dell’Aquarium,’ seminars devoted to philosophical debate in the biological sciences.
Anton Dohrn began his Stazione with departments for zoology, marine botany and physiology. The new structure of the research laboratories included biochemistry and molecular biology, neurobiology, biological oceanography, marine botany, cell biology, ecophysiology and benthic ecology. At the same time the technical facilities included one of the best libraries for marine biology and related fields, the Historical Archives and Scientific Collections (a Museum that houses about 3500 specimens of the Gulf of Naples, some of which are very rare, and an herbarium which serves as an archives for historically important material as well as the site for the permanent location of holotypes of new species described for the Gulf of Naples).
Research at the Stazione Zoologica had always focused on two aspects of marine organisms: their structures, functions and habits were studied to further knowledge of the inhabitants of the Mediterranean and they were also used to further the knowledge of the basic processes of life. This cultural attitude remains at the center of new research projects.
Fundamental biological research is associated with the identification of opportunities where marine biotechnology can contribute to solving the problem of sustaining marine natural resources, the research on the culture of marine invertebrates, the role of several biologically active substances from marine organisms, in particular the physiological roles of ‘hormones-peptides’ and the transglutaminase enzymes during cell development, the study of the marine environment in relation to biodiversity, focused on complex ecosystems such as the seagrass meadows, Poseidonia oceanica and Cymodocea nodosa.
Programs of an applied nature are given access to funds from numerous governmental and private funding agencies and have helped to create a new image, as the Stazione Zoologica is now seen as a focal point for the local community in the search for solutions to problems concerning the preservation of the sea.
Back to future’: Giorgio Bernardi
After the death of Gaetano Salvatore in 1997, Giorgio Bernardi, who had a long experience of contacts with the SZN and a high scientific visibility, was appointed as the new President. He took over the new developments and activities initiated by Nino Salvatore, maintaining the main objective of the Stazione: to produce scientific research to the highest standards in the fields of competence and to maintain its scientific and cultural role both in the international context and in the local community.
The creation of the new department of molecular evolution seems to realize, after a century and a quarter, Anton Dohrn’s scientific and philosophical dream of linking experimental biology to Darwinian theory. The new laboratory of molecular evolution in fact uses the best results produced by molecular biology to support Darwinian theory. The future of this prestigious institution harkens back to the vision of its founder, bringing together, in a dynamic institution, science, philosophy, culture and the constant ideal of producing something useful to mankind.
Conclusions
The unique character of the Stazione Zoologica stems from many complementary factors. First of all, there is the high level of scientific activity, aimed at fundamental biological questions and at improving living conditions and the quality of the environment. Secondly, there exists active and constant interaction and exchange with the international scientific community, the only way to ensure high-level scientific research. Thirdly a flexible organizational structure, and solid financial management make the Stazione independent from other academic and political organizations, a factor that greatly enhances the freedom to cooperate with Italian and foreign scientists and with scientific organizations. As well there are the unrivalled library facilities and the ability to use the best technical tools for biological research, from Zeiss microscopes and the Warburg apparatus, to the present-day advanced computer facilities and information retrieval. And finally one must note a cultural atmosphere that has contributed greatly to the exchange of ideas and experience, and to the creative interaction among different cultures.
These aspects have made the Stazione Zoologica an example and a leading institution in the field of biology. It has shown itself to be of optimal size in terms of space, staff, budget and facilities; larger establishments are not usually flexible enough to allow extensive programs of transformation, whereas smaller laboratories rarely achieve the critical mass which is vital to produce a significant impact on the international scientific community. Tradition and innovation are fused together in an Institute that always wants to remain at the edge of advanced biological research.
The ‘organism’ named Stazione Zoologica was able to survive in a difficult and very competitive environment, under strong selective pressure, not because it was itself strong enough to surmount all the adversities it encountered during its long history, but because it was able, at each crisis, to find new forces and new ideas brought by the community of all those who kept its project alive and found the best ways to revitalize it both culturally and materially.
Research institutes are ‘political structures’ in the large sense, as they delimit the organization of scientific enterprise, guide intellectual and institutional transformations, define problems to solve, and suggest and apply a given ‘image of science’ and ‘scientific style.’ Every scientific institute has its own specific nature, the result of its history and of the living and working experiences that have taken place within its walls. It is the result of a tradition which becomes an integral part of an institution’s history and its functioning, a specific ‘soul’ which permeates everybody who takes part in its activities, permanently or temporarily. This is particular true when the institution demonstrates unique and unrepeatable traits, when its finalities are original and specific. The Stazione Zoologica ‘Anton Dohrn’, because of its history which spans three centuries, its peculiar structure, its scientific life always at the leading edge of research and thinking, from Darwinian revolution to the molecular revolution, its particular mixing of international and national cultural characteristics, shows these traits of uniqueness and originality that make it an ‘institute with a soul’. The permanence of a tradition, in particular the ‘innovative tradition’, the ability to continuously renew in order to remain at the edge of creation; these are not obstacles but guarantees for the future.
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