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Thomas Hunt Morgan and His Legacy

by Edward B. Lewis
1995 Nobel Laureate in Physiology or Medicine


Thomas Hunt Morgan was awarded the Nobel Prize in Physiology or Medicine in 1933. The work for which the prize was awarded was completed over a 17-year period at Columbia University, commencing in 1910 with his discovery of the white-eyed mutation in the fruit fly, Drosophila.

Morgan received his Ph. D. degree in 1890 at Johns Hopkins University. He then went to Europe and is said to have been much influenced by a stay at the Naples Marine Laboratory and contact there with A. Dohrn and H. Driesch. He learned the importance of pursuing an experimental, as opposed to descriptive, approach to studying biology and in particular embryology, which was his main interest early in his career. A useful account of Morgan's life and works has been given by G. Allen (ref. 1).

Thomas Hunt Morgan
Thomas Hunt Morgan with fly drawings.
Courtesy of the Caltech Archives.
© Copyright California Institute of Technology. All rights reserved. Commercial use or modification of this material is prohibited.

In 1928 he moved with several of his group to Pasadena, where he joined the faculty of the California Institute of Technology (or Caltech) and became the first chairman of its Biology Division. What factors were responsible for the successes that Morgan and his students achieved at Columbia University and how did these factors carry over to the Caltech era first under Morgan's, and later G. W. Beadle's leadership? It is convenient to consider three time periods:


Morgan and the Columbia Period (1910 to 1928)

Morgan attracted extremely gifted students, in particular, A. H. Sturtevant, C. B. Bridges, and H. J. Muller (Nobel Laureate, 1946). They were to discover a host of new laws of genetics, while working in the "Fly Room," in the Zoology Department at Columbia.

A. H. Sturtevant
C. B. Bridges
H. J. Muller
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Throughout their careers Morgan and these students worked at the bench. The investigator must be on top of the research if he or she is to recognize unexpected findings when they occur. Sturtevant has stated that Morgan would often comment about experiments that led to quite unexpected results: "they [the flies] will fool you every time."

Morgan attracted funding for his research from the Carnegie Institution of Washington. That organization recognized the basic research character of Morgan's work and supported research staff members in Morgan's group, such as C. B. Bridges and Morgan's artist, Edith Wallace, who was also curator of stocks. The Carnegie grants required nothing more than an annual report from the investigators. Federal support had not yet started and although universities were able to finance costs associated with teaching they were usually unable to support basic research.

C. Bridges, P. Reed, T. H. Morgan, A. H. Sturtevant,
E. M. Wallace.
Courtesy of the Caltech Archives. © Copyright California Institute of Technology. All rights reserved. Commercial use or modification of this material is prohibited.

During the Columbia period Morgan was clearly in his prime. His style of doing science must have been of paramount importance. He was not afraid to challenge existing dogma. He had become dissatisfied, even skeptical, of the formalistic treatment that genetics had taken in the period between the rediscovery of Mendelism in 1901 and 1909. He ridiculed explanations of breeding results that postulated more and more hereditary factors without any way of determining what those factors were. He wanted to know what the physical basis of such factors might be. At that time it was generally assumed that chromosomes could not be the carriers of the genetic information. He wanted a suitable animal and chose Drosophila, because of its short life cycle, ease of culturing and high fecundity. Also, large numbers of flies could be reared inexpensively -- an important factor during this period when there were very few funds available to support basic research. Morgan was very thrifty when it came to purchasing laboratory equipment and supplies -- but, according to Sturtevant, generous in providing financial help to his students. At the start of the work hand lenses were used. Only later did Bridges introduce stereoscopic microscopes. Bridges also devised a standard agar-based culture medium. Prior to that, flies were simply reared on bananas. In addition, Bridges built the basic collection of mutant stocks, mapped virtually all of the genes and later, at Caltech, drew the definitive maps of the salivary gland chromosomes. His enormous research output may in part be attributed to his being a staff member of the Carnegie Foundation with consequent freedom from teaching and other academic obligations.

Morgan's first attempts to find tractable mutations to study were quite disappointing. Fortunately, he persevered and found the white-eyed fly. This led to his discovery of sex-linked inheritance and soon with the discovery of a second sex-linked mutant, rudimentary, he discovered crossing over.


A. H. Sturtevant in the Drosophila stock room of the Kerckhoff Laboratories.
Courtesy of the Caltech Archives. © Copyright California Institute of Technology. All rights reserved. Commercial use or modification of this material is prohibited.

Sturtevant (ref. 2) has described how chromosomes finally came to be identified as the carriers of the hereditary material. In a conversation with Morgan in 1911 about the spatial relations of genes in the nucleus, Sturtevant, who was still an undergraduate, realized that the sex-linked factors might be arranged in a linear order. He writes that he went home and spent the night constructing a genetic map based on five sex-linked mutations that by then had been discovered. In 1912 Bridges and Sturtevant identified and mapped two groups of autosomal (not sex-linked) factors and a third such group was identified by Muller in 1914. The four linkage groups correlated nicely with the four pairs of chromosomes that Drosophila was known to possess. Proof that this correlation was not accidental came when Bridges used the results of irregular segregation of the sex chromosomes (or non-disjunction) to provide an elegant proof that the chromosomes are indeed the bearers of the hereditary factors or genes as they are now known. Bridges published this proof in 1916 in the first paper of volume I of the journal Genetics.

Sturtevant often commented on Morgan's remarkable intuitive powers. Thus, Sturtevant describes how after explaining some puzzling results to Morgan, Morgan replied that it sounded like an inversion. Sturtevant went on to provide critical evidence, purely from breeding results, that inversions do occur; it was only later that inversions were observed cytologically.

It seems clear that Morgan was not only a stimulating person but one who recognized good students, gave them freedom and space to work, and inspired them to make the leaps of imagination that are so important in advancing science.

Morgan and the Caltech Period (1928 to 1942)

Robert A. Millikan with cosmic ray equipment.
Courtesy of the Caltech Archives. © Copyright California Institute of Technology. All rights reserved. Commercial use or modification of this material is prohibited.

Morgan was invited by the astronomer, G. E. Hale, to chair a Biology Division at the California Institute of Technology (Caltech). Hale had conceived the idea of creating Caltech some years earlier and had already recruited R. A. Millikan (Nobel Laureate in Physics, 1923) and A. A. Noyes to head the Physics and Chemistry Divisions, respectively. According to Sturtevant, Morgan told his group at Columbia of Hale's invitation and of how it was not possible to say no to Hale. Morgan accepted and came to Caltech in 1928. He brought with him Sturtevant, who came as a full professor, Bridges, and T. Dobzhansky, who later became a full professor. In addition to Sturtevant and Dobzhansky, the genetics faculty consisted of E. G. Anderson and S. Emerson. J. Schultz, who like Bridges was a staff member of the Carnegie Institution of Washington, participated in the teaching of an advanced laboratory course in genetics.

During this second period, many geneticists visited the Biology Division for varying periods of time. Those from foreign countries included D. Catcheside, B. Ephrussi, K. Mather, and J. Monod (1965 Nobel Laureate). Visiting professors included Muller and L. J. Stadler. B. McClintock (1983 Nobel Laureate) came as a National Research Fellow in the early 1930s.

Morgan was well known outside of the scientific community and attracted interesting people. Professor Norman Horowitz, who was a graduate student in the Biology Division during this period, tells me that he remembers Morgan giving a tour of the Biology Division to the well-known author, H. G. Wells.

back:(l to r) Wildman, Beadle, Lewis, Wiersma; standing: Keighley, Sturtevant, Went, Haagen-Smit, Mitchell, Van Harreveld, Alles, Anderson; seated (back row), Borsook, Emerson; (front row) Dubnoff, Bonner, Tyler, Horowitz.
Courtesy of the Caltech Archives. © Copyright California Institute of Technology. All rights reserved. Commercial use or modification of this material is prohibited.

J. R. Goodstein (ref. 3) has described how the Rockefeller Foundation and private donors provided financial support to the Biology and other Divisions during this period. Such assistance was essential at that time, since Caltech is a private institution and received no support from the state or the federal government.


Edward B. Lewis with Drosophila.
Courtesy of the Caltech Archives. © Copyright California Institute of Technology. All rights reserved. Commercial use or modification of this material is prohibited.

In the latter half of this period, Morgan returned to his interest in marine organisms and did not follow the newer developments in genetics. Instead it was largely Sturtevant who carried on the Morgan legacy as far as genetics was concerned. Sturtevant also allowed his graduate students considerable freedom to choose their thesis projects and to consult with him on those projects or indeed on any matter. I was fortunate to have been one such student, commencing in 1939. Sturtevant's door was always open to students and faculty. I well remember Morgan coming to Sturtevant's office to discuss matters affecting the Division.

Sturtevant told us that the award of the Nobel Prize to Morgan in 1933 was an important factor in elevating the prestige and status of the Biology Division at the Institute. At the time, the only other Nobel Laureate at Caltech was Millikan. From 1942 to 1946, the Division was managed by a committee chaired by Sturtevant.


Beadle and the Caltech Period (1946 to 1961)


Beadle and Pauling with molecular model.
Courtesy of the Caltech Archives. © Copyright California Institute of Technology. All rights reserved. Commercial use or modification of this material is prohibited.

In 1946, Sturtevant and Linus Pauling (who was awarded Nobel Prizes in Chemistry, 1954, and Peace, 1962) persuaded Beadle, who was then Professor of Biology at Stanford University, to become chairman of the Biology Division. Beadle carried on the Morgan tradition of strongly supporting basic research and maintaining a stimulating intellectual atmosphere. During the early 1930s Beadle had been a National Research Fellow in the Division. He had collaborated with Sturtevant on a monumental study of inversions and together they wrote a textbook of genetics. He had collaborated also during that time with Sterling Emerson, and with E. G. Anderson. Beadle was clearly a part of the Morgan legacy.


Beadle in lab coat.
George Beadle and B. Ephrussi using microscopes.
Courtesy of the Caltech Archives. © Copyright California Institute of Technology. All rights reserved. Commercial use or modification of this material is prohibited.

Beadle received the Nobel Prize in Physiology or Medicine in 1958 for work carried out at Stanford University on the biochemical genetics of the bread mold, Neurospora. In his biographical memoir on Beadle, Horowitz (ref. 4) describes how, while postdoctoral fellows in the Biology Division, Beadle and Ephrussi decided to pursue an early discovery by Sturtevant; namely, that a diffusible substance must be involved in the synthesis of the brown eye pigment of Drosophila. Sturtevant had shown that the vermilion eye color mutation is non-autonomously expressed in flies that are mosaic for the vermilion mutation and its wild-type allele. Beadle and Ephrussi designed at Caltech a set of experiments, involving transplantation of larval imaginal eye discs, to study the vermilion-plus hormone, as they called the diffusible substance. They carried out these experiments in Paris in Ephrussi's laboratory. They were able to show that another eye color gene, cinnabar, lacks a cinnabar-plus substance and that the wild-type vermilion and cinnabar genes control sequential steps in a biochemical pathway leading to the brown eye pigment. Beadle correctly realized that the fungus Neurospora would provide better genetic material for exploring such pathways. Beadle and E. Tatum (co-winner with Beadle of the Nobel Prize) and colleagues at Stanford were then successful in dissecting the biochemical pathways that are involved in the synthesis of vitamins and many amino acids in that organism. The Neurospora findings opened a new era, now known as molecular genetics.

During Beadle's tenure as chairman, N. H. Horowitz, H. K. Mitchell, R. D. Owen, and R. S. Edgar were added to the faculty in genetics. [I had come as an instructor in 1946 before Beadle had arrived]. Horowitz and Mitchell had been associated with Beadle at Stanford and played major roles in developing the one-gene one-enzyme hypothesis that led to the award of the Nobel Prize to Beadle and Tatum.

Beadle was responsible for persuading Delbrück to return to Caltech as a full professor. Delbrück had not been offered an appointment at Caltech after his tenure in the Division in the 1930s as a post-doctoral fellow and had taken a faculty position at Vanderbilt University. Other appointments during Beadle's chairmanship that added strength in animal virology were R. Dulbecco (1975 Nobel Laureate), and M. Vogt. Howard Temin was one of Dulbecco's graduate students and later a cowinner with Dulbecco of the Nobel Prize in 1975. R. Sperry (1981 Nobel Laureate) joined the faculty as a full professor in 1954 and continued his work on split brains that he had begun at the University of Chicago.

R. Dulbecco, G. Beadle, M. Delbrück, H. Temin.
Courtesy of the Caltech Archives. © Copyright California Institute of Technology. All rights reserved. Commercial use or modification of this material is prohibited.

Basic research gradually became well supported financially by Federal Agencies commencing with the Office of Naval Research, the Atomic Energy Commission and finally by the National Institutes of Health and the National Science Foundation. Such support was essential to obtain the personnel, equipment and supplies needed by the new fields of molecular and microbial genetics which flourished and indeed flowered during Beadle's chairmanship.

During this third period there were many postdoctoral research fellows in the Biology Division, including S. Benzer (Crafoord Prize in 1993), who was a post-doctoral fellow in Delbrück's group from 1949 to 1951, and was later recruited in 1967 as full professor. J. Weigle was a visiting professor and a valuable member of the Delbrück group. There were visits by F. Jacob (Nobel Laureate, 1965) and J. Watson (Nobel Laureate, 1962). B. McClintock returned in 1946 for a short visit, working with one of the graduate students, Jessie Singleton, perfecting a method of analyzing the chromosomes of Neurospora. Interestingly, R. Feynman, Caltech professor of physics (Nobel Laureate in Physics, 1965), spent part of an academic year working with R. Edgar and other members of the Delbrück group.

George Beadle at blackboard.
Courtesy of the Caltech Archives. © Copyright California Institute of Technology. All rights reserved. Commercial use or modification of this material is prohibited.

Beadle had remarkably versatile skills. He early abandoned his research on Neurospora in order to devote full time to being chairman. He was very successful in finding donors to endow postdoctoral fellowships and new buildings. The fellowships were often used to support visits by foreign scientists who otherwise would not have had been able to come to the USA. As in the previous period, teaching loads were kept light and much teaching was conducted in the form of seminars and journal clubs. The biology faculty was by and large a harmonious group and students were allowed considerable freedom to choose their professors. As one of a number of measures of the success of this atmosphere, the Nobel Prize in Physiology or Medicine was awarded to Professors Delbrück, Dulbecco and Sperry, as already noted, and in my case as well, for work carried out in the Division under the leadership of Beadle.

George Beadle and students.
Courtesy of the Caltech Archives. © Copyright California Institute of Technology. All rights reserved. Commercial use or modification of this material is prohibited.



1. Allen, G., Thomas Hunt Morgan, pp. 1-447, Princeton University Press, Princeton, N.J. (1978).
2. Sturtevant, A. H., A History of Genetics, pp. 1-165, Harper and Rowe, New York (1965).
3. Goodstein, J. R., Millikan's School, W. W. Norton and Co., New York. pp. 1-318 (1991).
4. Horowitz, N. H. Biographical Memoirs, vol. 59, pp. 26-52, National Academy Press, Washington, D. C. (1990).

How Golgi Shared the 1906 Nobel Prize in Physiology or Medicine with Cajal

by Gunnar Grant*


Camillo Golgi was nominated for the Nobel Prize in Physiology or Medicine as early as 1901, when the first prize was awarded. After that, his name came up every year until 1906, when he was finally awarded the prize together with Santiago Ramón y Cajal.

There were four proponents for Golgi that year, namely Hertwig, professor of comparative anatomy from Berlin, Kölliker, professor of anatomy from Würzburg, and two Swedes, Gustaf Retzius, former professor of anatomy from Stockholm, and Carl Magnus Fürst, professor of anatomy from Lund. Kölliker, Retzius and Fürst proposed Golgi and Cajal. Retzius, however, proposed Cajal alone as an alternative. Cajal's nomination was also supported by, in addition, Ziehen, professor of psychiatry and neurology from Berlin, and by Emil Holmgren, from Stockholm.

Kölliker had proposed Golgi in 1901. He nominated him again in 1905 and then, as in 1906, he proposed both Golgi and Cajal. Retzius sent in proposals for Golgi all the five years from 1902. The first three times he proposed Golgi and Cajal, but in 1905 he nominated Cajal, and after him, Golgi. Finally, as mentioned, in 1906 his suggestion was for Golgi and Cajal, or Cajal alone.

The Royal Caroline Medico-Chirurgical Institute (Kongl. Carolinska Mediko-Chirurgiska Institutet; today known as Karolinska Institutet) at Kungsholmen, close to the present City Hall in Stockholm, around 1900. Located in Solna since the mid-40s.


It may be of some interest that, in 1902, Emil Holmgren, professor of histology at the Caroline Institute in Stockholm, had been one of the proponents in favor of Golgi. It was Holmgren who was commissioned by the Nobel Committee to carry out the investigation on Golgi's and Cajal's work and to write the reports for all the five years from 1902, until the time they were awarded the prize in 1906.

The comprehensive report by Holmgren in 1906, corresponding to nearly 50 type-written pages of size A4 paper, was based on a careful and extensive analysis of the merits of the two candidates, who were also weighed against each other. Holmgren's conclusion was the following (translation from Swedish by G.G.): "If the achievements by Golgi, on the one hand, and Cajal, on the other, in the research on the nervous system are considered, one can not, in justice, evade the final conclusion that Cajal is far superior to Golgi." It may be noted that this was his conclusion during the later years. In his report, Holmgren made it clear that he would have given Golgi a higher priority if it had been some years earlier. Now, however, according to Holmgren, Cajal had made such important and principally valuable discoveries, and also interpreted his findings in a correct way, as had been confirmed by others. Because of this, Holmgren was predisposed to rank Cajal before Golgi.

Favouring Cajal over Golgi, Holmgren writes (translation from Swedish by G.G.): "Cajal has not served science by singular corrections of observations by others, or by adding here and there an important observation to our stock of knowledge, but it is he who has built almost the whole framework of our structure of thinking, in which the less fortunately endowed forces have had to, and will still have to put in their contributions."

Holmgren's evaluation included Cajal's more recent contributions based on his neurofibrillar impregnation method, both for a better understanding of the interior of the nerve cell and for studies of regeneration of peripheral nerve fibers - which had also been studied by Perroncito in Pavia, a pupil of Golgi - as well as for studies of outgrowth of axons during the embryonic development, demonstrating end bulbs (growth cones). These formed part of the basis for his support for Cajal's scientific superiority.

Regarding Golgi, Holmgren discussed some of the findings which had turned out to be wrong. The most important of these were Golgi's adherence to the reticular theory, against which the neuron doctrine had been put forward and gained acceptance by most neuroscientists during this period. Further to this was Golgi's view on the dendrites, which he regarded as nutritive elements for the neurons and not involved in the conduction of impulses, as well as his view on his type II cells, which he suggested to be involved in sensory function, sending axons out from the central nervous system to the periphery, on the sensory side.

Carl Sundberg, professor of pathology at the Caroline Institute, who was also Vice President at the Institute at that time, was thereupon asked for another evaluation of the candidates, after Holmgren's conclusions had become known to members of the Nobel Committee. Contrary to Holmgren's conclusions, Sundberg, for his part, put more stress upon Golgi's valuable contributions, citing not only the development of the Golgi method but also, for instance, his findings of collaterals both in the gray matter and in the longitudinally running white columns of the spinal cord. He tried to soften the weak points in Golgi's contributions and quoted passages from evaluations done by Holmgren during the earlier years that were in support of Golgi.

Before the final decision was taken on October 25th, written opinions were expressed both by Holmgren and Sundberg, and in addition, by Bror Gadelius, professor of psychiatry at the Caroline Institute. Gadelius supported Holmgren's views.

The final voting among the professors at the Institute resulted in a majority for a Nobel Prize shared by Golgi and Cajal. Only two were against - their names were not given, but it should not be difficult to guess who they were.

Of some interest may also be Gustaf Retzius' view on the decision that was taken. This is expressed in a passage in his autobiography (1948, p. 246; translation by G.G. - italics also in the Swedish text): "... Cajal...But it is true that already at his arrival in Stockholm, I thought that he had deserved receiving a full, and undivided Nobel Prize, and asked about this by the Nobel Council of the staff of professors at the Caroline Institute, I expressed this opinion of mine decidedly."

That Retzius was asked for his opinion but did not take part in the decision, is explained by the fact that he was no longer a member of the Medical Faculty at the Caroline Institute. He had resigned from his chair in anatomy in 1890, in protest over the failure to get his candidate appointed to a professorship in ophthalmology. Paradoxically, however, his membership both in the Royal Swedish Academy of Sciences and in the Swedish Academy, meant that he took part in the election of the Laureates both in Physics and Chemistry, and in Literature.

Going back to the nominations, this was the first time that the Nobel Prize was shared between two Laureates. Cajal writes about this (from the English translation of his autobiography, 1989, p. 546): "The other half was very justly adjudicated to the illustrious professor of Pavia, Camillo Golgi, the originator of the method with which I accomplished my most striking discoveries."



Material from the Nobel Archives was kindly provided by theNobel Committee for Physiology or Medicine.

Permission to reproduce the lecture given at the Golgi and Bizzozero symposium was given by the Giornale dell'Accademia di Medicina di Torino.


Ramón y Cajal S (1989) Recollections of my life; translated by E. Horne Craigie, with the assistance of Juan Cano. Cambridge, Massachusetts, The MIT Press, 638 pp.

Retzius, G (1948) Biografiska Anteckningar och Minnen, vol. II. Uppsala, Almqvist & Wiksell AB, 285 pp.


*This article is based in part on a lecture entitled "Camillo Golgi and the Nobel Prize", given at a symposium entitled "Golgi and Bizzozeronel centenario della scoperta dell'apparato reticulare interno" at the Accademia di Medicina di Torino, Italy, November 24, 1998. Original information was achieved from the archives of the Nobel Committee for Physiology or Medicine after a formal request by the author to get access to these archives.

Life and Discoveries of Camillo Golgi

by Marina Bentivoglio


Biographical Sketch and Scientific Work

Camillo Golgi was born in July 1843 in Corteno, a village in the mountains near Brescia in northern Italy, where his father was working as a district medical officer. He studied medicine at the University of Pavia, where he attended as an 'intern student' the Institute of Psychiatry directed by Cesare Lombroso (1835-1909). Golgi also worked in the laboratory of experimental pathology directed by Giulio Bizzozero (1846-1901), a brilliant young professor of histology and pathology (among his several contributions, Bizzozero discovered the hemopoietic properties of bone marrow). Bizzozero introduced Golgi to experimental research and histological techniques, and established with him a lifelong friendship. Golgi graduated in 1865 and was, therefore, a student during the last years of the fights for the independence of Italy (Italy became a united nation in 1870).

Seated left to right: Perroncito, Kölliker, Fusari
Standing left to right: Bizzozero, Golgi (here in his late fifties).

Golgi started his scientific career in 1869, with an article in which, influenced by Lombroso's theories, he stated that mental diseases could be due to organic lesions of the neural centers. However, convinced that theories had to be supported by facts, Golgi soon abandoned psychiatry and concentrated on the experimental study of the structure of the nervous system. Histological techniques, such as fixation procedures and tissue stainings (hematoxylin or carmine) had been introduced in the middle of the 19th century. However, these procedures were inadequate and unsatisfactory for the investigation of the structure of the nervous system, due to its complexity and peculiar organization in respect to other tissues.

In 1872, due to financial problems, Golgi had to interrupt his academic commitment, and accepted the post of Chief Medical Officer at the Hospital of Chronically Ill (Pio Luogo degli lncurabili) in Abbiategrasso (close to Pavia and Milan). In the seclusion of this hospital, he transformed a little kitchen into a rudimentary laboratory, and continued his search for a new staining technique for the nervous tissue. In 1873 he published a short note ('On the structure of the brain grey matter') in the Gazzetta Medica Italiana, in which he described that he could observe the elements of the nervous tissue "studying metallic impregnations... after a long series of attempts". This was the discovery of the 'black reaction' (reazione nera), based on nervous tissue hardening in potassium bichromate and impregnation with silver nitrate. Such revolutionary staining, which is still in use nowadays and is named after him (Golgi staining or Golgi impregnation) impregnates a limited number of neurons at random (for reasons that are still mysterious), and permitted for the first time a clear visualization of a nerve cell body with all its processes in its entirety.

 Hippocampus impregnated by the Golgi stain (from an original preparation from Golgi's laboratory kept in the Institute of Pathology of the University of Pavia).

 Purkinje cells of the cerebellum. The Golgi stain reveals their extensive dendritic branches (from an original preparation from Golgi's laboratory kept in the Institute of Pathology of the University of Pavia).

In 1875 Golgi published, in an article on the olfactory bulbs, the first drawings of neural structures as visualized by the technique he had invented. In 1885, Golgi published a monograph on the fine anatomy of the central nervous organs, with beautiful illustrations of the nerve centers he had studied with his method.

Golgi's drawing of the hippocampus impregnated by his stain (from Golgi's Opera Omnia).

In the same year, Golgi returned to Pavia, where he was appointed in 1876 as Professor of Histology. In 1877 he married Lina Aletti (Bizzozero's niece). They had no children, and adopted Golgi's niece Carolina.

Camillo Golgi and his wife Lina by the sea in his late seventies.

In 1881 Golgi was appointed to the chair of General Pathology at the University of Pavia, and he also maintained his teaching in histology.

Golgi at the age of 77 in his laboratory in Pavia.

Golgi established in the Institute of General Pathology a very active laboratory, with international contacts, and was especially gifted in stimulating his students and foreign guests, including the Norwegian histologist and explorer Fridtjof Nansen (1861-1930), Nobel Laureate in Peace 1922. In Golgi's laboratory, Adelchi Negri (1876-1912) discovered the intraneuronal inclusions (the Negri bodies) which represent specific features of rabies and provided a histopathological diagnostic criterion for such infection. In Golgi's laboratory, Emilio Veratti (1872-1967), described for the first time the sarcoplasmic reticulum in skeletal muscle fibers. In 1906 Golgi shared the Nobel Prize with Santiago Ramón y Cajal (1852-1934) for their studies on the structure of the nervous system.

The Nobel diploma Golgi received in 1906.

Highly respected, Golgi was dean of the Faculty of Medicine of the University of Pavia, and rector of this university for several years. Golgi also received honours from several European universities. He took an active part in public life; he was especially concerned with public health, and became a senator in 1900. He retired in 1918 but remained as professor emeritus at the University of Pavia. Golgi died in Pavia in January 1926. His publications are collected in the Opera Omnia (published by Hoepli Editore, Milan). The first three volumes of Opera Omnia appeared in 1903 and the fourth volume was edited by Golgi's co-workers (L. Sala, E. Veratti, G. Sala) and appeared in 1929.


Scientific Debates and the Impact of Golgi's Discoveries

Golgi's discovery of the black reaction and his subsequent investigations provided a substantial contribution to the advancement of the knowledge on the structural organization of the nervous tissue. The theory that tissues are composed of individual cellular elements (the cell theory) had been enunciated in 1838-1839 by Matthias Jacob Schleiden (1804-1881) and Theodor Schwann (1810-1882), but had not been extended to the nervous tissue. However, Golgi believed that his own observations of ramified nerve fibers could support the 'reticular theory', which postulated that the nervous system was a syncytial system, consisting of nervous fibers forming an intricate network, and that the nervous impulse propagated along such diffuse network. In the meantime, the theory that the nervous system as the other tissues was composed of cells, which were christened as 'neurons' by Wilhelm Waldeyer (1836-1921) in 1891, was receiving wide support, also from studies pursued in other laboratories by means of the Golgi's new staining. Cajal was the main supporter of the 'neuron theory', which correctly interpreted the nervous system as composed of anatomically and functionally distinct cells, not in cytoplasmic continuity.

Human cerebellar cortex as drawn by Golgi (from the Opera Omnia).

Golgi was an exceptionally acute and prolific investigator, who provided a number of outstanding observations. Although he misinterpreted the overall view of the organization of the nervous system, he contributed highly to the modern knowledge of its structure. Among other findings, Golgi described the morphological features of glial cells (that are also impregnated by his staining) and of the relationships between glial cell processes and blood vessels. He also described two fundamental types of nerve cells, still named after him as neurons 'Golgi type I', extending their axons at a distance from the cell body (the 'projection neurons' of the modern nomenclature), and 'Golgi type II', with axons ramifying in the vicinity of the cell body (corresponding to the 'local circuit neurons' and 'interneurons' of the modern nomenclature).

Among his other discoveries, in 1878 Golgi described the tendinous sensory corpuscles that bear his name (the Golgi tendon organs). In the years 1886-1892, Golgi provided fundamental contributions to the study of malaria: he elucidated the cycle of the malaria agent, the Plasmodium, in red blood cells, and the temporal coincidence between the recurrent chills and fever with the release of the parasite in the blood. Golgi also studied the efficacy of the administration of quinine during the disease.

 Golgi's drawing of the tendon organ that now bears his name (from Opera Omnia).

 A typical rosette shape of the malarian parasite on the top, among red blood cells. Photograph of an original Golgi preparation preserved at the Museum for the History of the University of Pavia.

In 1897, studying the nervous system with his black reaction, Golgi noticed in neurons an intracellular structure, whose existence he officially reported in April 1898. This structure was designated by Golgi "internal reticular apparatus" and was soon named after him as Golgi apparatus (or much later as the Golgi complex and is frequently referred to nowadays only as "the Golgi"). The discovery of this cell organelle was a real breakthrough in cytology and cell biology. However, the existence of the Golgi apparatus was debated for decades (many scientists believed that it only represented a staining artefact) and was only confirmed in the mid-1950s by the use of the electron microscope. The Golgi apparatus plays a key role in the intracellular sorting, trafficking and targeting of proteins. This organelle makes Golgi the most frequently cited scientist in cell and molecular biology. The year 1998 marks the centenary of this discovery, celebrated in many scientific journals and meetings.

 Golgi's drawings of the "internal reticular apparatus" that he observed in spinal ganglia (the different drawings illustrate the variety of features Golgi observed with his metal impregnation, from Opera Omnia). This intracellular structure is universally known nowadays as "Golgi apparatus".

Rigorous, determined, highly motivated scientist and stimulating teacher, Golgi left a heritage of passionate studies that exerted a profound influence on biomedical research in the 20th century.


The Golgi Hall in the Museum for the History of the University of Pavia. In the "high altar" (to the right), exclusively devoted to Golgi, his Nobel diploma can be seen.

The Italian 'Ufficio Principale Filatelico' issued this stamp in 1994 to celebrate the Nobel Laureate Camillo Golgi. Reproduced with the permission of Mrs Maria Ciraci, The Director of Ufficio Principale Filatelico, Rome, Italy.



Thanks are due to Dr. Paolo Mazzarello for his help and advice.
Photos were kindly provided by Museo di Storia dell'Università di Pavia - Museum for the History of the University of Pavia, Director: Dr. Alberto Calligaro and from the book by Dr. Paolo Mazzarello "La struttura nascosta", Cisalpino, Istituto Editoriale Universitario - Monduzzi Editore S.p.A, 1996.

For iconographic material on Golgi's birthplace as well as on Golgi's life, the contact person is Antonio Stefanini.

The Role of Science and Technology in Future Design

by Jerome Karle
1985 Nobel Laureate in Chemistry



The role of science and technology in future design will be discussed from the perspective of someone who has lived all his life in the United States and whose scientific experience has spanned the years since the late 1930s. It is likely that the reader will find in my discussion characteristics that apply to many developed countries and developing ones. Inasmuch as scientific progress is highly dependent on financial support and, in modern times, on general societal support, it is appropriate to discuss the interaction of science and society. Using the United States as an example, some of the topics to be discussed are the views of public officials who influence the distribution of research funds, the response of funding agencies and the views of scientists. Finally, we shall look at the co-evolution of science and society and attempt to draw some conclusions concerning their related future and the implications for the future of technology.

Views of Public Officials

Public officials who are involved in setting or influencing science policy have expressed opinions that indicate that they intend to change the basis for supporting research and development. They speak in terms of a "paradigm shift" based on some new perception of the role of science in society. The word paradigm has several meanings, but in the way it is used here the words "pattern" or "model" may be good substitutes. In other words, the public officials wish to alter somewhat the pattern of funding for science. Their motivation is to orient research more toward programs that, for example, ensure a stronger economy and improvements in the environment. It is becoming increasingly apparent that those public officials who control public funds, will be reluctant to fund research programs that they consider unrelated to national needs.

An example of priority-setting by public officials was the vote in the House of Representatives against further construction of the high energy accelerator known as the superconducting super collider. This shift in spending priorities implies that nuclear physics may receive less support in the future if it continues to be viewed as less related to the new national priorities than other scientific disciplines.

Views of Funding Agencies

The effect of the intention of federal officials to shift public research funds toward research programs that serve the national priorities has already affected the nature of the funding available at the funding agencies. For example, at the National Science Foundation, a small increase in funding for the chemistry division is directed toward so-called strategic research initiatives that involve, for example, advanced materials and processing, biotechnology, environmental chemistry and high-performance computing. It is likely that this trend will continue. The Federal Coordinating Council on Science, Engineering and Technology identified the current national priority areas as high-performance computing, advanced materials, manufacturing research and education, biotechnology and global change. The expressed intention is to get more effort into those areas, but not to have them be entirely exclusive.

Views of Scientists

Many questions arose in the scientific community as a consequence of the use of words such as "new paradigm," "strategic areas", "priorities," and "national competitiveness" in statements concerning the future funding of science. The questions concerned many aspects of the support of science, such as, is the paradigm really new, who decides which areas are strategic and who sets the priorities, and are the important contributions of curiosity-driven basic research to be largely sacrificed.

The indications so far are quite clear that the government expects to shift publicly funded research activity into the areas that are deemed strategic. Is this a new paradigm or merely a shift in emphasis? Quite apparently there has been over the years heavy funding and much research in the strategic (priority) areas. There also has been in the United States, a major Industry-University cooperative research program conducted by the National Science Foundation. It celebrated its 20th year of operation in January, 1994. An account of this very successful and extensive program has been presented in the January 24, 1994 issue of Chemical and Engineering News published by the American Chemical Society. The motivation of this cooperative program is to develop and transfer industrially relevant technologies from the university into practice. There are currently more than 50 active centers involving about 1,000 faculty members, about 1,000 graduate students and 78 universities. More than 700 organizations sponsor the centers, including government agencies, national laboratories and about 500 industrial firms. A table in the article lists 55 research topics covering a broad array of technologies. It is pointed out that the success rate is very high, namely only 6% of the centers have failed. Major investments have been made by sponsor organizations, based on center technologies. There are also many other industry-university collaborations that are not part of the National Science Foundation program.

Do we really have a "new paradigm" and, if so, what is it? Performing research in the interest of national needs is not new. Cooperating with industry is not new. Setting priorities is not new. What could be new? It is indicated that what is new is that by control of public funds curiosity driven research is to be curtailed to some unspecified degree in favor of research perceived to be in the national interest. This, I believe is the source of the apprehension among scientists. The major developments in science and technology generally derive from curiosity driven research and these developments have had over time great impact on the national interest, enriching the country with whole new industries and making contributions to the health, welfare, comfort and security of society. Is curtailing curiosity driven research in the national interest?

The Impact of Curiosity Driven Basic Research

Many scientific groups have produced literature that describes, in terms of many examples, how curiosity driven research has led to important developments in the interest of society. The October, 1993 issue of Physics Today celebrated the one hundredth anniversary of the journal, Physical Review. A major part of this issue was devoted to the matter of basic research. An article by Robert K. Adair and Ernest M. Henley pointed out that "a century of fundamental physics research has appeared in the Physical Review. Such research is the seed corn of the technological harvest that sustains modern society." In an article on the laser, Nicolaas Bloembergen points out that "the first paper reporting an operating laser was rejected by Physical Review Letters in 1960. Now lasers are a huge and growing industry, but the pioneers' chief motivation was the physics." In an article on fiber optics, Alister M. Glass notes that "fundamental research in glass science, optics and quantum mechanics has matured into a technology that is now driving a communications revolution." In an article on superconductivity, Theodore H. Geballe states that "it took half a century to understand Kamerlingh Onnes' discovery, and another quarter-century to make it useful. Presumably we won't have to wait that long to make practical use of the new high-temperature superconductors." Other articles concerned nuclear magnetic resonance, semiconductors, nanostructures and medical cyclotrons, all subjects of great technological and medical importance that originated in basic physical research.

In a preface for a publication of the American Chemical Society, Science and Serendipity, the President of the ACS in 1992, Ernest L. Eliel, writes about "The Importance of Basic Research." He writes that "many people believe - having read about the life of Thomas Edison - that useful products are the result of targeted research, that is, of research specifically designed to produce a desired product. But the examples given in this booklet show that progress is often made in a different way. Like the princes of Serendip, researchers often find different, sometimes greater, riches than the ones they are seeking. For example, the tetrafluoroethylene cylinder that gave rise to Teflon was meant to be used in the preparation of new refrigerants. And the anti-AIDS drug AZT was designed as a remedy for cancer." He goes on to say that "most research stories are of a different kind, however. The investigators were interested in some natural phenomenon, sometimes evident, sometimes conjectured, sometimes predicted by theory. Thus, Rosenberg's research on the potential effects of electric fields on cell division led to the discovery of an important cancer drug; Kendall's work on the hormones of the adrenal gland led to an anti-inflammatory substance; Carothers' work on giant molecules led to the invention of Nylon; Bloch and Purcell's fundamental work in the absorption of radio frequency by atomic nuclei in a magnetic field led to MRI. Development of gene splicing by Cohen and Boyer produced, among other products, better insulin. Haagen-Smit's work on air pollutants spawned the catalytic converter. Reinitzer's discovery of liquid crystals is about to revolutionize computer and flat-panel television screens, and the discovery of the laser - initially a laboratory curiosity - is used in such diverse applications as the reattachment of a detached retina and the reading of barcodes in supermarkets. All of these discoveries are detailed in this booklet (Science and Serendipity). Ernest Eliel goes on to point, out that "the road from fundamental discovery to practical application is often quite long, ranging from about 10 years in the example of Nylon to some 80 years in the case of liquid crystals." He concludes that "if we stop doing fundamental research now, the 'well' that supplies the applications will eventually run dry. In other words, without continuing fundamental research, the opportunities for new technology are eventually going to shrink."

Some of the other topics in the brochure on Science and Serendipity, that were included to document further the importance of basic research, concerned several examples of the impact of chemistry on medicine. There are, in fact, countless such examples. The Federation of American Societies for Experimental Biology (FASEB) in their Newsletter of May, 1993 considered basic biomedical research and its benefits to society. I quote from the FASEB Public Affairs Bulletin of May, 1993. "There have been recent suggestions that tighter linkage between basic research and national goals should become a criterion for research support. Concerns also have been raised that science is being practiced for its own sake, and that it would be better for the nation if research were oriented more toward specific industrial applications." They go on to point out that "the available evidence, however, clearly indicates that the desired linkage already exists. Indeed, a majority of scientists are intimately involved in the study and treatment of common human diseases and collaborate closely with clinical scientists. Industries involved in biomedical development have been remarkably efficient in commercial application of treatment modalities based on discoveries resulting from fundamental research funded primarily by the federal government.

"A critical factor in sustaining the competitive position of biomedical-based industries is for basic research to continue to provide a stream of ideas and discoveries that can be translated into new products. It is essential to provide adequate federal support for a broad base of fundamental research, rather than shifting to a major emphasis on directed research, because the paths to success are unpredictable and subject to rapid change.

"History has repeatedly demonstrated that it is not possible to predict which efforts in fundamental research will lead to critical insights about how to prevent and treat disease; it is therefore essential to support a sufficient number of meritorious projects in basic research so that opportunities do not go unrealized. Although its primary aim is to fill the gaps in our understanding of how life processes work, basic research has borne enormous fruit in terms of its practical applications. We recognize that during a time when resources are constrained, it may be tempting to direct funding to projects that appear likely to provide early practical returns, but we emphasize that support for a wide-ranging portfolio of untargeted research has proven to be the better investment. This provides the broader base of knowledge from which all new medical applications arise. Decisions regarding what research to fund must be based on informed judgments about which projects represent the most meritorious ideas."

FASEB continues with a discussion of economic benefits and a number of examples of basic research-driven medical breakthroughs. "Society reaps substantial benefit from basic research. Technologies derived from basic research have saved millions of lives and billions of dollars in health care costs. According to an estimate by the National Institutes of Health on the economic benefits of 26 recent advances in the diagnosis and treatment of disease, some $6 billion in medical costs are saved annually by those innovations alone. The significance of these basic research-derived developments, however, transcends the lowering of medical costs: the lives of children as well as adults are saved, and our citizens are spared prolonged illness or permanent disability. Fuller, more productive lives impact positively on the nation's economic and social progress."

FASEB continues with thirteen examples of contributions by basic research to the diagnosis and treatment of numerous diseases, most of them very serious. Also noted in this Public Affairs Bulletin is that "our ability to know in advance all that is relevant is very poor" (Robert Frosch) and that, in suggesting new ideas for the management of funding for science, never considered were "the serious consequences of harming the system."


Up to this point, we have been concerned with basic science and its support by government funds in a modern society. Although there is also some support by private institutions established for that purpose and also some industrial investment in generally product-oriented basic research, the greatest amount of support by far comes from public funds. One of the ways that the public is repaid for their support is through the technology that fundamental research generates. I suspect that the economic return from technology alone more than compensates for the monies expended for the entire basic research effort. I have no estimate, however, of whether my suspicion is true or not. It should be noted that the public gains much more than the economic value of technology. It gains culture, comfort, convenience, security, recreation, health and the extension of life. What monetary value can be put on the triumphs of health over debilitating or fatal disease? The monetary value has to be higher than the purely economic savings that were noted above in the 26 examples referred to in the FASEB Bulletin.

The word "technology" means industrial science and is usually associated with major activities such as manufacturing, transportation and communication. Technology has been, in fact, closely associated with the evolution of man starting with tools, clothing, fire, shelter and various other basic survival items. The co-evolution persists and, since basic science is now very much a part of developing technologies, the term co-evolution of science and society which is used at times very much implies the co-evolution of both basic science and industrial science with society. Advances in technology are generally accompanied by social changes as a consequence of changing economies and ways of carrying out life's various activities. An important question arises concerning how basic scientific discoveries eventually lead to new technologies and what that may mean to the rational support of basic research and the future of science and technology in the developed and developing world.

There are great uncertainties in the process that starts with basic research and ends with an economically successful technology. The successful discovery of a new development in research that appears to have technological significance does not ensure the economic success of technologies that may be based on it.

Nathan Rosenberg of Stanford University, in a speech, "Uncertainty and Technological Change", before the National Academy of Sciences (April, 1994), pointed out that there are great uncertainties regarding economic success even in research that is generally directed toward a specific technological goal. He notes that uncertainties derive from many sources, for example, failure to appreciate the extent to which a market may expand from future improvement of the technology, the fact that technologies arise with characteristics that are not immediately appreciated, and failure to comprehend the significance of improvements in complementary inventions, that is inventions that enhance the potential of the original technology. Rosenberg also points out that many new technological regimes take many years before they replace an established technology and that technological revolutions are never completed overnight. They require a long gestation period. Initially it is very difficult to conceptualize the nature of entirely new systems that develop by evolving over time. Rosenberg goes on to note that major or "breakthrough" innovations induce other innovations and their "ultimate impact depends on identifying certain specific categories of human needs and catering to them in novel or more cost effective ways. New technologies need to pass an economic test, not just a technological one."

What does this mean with regard to government managed research? I quote from Rosenberg's speech.

"I become distinctly nervous when I hear it urged upon the research community that it should unfurl the flag of 'relevance' to social and economic needs. The burden of much of what I said is that we frequently simply do not know what new findings may turn out to be relevant, or to what particular realm of human activity that relevance may eventually apply. Indeed, I have been staking the broad claim that a pervasive uncertainty characterizes, not just basic research, where it is generally acknowledged, but the realm of product design and new product development as well - i.e., the D of R&D. Consequently, early precommitment to any specific, large-scale technology project, as opposed to a more limited, sequential decision-making approach, is likely to be hazardous - i.e., unnecessarily costly. Evidence for this assertion abounds in such fields as weapons procurement, the space program, research on the development of an artificial heart, and synthetic fuels.

"The pervasiveness of uncertainty suggests that the government should ordinarily resist the temptation to play the role of a champion of any one technological alternative, such as nuclear power, or any narrowly concentrated focus of research support, such as the War on Cancer. Rather, it would seem to make a great deal of sense to manage a deliberately diversified research portfolio, a portfolio that will illuminate a range of alternatives in the event of a reordering of social or economic priorities. My criticism of the federal government's postwar energy policy is not that it made a major commitment to nuclear power that subsequently turned out to be problem-ridden. Rather, the criticism is aimed at the single-mindedness of the focus on nuclear power that led to a comparative neglect of many other alternatives, including not only alternative energy sources but improvements in the efficiency of energy utilization."

To these words, I add those (noted by FASEB) of Bruce Ferguson, Executive Vice President of Orbital Sciences Corporation, a space technology firm. Ferguson said, "The federal government should focus its research and development spending on those areas for which the benefits are diffuse and likely to be realized over many years, rather than areas for which benefits are concentrated on particular products or firms over a few years. These areas are not well covered by corporate investment, yet are vital to the long-term economic strength of the country."

Some reactions to "strategic" research are recounted in an article in Nature of February 10, 1994 (Vol. 367, pp. 495-496) from which I quote some passages. The concept of strategic research "is not an unfamiliar cry, witness last year's debate in Britain about harnessing of research to 'wealth creation.' Nor, of course, is the objective in any way disreputable; what scientist would not be cheered to know that his or her research won practical benefits for the wider world as well as a modicum of understanding? The difficulties are those of telling in advance which particular pieces of research will lead to 'new technologies' and then to 'jobs'.

"The recent past is littered with examples of adventurous goal-directed programmes of research and development which have failed for intrinsic reasons or which, alternatively, have been technically successful, but unusable for economic or other reasons."

The article goes on to say that the affection for strategic research in the United States may prove short-lived. "In Britain, much the same seems to be happening. Having pinned its reorganization of research on the doctrine of science for wealth-creation, the government appears now to be more conscious of the problems it has undertaken to solve. Indeed, the prime minister, John Major, seemed to be suggesting in a speech last week that the British part of the research enterprise deserves respect of the kind accorded to other social institutions at the heart of his 'back to basics' rhetoric. After more than a decade of needless damage-doing, that would be only prudent."

As a final remark, the article ends with the statement: "On the grander questions, on both sides of the Atlantic, it seems likely that the first flush of enthusiasm for turning research into prosperity will be abated by the reality of the difficulties of doing so. When governments discover in the course of seeking radical reorganization that the best they can do with their parts of the research enterprise is to cherish them, the lessons are likely to be remembered. If the outcome in the research community is a more vivid awareness of how much the world at large looks to research for its improvement, so much the better."

The Future of Science, Technology and Society

In discussing the future of science (including industrial science) and society, it is valuable to recount some of the important points that emerged from the previous discussion.

1. As a consequence of recognizing the economic benefits that derive from the development of novel, successful technologies, governments have been attempting to direct research, supported with public funds, toward subjects that are perceived as national priorities. This contrasts with broad-based "curiosity" oriented basic research.

2. The views of scientists, a distinguished economist, some industrial leaders and an editorial comment in a distinguished science journal provide very strong indications that governmental management of goal-oriented research is replete with uncertainties and pitfalls and, although well-motivated, may cause serious damage to the scientific culture. This, of course, would defeat the original purpose, since the co-evolution of science and society is a very-well documented and irrefutable phenomenon.

3. Strong arguments are presented in this article by individuals and groups that support the current system of governmental funding of a very broad range of scientific efforts as probably being as close to optimal with regard to national priorities as is possible. No one can predict with any certainty what the most successful inventions and technologies will be in the future. The economic return on federally supported funding was the subject of a report by the Council of Economic Advisors to President Clinton. This report was released in November 1995. It documents high returns to the economy and the importance of governmental involvement. 1

4. By any measure, basic scientific research has made monumental contributions to technology and national priorities. The bond between basic research and the development of both novel and current technologies has been and is well in place.

There is no question that science and society will continue to co-evolve. The nature of this evolution will certainly be affected by the extent to which governments set funding priorities. Societies whose governments recognize the dependence of the development of successful novel technologies on broadly supported basic research are more likely to be healthier and economically prosperous in the future than those that do not. Because of the unpredictability of the details of the new science and technology that will evolve, the details of social evolution are also unpredictable.

1 The CEA Report on Economic Returns from R&D is available on the World Wide Web at