(b. Pretare, near Arquata del Tronto, Italy, 17 July 1864; d. Baragazza, near Castiglione dei Pepoli, Italy, 7 September 1929), histology, embryology.
Ruffini, son of Giacomo Ruffini, a magistrate, and of Vincenza Saladini, received his secondary education in Ascoli Piceno. In 1884 he began to study medicine at Bologna, graduating in 1890. While a student Ruffini worked on histology in the laboratory of comparative anatomy, and in 1888 he set up a small histology laboratory in the workhouse at Bologna. In 1890 the clinician Augusto Murri appointed him manager of the microscopy laboratory of the Bologna Medical Clinic. Four years later Ruffini had qualified himself to teach histology. Ironically, just when his aim of teaching seemed to be attained, financial needs forced him to accept a job as a country physician: in January 1895 he was director of the little hospital of Lucignano, near Arezzo. Ruffini remained there for six years: and in this hospital he equipped, at his own expense, a laboratory for histological research. Thus he was able to carry out and publish some of his most important work. While at Lucignano, Ruffini gave a free course in histology at the Institute of Anatomy of the University of Siena; and in 1901 he was appointed professor of embryology at Siena. In 1912, having won a national competition, he was appointed professor of histology and general physiology at the University of Bologna, a post he held until his death.
From 1890 to 1906 Ruffini worked on the structures of proprioceptive sensibility. In 1888, while still a student, he had begun research on nerve receptors, and he was thus the first to see the ultraterminal plates, formed at the extremity of a nerve stem that already has a motor nerve ending. In 1891, shortly after receiving his degree, he discovered in man a new form of terminal nerve expansion now known as Ruffini’s corpuscle: he published his discovery in 1894 as “Di un nuovo organo nervoso terminale e sulla presenza dei corpuscoli Golgi-Mazzoni nel connettivo sottocutaneo dei polpastrelli delle dita dell’uomo.” Ruffini described two new forms of nerve expansion in the dermal papillae that also are named for him— the papillary brushes and the interlaced ansae— thereby demonstrating that all the papillae, even those lacking Meissner’s corpuscles, were innervated. In addition, he discovered minute amyelinic networks in the subpapillary stratum of the skin.
Ruffini also was the first to observe Golgi-Mazzoni corpuscles in the subcutaneous cellular structure, and he illustrated (1900) the intermediate forms that represent links between the classical Pacinian corpuscles and the more typical forms of the Golgi-Mazzoni corpuscles. Of great importance was his contribution (1905) to the study of the thin sheath interposed in the sensory nerve fiber between the myelinic sheath and Henle’s sheath, today known as the subsidiary sheath of Ruffini. Ruffini also made a valuable contribution to knowledge of the minute structure of the organs of Golgi. In addition, he worked diligently to present the first complete, exhaustive description of the expansions in the neuromuscular spindles. His discovery in 1900 and 1902 of the ultraterminal fibrils and of fibrils that establish anastomosis between nerve corpuscles gave him the fundamental basis with which to oppose the too rigid outlines, then accepted, of the neuron doctrine.
Ruffini’s fame is still essentially linked with his illustration of the nerve expansions of proprioceptive sensibility. Nevertheless, his observations on the earliest stages of the development of the fertilized amphibian egg were far more important and original. This research, begun in 1906 and pursued throughout his life, increased knowledge of embryogeny. Ruffini’s embryological work, collected in his book Fisiogenia, may be placed between the work of Wilhelm Roux and that of Hans Spemann, who received the 1935 Nobel Prize for medicine or physiology. Ruffini published his Fisiogenia in 1925, one year after the death of Roux, the celebrated theorist of Entwicklungsmechanik; and in the year of Ruffini’s death (1929) Waldemar Schleip published a book on the same subject, the determinism of the embryo’s early development.
Ruffini was convinced that classical embryology, such as that of Wilhelm His, had become inadequate to understanding of the formation of the germ layers. On the other hand, he was convinced that Roux’s views were excessively one-sided. Roux’s imposing work was based on the action of mechanical forces exclusively (see Castaldi 1925); but Ruffini demonstrated, by means of a very precise technique, that the phenomena of gastrulation are dominated by physiological processes: cellular secretion and a peculiar form of cellular movement that he called “stichotropismus” (“a file movement of the cells”). He also first demonstrated that the cellular fields from which the germ layers develop are already well distinguished in the blastula of Amphibia. Yet Ruffini, strongly convinced of protential cellular specificity, neglected the importance of morphogenetic correlations, which were emphasized by Spemann. Although Ruffini’s Fisiogenia is thus preformist, it is nevertheless a wideranging and exhaustive treatise on the problems of general embryology.
Ruffini, who was awarded a gold medal in 1910 by the Accademia Nazionale dei Quaranta and (on the suggestion of Sherrington) a prize by the Royal Society, was a great teacher and observer; the Ruffini Institute of Histology and of General Embryology at Bologna is named in his honor.
I. Original Works. Ruffini’s writings include “Su due casi di anastomosi diretta fra i prolungamenti protoplasmatici delle cellule gangliari del cervello,” in Bolletino delle scienze mediche, 6th ser., 24 (1889); “Di una particolare reticella nervosa e di alcuni corpuscoli del Pacini che si trovano in connessione cogli organi muscolo-tendinei del gatto. Nota preventiva,” in Atti dell’ Accademia nazionale dei Lincei. Rendiconti. Classe di scienze fisiche, matematiche e naturali, 5th ser., 7, no. 1 (1892), 442–446: “Sulla terminazione nervosa nei fusi muscolari e sul loro significato fisiologico. Nota preventiva,” ibid., no. 2(1892), 31–38; “Considerazioni critiche sui recenti studi dell’apparato nervoso nei fusi neuromuscolari,” in Anatomischer Anzeiger, 9 (1894), 80–88; “Di un nuovo organo nervoso terminale e sulla presenza dei corpuscoli Golgi-Mazzoni nel connettivo sottocutaneo dei polpastrelli delle dita dell’uomo,” in Atti dell’Accademia nazionale dei Lincei. Memorie. Classe di scienze fisiche e naturali, 4th ser., 7 (1894), session of 12 Nov. 1893, also in Monitore zoologico italiano, 6 (1895), 196–203; “Sulla fine anatomia dei fusi neuromuscolari del gatto e sul loro significato fisiologico,” in Monitore zoologico italiano, 7 (1896), 49–52, also in Journal of Physiology, 23 , no. 3 (1898), 190–208; and Sulla presenza di nuove forme di terminazioni nervose nello strato papillare e subpapillare delta cute dell’uomo (Siena, 1898).
See also “Sulle fibrille nervose ultraterminali nelle piastre motrici dell’uomo,” in Rivista di patologia nervosa e mentale, 5 (1900), 433–444, with notes by the Hungarian histologist István Apáthy; “Contributo allo studio della cute umana,” in Monitore zoologico italiano, 11 (1900), 117–118, 282–289; “Le fibrille nervose ultraterminali nelle terminazioni nervose di senso e la teoria del neurone,” in Rivista di patologia nervosa e mentale, 6 (1901), 70–82; “Di una nuova guaina (Guaina sussidiaria) nel tratto terminale delle fibre nervose di senso dell’ uomo,” in Zeitschrift für wissenschaftliche Zoologie, 79 (1905), 150–170; “Le espansioni nervose periferiche alla luce dell’ analisi moderna,” in Monitore zoologico italiano, 17 (1906), 16–33, 6887: “Contributo alla conoscenza della ontogenesi degli anfibi anuri e urodeli,” in Atti dell Accademia dei fisiocritici in Siena. 4th ser., 18 (Nov. 1906). and 19 (Nov. 1907), also in Archivio italiano di anatomia e di embroilogia, 6 (1907). 129156 : and Fisiogenia. La biodinamica dello sviluppo e i fondamentali problemi morfologici dell’ embriologia generarle (Milan, 1925).
II. Secondary Literature. See L. Castaldi, “La vita e l’opera di Wilhelm Roux: 1850–1924,” in Rivista di biologia, 7 (1925), 97–104. with a complete list of Roux’s works : M. Clara, “Neue Untersuchungen zur Frage der Teilung bei den Talgdrüsen. Zugleich ein Beitrag zur Frage des ‘Stiehotropismus’ in der Formbildung,” in Zeitschrift für mikroskopisch-anatomische Forschung. 18 (1929), 487–519, which summarizes Ruffini’s views on stichotropismus and reproduces his drawings concerning gastrulation: G. Cotronei, “Angelo Ruffini,” in Rivista di biologia. 12 (1930), 198–202: E. Giacomini. “La vita e l’opera di Angelo Ruffini,” in Monitore zoologico italiano, 40 (1929), 277–292. with a complete list of Ruffini’s 85 works: G. Lamhertini, “L’opera neurologica di Angelo Ruffini,” in Essays on the History of Italian Neurology. Proceedings of the International Symposium on the History of Neurology: Varenna 30 VIII│I IX. 1961 (Milan. 1963). 195–202: G. Levi, “La Fisiogenia di Angelo Ruffini,” in Monitore zoologico italiano, 36 (1925), 176–180: and “Determinazione e specificità dei tessuti,” in Archivio italiano di anatomia e istologia patologica. 1 ( 1930), 55–84: and W. Schleip. Die Determination der Primitiventwicklung (Leipzig, 1929).