Kovalevsky, Aleksandr Onufrievich

(b.Daugavpils district, Vitebsk region, Russia [now Latvian S.S.R.], 19 November 1840; d. St. Petersburg, Russia, 22 November 1901)

embryology.

Kovalevsky, the leading Russian embryologist of the late nineteenth century, was an adherent of Darwin’s evolutionary theory. His numerous embryo. logical-studies of vertebrates and invertebrates estab. lished the occurrence of gastrulation by blastular invagination in a wide range of organisms and made a major contribution to the theory of germinal layers. His father, Onufry Osipovich Kovalevsky, was a Russianized Polish landowner of modest means; his mother, Polina Petrovna, was Russian. In 1856 Kovalevsky entered an engineering school in St. Petersburg; but in 1859, against the wishes of his father, he left it and enrolled in the natural sciences division of’ the Physico-Mathematics Faculty of St. Petersburg University, where he studied histology and microscopy under L. A. Tsenkovsky and zoology under S. S. Kutorga.

In the fall of 1860 Kovalevsky went to Heidelberg, where he worked in the laboratory of Ludwig Carius, publishing two works in organic chemistry, and attended lectures in zoology by G. K. Bronn. He spent three semesters at Tübingen before returning to St. Petersburg in 1862 to lake his examinations and to prepare a thesis. He returned in August 1863 to Tübingen, where he studied microscopy and histology with F. Leidig.

In the summer of 1864 Kovalevsky traveled to Naples to begin the embryological investigations on amphioxus, tunicates (simple and complex ascidians), hotothurians, Chaetognatha, Phoronis, and Cteno. phora that launched the studies in comparative embryology which were to be almost his sole scientific concern for the next thirty-five years and which formed the basis for both his master’s thesis (on amphioxus, 1865) and his doctoral dissertation (on Phoronis, 1866). These and later studies proved that a wide variety of organisms—coelenterates, echino. derms, worms, ascidians, and amphioxi—develop from a bilaminar sac (gastrula) produced by invagina. tion. His work also showed that later developmental stages of the larvae of ascidians and amphioxi are similar (which rinding contributed to their revised classification as chordates rather than mollusks), as are the mode of origin of equivalent organs in the embryos of worms, insects, and vertebrates, and that the nerve layers of insects and vertebrates are homo. logous. Theoretically,.his work was seen as providing embryological evidence for the descent theory and as refuting the widely accepted view, implicit in Cuvier’s work, that the organs of organisms from different embranchements cannot be homologous.

Kovalevsky apparently reached Naples in 1864 with a detailed plan of research which he subsequently followed. How this plan was formulated and how his intellectual outlook was formed are unclear: the relative importance of Tsenkovsky, N. D. Nozhin, Bronn, Leidig, Pagenstecher, Karl Ernst von Baer, Darwin’s Origin of Species, and Fritz Müller’s Für Darwin is disputed in the literature. But the impor. tance of Kovalevskv’s studies was quickly recognized by Baer, who nonetheless criticized their evolutionary tone; by Haeckel, who was greatly excited and gener. alized them well beyond Kovalevsky’s conclusion into his own theory of the gastrula; and by Darwin, who saw them as providing embryological proofs for his theory of descent.

In the fall of 1866 K. F. Kessler, zoologist and rector of St. Petersburg University, appointed Kovalevsky curator of the zoological cabinet and privatdocent. He subsequently served on the faculties of Kazan University (1868-1869); the University of St. Vladimir in Kiev (1869-1873); Novorossisk University in Odessa (1873-1890), where for a time he served as prorector; and St. Petersburg University (1891-1894).

Kovalevsky is described by contemporaries as a shy man who had almost no social life, a man totally dedicated to science, a demanding and thorough teacher who much preferred research. His only non. scientific interest seemed to be his family; in 1867, the year his father died, he married Tatiana Kirillovna Semenova; they had three daughters. He also main. tained close contacts with his younger brother, Vladimir, a paleontologist, and his sister-in-law Sonya, the mathematician.

Kovalevsky was active as a scientific organizer. He used his research trips to Naples, Trieste, Messina, Villefranche, Marseilles, the Red Sea, Algeria, and Sevastopol—which were almost annual—to make collections for Russian universities. At every uni. versity where he taught, he helped to found or was active in a natural history society; and he was instru. mental in promoting Russian biological stations at Villefranche and Sevastopol and in furthering Russian participation in the Naples Station and at Messina.

During his lifetime Kovalevsky published nothing about politics; but privately he was not totally apolitical, especially in his youth, when a number of his closest friends were politically active. Both at Kiev (1873) and at Odessa (1881) he was distressed by the government’s increasing interference in faculty appointments and university affairs; and in the 1880’s Kovalevsky seriously considered leaving Russia to join A. F. Marion at Marseilles or A. Dohrn at Naples, where he hoped to find less interference and greater appreciation of his talents. By 1886 he was an honorary member of the Cambridge Philosophical Society and the Society of Naturalists of Modena; a corresponding member of the academies of sciences of Brussels and Turin; and a foreign member of the Royal Society; and he had won two prizes (1882, 1886) awarded by the French Academy of Sciences. He became a member of the Russian Academy of Sciences in 1890 and had to teach at St. Petersburg University as a professor of histology (1891-1894) in order to receive a pension.

BIBLIOGRAPHY

I. Original Works. During his lifetime Kovalevsky published over 100 monographs on vertebrate and inverte. brate embryology. His writings include Istoria razvitia Amphioxus lanceolatus ili Branchiostoma lumbricum (“The Developmental History of Amphioxus lanceolatus or Branchiostoma lumbricam”), his master’s thesis (St. Peters. burg, 1865); “Beiträge zur Anatomie und Entwickelungs. geschichte des Loxosoma neapolitanum, “in Mémoires de l’Académie impériale des sciences de St.-Pétersbourg, 7th ser., 10 , no. 2 (1866); “Anatomie des Balanoglossus delle Chiaje,” ibid., no. 3 (1866), 1-18; “Entwickelungs. geschichte der Rippenquallen,” ibid., no. 4 (1866): “Ent. wickelungsgeschichte der einfachen Ascidicn,” ibid., no. 15 (1866); “Entwickelungsgcschichte des Amphioxus lanceolatus,” ibid., 11 , no. 4 (1867); “Beilräge zur Ent. wickelungsgeschichte der Holothurien,” ibid., no. 6 (1867);“Anatomia i istoria razvitia Phoronis” (“Anatomy and Developmental History of Phoronis”), his doctoral disser. tation, preface to Zapiski Akademii nauk (St. Petersburg), 11 , no. 1 (1867); “Untersuchungen üer die Entwickelung der Coclenteraten,” in Nachrichten von der Gesellschaft der Wissenschaften zu Göttingen (1868), no. 7. 154-159; “Bei. träg zur Entwickelungsgeschichte der Tunikaten,” ibid., no. 19, 401-415; Kratkii uchebnik zoologii (“Short Text. book of Zoology”). 2nd rev. ed. (St. Petersburg, 1869);“Embryologische Studien an Würmern und Arthro. poden,” in Mémoires de l’Académie impériale des sciences de St.-Pétersbourg, 7th ser., 16 , no. 12 (1871), pp. 1-70;“Weitere Studien über die Entwickelung der einfachen Ascidien,”in Archiv für mikroskopische Anatomie, 7 (1871), 101-130: “Sitzungsberichte der zoologischen Abtheilung der III. Versammlung russischer Naturforscher in Kiew,”. in Zeitschrift für wissenschaftliche Zoologie, 22 , no. 3 (1872), 283--304; “Nabliudenia nad razvitiem Coelentera. ta” (“Observations on the Development of Coelentera-a”), in Izvestia Obshchestva liubitelei estestvoznanii, antropologii i etnografii, 10 , no. 2 (1873), 1-36; “Nabliudenia nad razvi. tiem Brachiopoda” (“Observations on the Development of Brachiopoda”), ibid., 14 (1874), 1-40; “Ueberdie Knospung der Ascidien,” in Archiv für mikroskopische Anatomie, 10 (1874), 441-470; “Ueber die Entwickelungsgeschichte der Pyrosoma,” ibid., 11 (1875), 598-635; “Weitere Studienüber die Entwickelungsgeschichte des Amphioxtis lanceola. tus, nebst einem Beitrag zur Homologie des Nervensystems dcr Würmer und Wirbelthiere,” ibid., 13 (1876). 181-208;“Documents pour l’histoire embryogénique des Alcyo. naires,” in Annales du Musée d’histoire naturelle de Mar. seilles, 1 , no. 4 (1883), 7-43, written with A. F. Marion;“Embryogéni du Chiton Polii (Philipii) avec quelques remarques sur ie développement des autres Chitons,” ibid., Zoologie, 1 , no. 5 (1883), 5-37; Étude sur l’embryologie du Dentale, “ibid., no. 7 (1883), 7-46;” Matériaux pour servir à l’’histoire de l’Anchinie, “in Journal del’anatomic et de la physiologie, 19 (1883), 1-22, written with J. Barrois;“Beiträge zur Kenntnis der nachcmbryonalcn Entwicklung der Musciden. 1 Theil.” in Zeitschrift für wissenschaftliche Zoologie, 45 (1887). 542-588; “Ein Beitrag zur Kenntnis der Excretionsorgane,” in Biologisches Zentralblatt, 9 , no. 2 (1889), 33-47; no. 3 (1889). 65-76; no. 4 (1889), 127-128; “Études expérimentales sur les glandes lympha. tiques des invertébrés (communication préliminaire),”. in Mélanges biologiques, Bulletin de l’Académie impériale des ciences de St.-Pétersbourg, 13 (1894), 437-459;“Étude des glandes lynphatiques de quelques Myriapodes,” in Archives de zoologie expérimentale et générale, 3 (1896), 591-614; “Étude biologique de l’Haementeria costata Müller,” in Mémoires de l’Académie impériale des sciences de St.-Pétersbourg, 8th ser., 11 , no. 10 (1900), 1-19;“Études anatomiques sur le genre Pseudovermis,” ibid., 12 , no. 4 (1901), 1-32;” Les Hedilidées, étude anatomique,” ibid., no. 6 (1901).

II. Secondary Literature. See Biograficheskii slovar’. professorov i prepodavatelei imp. S.-Peterburgskago uni. versiteta za istekshuiu tret’iu chetvert’ veka ego sushchestvo. vaniia, 1869-1894 (“Biographical Dictionary of Professors and Teachers of the Imperial St. Petersburg University During the Third Quarter Century of its Existence, 1869-1894”), I (St. Petersburg. 1896), 320-324, with a bibliog. raphy of works by Kovalevsky; L. I. Bliakher, Istoria embriologii v Rossii (s serediny XIX do serediny XX veka).Bespozvonochnye (“The History of Embryology in Russia [From the Mid-nineteenth to the Mid-twentieth Century].Invertebrates”; Moscow, 1959), with bibliographies of works by and on Kovalevsky; P. Buchinskii, “A. O. Kova. levskii. Ego nauchnye trudy i ego zaslugi v nauke”(“A. O. Kovalevskii. His Scientific Works and His Services to Science“), in Zapiski Novorossiiskago obshchestva estestvoispytatelei, 24 , no. 2 (1901-1902), 1-23; K. N. Da. vydov, “A. O. Kovalevskii. ego rol” v sozdanii sravnitel’noi embriologii” (“A. O. Kovalevskii and His Role in the Creation of Comparative Embryology”), in Priroda (1916), no. 4, 463-467; nos. 5-6. 579-598; and “A. O. Kovalevskii kak chelovek i kak uchenyi (Vospominaniia uchenika)”(“A. O. Kovalevskii as a Person and as a Scientist [Memoirs of a Student]”), in Trudy Instituta istorii estestvoznaniia i tekhniki. Akademii nauk SSSR, 31 , no. 6, 326-363; V. A. Dogcl, “Embriologicheskie raboty A. O. Kovalevskogo v 60-80kh godakh XIX v” (“Embryol. ogical Works by A. O. Kovalevskii From the 1860’s Through the 1880’s”), in Nauchnoe nasledstvo, nat. sci. ser., 1 (1948), 206-218; and A. O. Kovalevskii (1840-1901)(Moscow-Leningrad, 1945), with bibliographies of works by and on Kovalevsky; A. E. Gaisinovich, “A. O. Kova. levskii i ego rol” v vozniknovenii evoliutsionnoi embriolo. gii v Rossii” (“A. O. Kovalevskii and His Role in the Origin of Evolutionary Embryology in Russia”), in Uspekhi sovremennoi biologii, 36 (1953), 252-272; L. L. Gelfenbein, Russkaia embriologia vtoroi poloviny XIX veka (“Russian Embryology in the Second Half of the Nine. teenth Century”; Kharkov, 1956); V. S. Ikonnikov, ed., Biograficheskii slavor’ professorov i prepodavatelei imp. Universiteta Sv. Vladimira (1834-1884) (“Biographical Dictionary of Professors and Teachers of the University of St. Vladimir [1834-1884]”; Kiev, 1884), 264-268. with a bibliography of Kovalevsky’s writings; P. P. Ivanov,“A. O. Kovalevskii i znachenie ego embriologieheskikh rabot” (“A. O. Kovalevskii and the Significance of His Embryological Works”), in Izestia Akademii nauk SSSR, biological ser. (1940). no. 6, 819830; A. (i. Knorre,“A. O. Kovalevskii—osnovopolozhnik sravnitel’noi cm. briologii (k 100-letiiu so dnia rozhdeniia)” (“A. O. Kova. levskii—Founder of Comparative Embryology [for the 100th Anniversary of His Birth]”), in Uspekhi sovremennoi biologii, 13 , no. 2 (1940), 195-206; V. A. Kovalevskaia. Chistovich, “Aleksandr Onufrievich Kovalevskii. Vospo. minaniia docheri” (“Aleksandr Onufrievich Kovalevskii. Memoirs of a Daughter”), in Priroda (1926), nos. 7-8, 5-20; T. V. Makarova, “Aleksandr Onufrievich Kova. levskii v Peterburgskom universitete” (“Aleksandr Onu. frievich Kovalevskii at Petersburg University”), in Trudy Instituta istorii estestvoznaniia I tekhniki, Akademii nauk SSSR, 24 , 222-254; A. I. Markevich, Dvadtsatipiatiletie’a imp. Novorossiiskogo universiteta. Istoricheskaia zapiska (“Twenty-fifth Anniversary of the Imperial Novorossisk University. Historical Note”; Odessa, 1890), pp. 457-661; V. F. Mirek,” Aleksandr Onufrievich Kovalevskii (1840-1901),” in Liudi Russkoi nauki (“People of Russian Science”), II (Moscow-Leningrad, 1948), 705-715. with a bibliography of works on Kovalevsky; E. Ray-Lankastcr,“Alexander Kowalevsky,” in Nature, 66 , no. 1712 (1902), 394-395; V. Shimkevich. “A. O. Kovalevskii (nekrolog),”. in Obrazovanie (1901), no. 11, 107-1’4; A. D. Nekrasov and N. M. Arlemov, eds., A. O. Kovalevskii. Izbrannye raboty (“A. O. Kovalevskii. Selected Works”; Moscow. Leningrad, 1951), with biographical essay, pp. 536-621, commentary by the eds., and bibliographies of works by and on Kovalevsky; V. L. Omelianskii, “Razvitie estest. voznaniia v Rossii v posledniuiu chetvert’ veka” (“The Development of Science in Russia in the Last Quarter. Century”), in Istorla Rossii o XIX veke, IX, 116-142;Iu. I. Polianskii, I. I. Sokolov, and L. K. Kuvanova, eds., Pis’ ma A. O. Kovalevskogo k J. I. Mechnikovu (1866-1900) (“A. O. Kovalevskii’s Letters to I. I. Mechnikov [1866-1900]”; Moscow-Leningrad, 1955); I. I. Puzanov, “Alek. sandr Onufrievich Kovalevskii, ego zhizn’ i znachenie v mirovoi nauke” (“Aleksandr Onufrievich Kovalevskii, His Life and Significance for World Science”), in Trudy Odesskogo derzhavnogo universiteta, 145 (1955), 5-19;S. la. Shtraikh, Sem’ia Kovaleskikh (“The Kovalevskii Family”; Moscow, 1948); and “Iz perepiskl V. O. Kova. levskogo” (“From the Correspondence of V. O. Kova. lcvskii”), in Nauchnoe nasledstvo, nat. sci. ser., 1 (1948), 219-423; and V. V. Zalenskii, “A. O. Kovalevskii,” in Izvestia Akademii nauk, 15 (1901), xci-xciv; and “Spisok sochinenii Akademika A. O. Kovalevskago” (“List of the Works of A. O. Kovalevskii”), in Izvestia imperatorskoi Akademii nauk, 22 , no. 1 (1905), 1-4.

Mark B. Adams