Giuseppe Sterzi (1876-1919) of the University of Cagliari: a brilliant neuroanatomist and medical historian.
First of all, let me explain why we are now writing about Sterzi. Approximately two years ago I (A.R.) received a message from Prof. Colin Wendell Smith, the secretary of the FCAT, who happens to live in the antipodal (for me) island of Tasmania, asking me if I could find him the article on subcutaneous tissue written by an Italian professor named Sterzi. This article had been mentioned by Prof. DiDio during one FCAT meeting and Prof. Wendell Smith had already unsuccessfully tried to obtain it via the usual library system. Since I was aware that Sterzi had been one of my predecessors in the Anatomy Chair in Cagliari, it was easy for me to send a copy of the article to Prof. Wendell Smith, who was surprised to hear that Sterzi had been working in the antipodal (for him) island of Sardinia and so was indirectly associated with me. In order to provide him with an English summary, I had to read the 172-page article myself. I was greatly impressed by the rigorous discussion of previous reports and by the amount of new findings obtained also through a comparative anatomical/embryological approach. Prompted by this, we became interested in Sterzi's life and scientific achievements, being manifest that the studies on the subcutaneous tissue and some others such as that on the endolymphatic sac, though important and superbly carried out, had been just a parenthesis in his production devoted almost exclusively to neuroanatomy and the history of anatomy. Favaro (1921) maintains, in fact, that Sterzi made them mainly to show his critics that his production was not monothematic. Giuseppe Sterzi (he never used his second christian name: Nazzareno) was born on March 19, 1876 in Cittadella, a town in the province of Padua famous for its intact city walls. His father, who in his student years in Padua had been involved in the patriotic uprisings against the Austrian government, was an engineer working for the Italian rail system and, consequently, the Sterzi family had to change residence several times. Thus, young Giuseppe attended elementary school in Suzzara and Verona and secondary school in Carrara and Massa. In 1893 he matriculated in the University of Pisa and, following a brief period in mathematics into which he entered just to comply with the wishes of his father who wanted to make of him an engineer, he joined the medical school. From the beginning, Sterzi's proficiency in basic sciences was so outstanding that the Chairman Professor Guglielmo Romiti asked him to enter the Anatomy department as “studente interno” and, in 1895, gave him the job of “proassistente” (instructor) and second dissector. On July 12, 1899 he brilliantly defended a graduation thesis on the morphology of meninges and a few months later, because of his exceptional ability, he was appointed “Aiuto” (senior lecturer) in the glorious Anatomy Department of the University of Padua then directed by Professor Dante Bertelli. In 1906 he became Professor of Topographical Anatomy on annual contract and in 1910, having been selected by the National Search Committee, he was made full Professor and Chairman of the Anatomy Department of the University of Cagliari (Fig. 1). In Cagliari for 2 years (1911–1912) he also directed the Marine Biological Station. In 5 years of hard work, even giving up vacations (Sterzi, 1915), he completely reorganized and modernized the Anatomy Department, building inter alia a dissecting room for students and new laboratories for histology and embryology. Thanks to his skill in teaching and enthusiasm in research, he attracted many young students to the Anatomy Department. Two, among them, must be remembered for they both later became true stars of the University of Cagliari. The first: Mario Aresu (1892–1963), who in his student years worked in Sterzi's laboratory and published under his guidance two articles on neuroanatomy, became Professor of Internal Medicine and, for over 30 years, a prominent figure (Cosmacini, 1989; Dodero, 1999) in the Italian health system. The second and younger of the two: Giuseppe Brotzu (1895–1976) also attended the anatomy lab as documented by a photograph taken in 1915 (Fig. 2). In 1945 in Cagliari, where he was Professor of Hygiene, Brotzu discovered (Goodman Gilman et al., 1990; Dodero, 1999) the microorganism Cephalosporium acremonium, producer of the potent antibiotic cephalosporin (Brotzu, 1948). Giuseppe Sterzi in 1910. The laboratory of the Department of Anatomy in the Spring of 1915. In the first row, the first student on the left is Giuseppe Brotzu while the smoking one in the center is Mario Aresu. The instructor in the middle of the group is Sterzi's “Aiuto” Professor Efisio Orrù. In the early summer of 1915, Sterzi, to whom in the meantime had been unanimously offered the chair of Anatomy of the University of Messina (a position that he would never be able to claim), volunteered for the Italian Army where he served as medical officer. After the end of World War I, Sterzi, who had the rank of lieutenant colonel and was entitled to be released from the army, valiantly chose to remain in his office as director of the military hospital of Arezzo during the postwar Spanish flu epidemic. During the winter of 1919 he contracted a fever and, after several days marked by fluctuating states, died on February 17, at the age of 43. He left a loving wife and four sons, the youngest of whom followed his father into the grave just one year later. In his will he donated to the Department of Anatomy (since 1992 Department of Cytomorphology) of the University of Cagliari the entire series of his manuscripts and histological preparations, in addition to many old and precious anatomy books. Although his research activity encompassed no more than 15 years, Sterzi's scientific production consists of three large textbooks (or four if including the annotated translation of Hertwig's Organogenesis) and 30 articles, some of which are more than 200 pages long, and of which he was the sole author. As already stated, apart from the article on the endolymphatic sac and the already mentioned one on subcutaneous tissue, the themes treated by Sterzi are relevant to neuroanatomy and history of anatomy. Not being active neuroanatomists we will comment only briefly on the former topic. Since the times of his graduation Sterzi refuted the merely descriptive approach then used among Italian anatomists: In his 1910 curriculum vitae he wrote: “To describe is good, but it is only the interpretation of what it is described that makes real science” (Sterzi, 1910a). He also added that although human anatomy had been the aim of his research, comparative anatomy and embryology had been the tools that enabled him to understand human anatomy. According to the classification used in his curricula vitae (Sterzi, 1910a, 1915), the following topics may be recognized in his publications on neuroanatomical subjects. In a series of seven articles published from 1899 to 1902 that represents the progression of his thesis and that includes a long (over 260 pages) monograph on medullary meninges, Sterzi illustrates the comparative anatomy of meninges from the amphioxus to man. Contrary to previous reports the spinal meninges, both in the adult lower vertebrates and in the early developmental stages of the more advanced vertebrates, are very simple. From the mesenchyme surrounding the neural tube only a single leaflet forms the primitive meninx. In the following phylo- and ontogenetic stages, the latter divides into an internal leaflet: the secondary meninx, and into an external one: the dura mater. Finally, in higher vertebrates, even the secondary meninx divides into the arachnoid and the pia. In the same animals, Sterzi demonstrated that, while in the spinal medulla the dura keeps its identity, in the skull it fuses with the periosteum (Sterzi, 1902). He also demonstrated the continuity of all meninges with the envelopes of nerves and with the filum terminale. To describe is good, but it is only the interpretation of what it is described that makes real science.— Giuseppe Sterzi This second group of studies started with a communication read by the young Sterzi, in front of nearly all of the most important morphologists of the time, at the momentous 14th Congress of the German Anatomical Society in 1900, chaired by Golgi in Pavia. This work was continued until his Cagliari years. It includes a beautifully illustrated paper of 370 pages written in German for the journal Anatomische Hefte (Sterzi, 1904a), dealing with the comparative anatomy and embryology of these vessels from the cyclostomes (petromyzontes) to man. Sterzi showed that, both in petromyzontes and in the precocious developmental stages of higher vertebrates, the spinal medulla receives its blood from the superficial vessels. It is only in later stages that vessels penetrate the spinal medulla in various patterns. He also demonstrated that, whereas in lower vertebrates there are portions of the spinal medulla supplied by venous blood and others by the arterial one, in later phylo- and ontogenetic stages the blood supply becomes uniform for the formation of longitudinal tracts among the primitive metameric systems. In 1913, Sterzi published a study on the development of mammalian central arteries in the spinal medulla, medulla oblongata, and pons (Sterzi, 1913). He observed that these arteries maintain their symmetrical arrangement in the pons, whereas in the lower segments, owing to the formation of the median fissure, they give rise to a single median tract. According to Franceschini (1972) these findings of Sterzi have been fully confirmed by modern angiographic studies. Sterzi studied the hypophysis of petromyzontes and, at variance with previous findings, demonstrated that no infudibular gland exists in these cyclostomes (Sterzi, 1904b). He also investigated the comparative anatomy of this organ in all vertebrates and, although his terminology (pars cromophoba for what is now known as the pars intermedia, and pars cromophila for today's pars anterior) has not been further accepted, he correctly stated that all secretion goes into sinusoids. According to Sterzi, there are in this region organs which are single (epiphysis and paraphysis) and organs that are originally double (pineal and parapineal organs; Sterzi, 1905). The latter, however, in adult petromyzontes become both asymmetrical acquiring a median location below the cranial vault, which is fissured in their correspondence. They have the structure of a sort of eyes (parietal eyes). Under this heading are the articles on the development of the longitudinal cerebral fissure (Sterzi, 1912) and that on the significance of the human encephalon and telencephalon (Sterzi, 1914). Concerning the first topic, Sterzi demonstrates that, contrary to earlier reports, the longitudinal fissure is not due to pressure exerted by the falx cerebri but that it is the product of the rapid outgrowth of the cerebral hemispheres from the lateral portions of the telencephalic vesicle. He also maintains that the third ventricle consists, not only of the posterior diencephalic portion, but of a smaller anterior one of telencephalic origin. In the second article (Sterzi, 1914), Sterzi criticizes the concept put forward by Edinger (1911) that the human brain consists of a neoencephalic portion: the cerebral cortex, and of a paleoencephalic one: the remaining encephalon. He demonstrates, in fact, that even in the portion that Edinger considers paleoencephalon there are neoencephalic derivatives. Concerning whether all portions of the brain really evolve from lower craniates to man, he demonstrates that this is not the case because, in the course of phylogenesis, some parts evolve and others regress. Sterzi authored the following textbooks: Il Sistema nervoso centrale dei vertebrati (The vertebrate central nervous system) Vol. 1: Ciclostomi (Cyclostomes). Draghi: Padova, 1907, XIV–732 p., 194 Figs.; Il Sistema nervoso centrale dei vertebrati (The vertebrate central nervous system) Vol. 2: Pesci (Fishes). Libro 1: Selaci (Selachians). Parte 1: Anatomia. Draghi: Padova, 1909, XII–986 p., 386 Figs.; Il Sistema nervoso centrale dei vertebrati (The vertebrate central nervous system) Vol. 2: Pesci (Fishes). Libro 1: Selaci (Selachians). Parte 2: Embriologia. Draghi: Padova, 1912, VIII–375 p., 159 Figs.; and Anatomia del sistema nervoso centrale dell'uomo (Anatomy of the human central nervous system) 2 Vol. Draghi: Padova, 1914, 1915. Vol. I, XV–566 p., 278 figs., Vol. II, XVI–1161 p., 415 Figs. Following the publication of the first book, John Black Johnston (1908), one of Sterzi's illustrious American contemporaries, wrote in a long review published in The Anatomical Record: “Sterzi's work promises to be the most extensive work in any language dealing with the central nervous system of vertebrates.” He also stated that the book “should find a place in every anatomical library.” Judging by the titles, “the sad circumstances of his life allowed Sterzi to accomplish only the extremes of his projected monumental work” (Favaro, 1921). What he left testifies to his prodigious activity, especially when one realizes that these works were based mostly on his own comparative anatomical and embryological findings, and that even most of the illustrations were his original drawings (Fig. 3). Original drawing by Sterzi of the human hippocampal cortex. Reproduced as Figure 327 in the second volume (1915) of his textbook on human neuroanatomy. The first part of Sterzi's article on the endolymphatic sac (Sterzi, 1909a) is devoted to the anatomy and embryology of this organ in man, whereas the second part, less detailed, deals with the other craniates. In humans the sac is included into the endocranium, while in lower vertebrates it is located in between the dura and the endocranium. It is absent in selachians and teleosteans while its presence is doubtful in petromyzontes. In this work, the subcutaneous tissue is carefully studied in its general characteristics: development, chronology, gender, and nutritional conditions (Sterzi, 1910b). In addition it is described in the various regions of the human body. According to Sterzi (and his conclusions were fully accepted by Italian anatomists), the subcutaneous tissue is divided into two layers: superficial and deep, that in the head, neck, trunk, and proximal limbs are separated from each other by an intermediate layer homologous to the muscular one found in other mammals. This intermediate layer in man gives rise to the cutaneous muscles of the head and neck and to the subcutaneous and superficial parts of the external anal sphincter. In the other regions it constitutes the superficial fascia, which is lacking in distal limbs. The sad circumstances of his life allowed Sterzi to accomplish only the extremes of his projected monumental work. By searching in the archives of the University of Padua, Sterzi came to know that Fabricius (Fig. 4) had willed to the government of Venice the famous Tabulae Anatomicae that had been considered lost for 200 years. He found them, together with three other works of Fabricius, in the Marciana Library in Venice, the former state library of the Republic. Of the 300 tables mentioned by Fabricius, there were only 167, bound into eight volumes (seven devoted to human and one to comparative anatomy). Besides being the most important anatomical work of the 16th and 17th centuries, the Tabulae Pictae, which are in folio and mostly in good condition, represent (Premuda, 1993) the first coloured (hand-painted) anatomical atlas in history, while according to the same author (Premuda, 1993) the first color (printed) anatomical tables are those illustrating Aselli's (1627) work on chiliferous vessels. Some of the Tables, like the 20 on human abdominal organs, are veritable masterpieces of figurative art (Figs. 5 and 6). Although impressed by their quality, Sterzi was not able to identify the painter (Sterzi, 1909b). He could ascertain, however, that Fabricius incorporated into these drawings findings obtained by his former dissectors Giulio Casseri and Giulio Cesare Sala. According to modern experts (Semenzato, 1986), some of the Tabulae may be the work of Dario Varotari, a skilled painter and father of the famous Padovanino and friend of Fabricius. Portrait of Fabricius ab Aquapendente. From Premuda L, editor. I secoli d'oro della medicina: 700 anni di scienza medica a Padova. Modena: Panini. 1986. Tabulae Anatomicae (Pictae) by Fabricius. From Premuda L, editor. I secoli d'oro della medicina: 700 anni di scienza medica a Padova. Modena: Panini. 1986. Tabulae Anatomicae (Pictae) by Fabricius. From Premuda L, editor. I secoli d'oro della medicina: 700 anni di scienza medica a Padova. Modena: Panini. 1986. In this monograph of nearly 170 pages complete with documents found in Padua archives, Sterzi (1910c) gives a fascinating report not only of the biography and achievements of this illustrious anatomist (Fig. 7), but also of the teaching of Anatomy and of the activity of the Nationes Artistarum in the most glorious period of Padua Medical School. Of particular interest are the documents relevant to the manner in which cadavers for dissection were obtained. Apart from the two bodies supplied yearly by the government, the main source was that of stealing them from churches or even at funerals, most frequently, those of Jewish people. The life of Casserius, who started as house servant of Fabricius and then became dissector and eventually the first Professor of Surgery in Padua, is described and documented in detail. So is the long quarrel that later developed between him and his former master and mentor, Fabricius. Casserius (Giulio Casseri) at 39 years of age. From Sterzi (1910c). All of Casserius works (De Vocis Auditusque Organis, Pentaestheseion, Tabulae Anatomicae) are carefully examined page by page. Sterzi, who does not hide a great admiration toward their author, correctly acknowledges not only the discoveries but also the mistakes made by Casserius. Besides those on Fabricius and Casserius, Sterzi published in 1910 three minor works on the history of medicine. In the first one, Sterzi demonstrated that Botallus, and not Carcano (as Scarpa believed), had been the first discoverer of the foramen ovale. His conclusions were soon universally accepted. The second work is about the period of lectorship in Padua of the famous Polish physician Joseph Struthius (1510–1568) and contains an interesting report on the Institute of Lectorship in the University of Padua of that time. The third publication, I Progressi della Nevrologia, is the inaugural speech given by Sterzi on the occasion of his appointment as Professor of Anatomy at Cagliari in 1910. It is a concise and passionate history of neuroscience based, as Sterzi states, on a direct reading of classic works. Among the old theories reported by Sterzi is that of Plato, who stated that man has three souls with different functions, an idea that resonates with the modern concept of the triune brain (MacLean, 1970). He also vividly reported the progresses of neuroscience from the Renaissance to the 19th century. He stressed the importance of the anatomical and embryological comparative approach started in old times by Willis and more recently revived by Cuvier, His, and many other European scientists. As Sterzi (1910a, 1915) himself reports in his curricula vitae, his research on comparative neuroanatomy and embryology were enthusiastically acknowledged by his most illustrious contemporaries (Bardeleben, Chiarugi, Edinger, Eisler, Johnston, Krause, Nicolas, Obersteiner, Sobotta, etc.) and many of his discoveries were soon incorporated into classical anatomy textbooks. During his life Sterzi was awarded several prestigious scientific prizes, among which were the particularly important ‘Premio Fossati’ (chaired by Camillo Golgi) of the Reale Istituto Lombardo di Scienze e di Lettere, Milano in 1909 and the ‘Prix Lallemand’ of the Academie des Sciences de L'Institut de France, Paris in 1912. Despite their fortune among contemporaries, reference to Sterzi's research is found almost exclusively in classical textbooks (Ariëns Kappers, 1947; Clara, 1959) devoted to comparative neuroanatomy. This possibly is due to World War I, and to his untimely death, which obviously prevented him from completing his great project of studying the comparative neuroanatomy and embryology of all vertebrates. A second reason might also lie in the lack of translation of his textbooks that were written in Italian, a language, even then considered marginal to international science. Curiously enough, because it is linked to an author that he seems to like much less than his beloved Casserius, Sterzi's actual fame rests mostly on the discovery of Fabricius' Tabulae Pictae that is cited even in recent textbooks on the history of anatomy (Roberts and Tomlinson, 1992; Premuda, 1993). For his many outstanding achievements, both as anatomist and historian, Sterzi's work, which still awaits a critical review, clearly a and for the and for are to Prof. of the University of Cagliari, for his and with is Professor of Anatomy and of Medicine at the Medical of the University of Cagliari, where he also as of the of and of the famous of Anatomical of the and of the on Anatomical he is of the of Italian and of The European of His main research which he with his wife Professor of and are in and other and in history of anatomy. is the of the Library of the University of Cagliari at He is of the of the Italian Library His main are in Library and of
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