and allied subjects. At Zurich he listened to the stimulating lectures of Oken on zoology; in 1839, in Bonn, he attended the clinic of Nasse, who gave his lectures in Latin. The future anatomist tells how at the clinic at Bonn he was unable to find the vein in the arm of a very fat lady patient who was ordered to be bled. The turning point in his career came in Berlin in 1839, where he fell under the spell of Johannes Müller and Jakob Henle, whose influence on him was powerful. Under Müller he studied comparative anatomy and pathological anatomy, and under Henle normal histology. Henle instilled into him the epoch-marking doctrines of Schwann and directed his attention to the microscopical structure of the body. There also he got much encouragement from Ehrnberg, Meyen, and Robert Remak. He took a private course under Remak, who gave lectures and demonstrations at his own house on "Development of the chick." This led in future years to a study of development in mammals and many other animals and to the publication in 1861 of his Entwicklungsgeschichte des Menschen und der höheren Thiere.

Kölliker tells how, at his Staats-examen, he knew all about the finest branches of the cranial nerves, the structure of the ear, brain, and eye, yet he could not answer a simple question on the portal vein. In the session of 1841-42 he became assistant to Henle, who at that time was professor of anatomy at Zurich. In 1842 began the first of a series of journeys to other lands, which were always utilized for scientific purposes. The first was undertaken with Nägeli to Naples, where he devoted his time to comparative anatomy of aquatic forms, a subject which began to be studied in the thirties and forties by Stannius, W. Peters, J. Müller, Milne-Edwards, and Quatrefages. While there he made eight researches in all, on amphioxus, cephalopoda, on the hectocotylus of Argonauta, and kindred subjects. After Henle left Zurich in 1844 to go to Heidelberg the chair in Zurich was divided, and Kölliker, at the age of 27, became professor of physiology and comparative anatomy, with an income of 1,200 francs. In 1847, when 30 years of age, he was called to Würzburg, largely through the influence of Henle and the then rector, Professor Rinecker. Originally at Heidelberg he taught physiology, but Kölliker made it a condition that as soon as the chair of anatomy became vacant he was to have it, for he desired to devote himself to microscopical anatomy. During the Zurich period he published papers on the Pacinian corpuscles, the tissues of the tadpole, independence of the sympathetic nervous system, development of C'ephalopoda, on blood, spermatozoa, and the structure of smooth muscle. By the use of nitric acid he showed that smooth muscle is made up of fusiform cells. He also found that cellulose existed in the skin of tunicates.

At Würzburg he taught physiology, and added to this microscopic anatomy and development, so that in 1849, when the chair of anatomy became vacant, he taught all these subjects and was the head of two institutes. Kölliker, in his first decennium in Würzburg, was particularly fortunate in his associates and assistants, for he had as friends the gifted Heinrich Müller, Carl Gegenbaur, and Franz Leydig. Professor Rinecker had a microscopical institute, with Franz Leydig as an assistant, but Kölliker gave the first microscopical course in 1848. On the death of Heinrich Müller, in 1864, physiology was separated from anatomy, and Prof. A. von Bezold was called to fill the chair of physiology.

In 1849 Kölliker had as prosecutor Gottfried von Siebold, and in the same year these two founded and edited the Zeitschrift für wissenschaftliche Zoologie. Many papers on zoological subjects were contributed by him to this journal. Among his later assistants were Eberth, Forel, C. Hasse, M. Flesch, R. Fick, Grenacher, Eimer, M. v. Lenhossék, M. Heidenhain, and Ph. Stohr, who succeeded him on his retirement from active duty as an anatomist in 1897, when he had completed his eightieth year and his fiftieth as an active professor of anatomy. He still retained what he called his second institute, namely, that for comparative anatomy, microscopy, and embryology.

Kölliker, on arriving in Würzburg, found the want of a scientific society, and to him was largely due the foundation, in 1849, of the well-known Die physikalisch-medicinische Gesellschaft in Würzburg.

As showing the width of his training Kölliker lectured on human anatomy, physiology, comparative anatomy, gave a microscopical course on normal histology, and lectures on topographical histology and comparative histology, and also courses on embryology-human and comparative. He also gave short courses on comparative anatomy and physiology and on topographical anatomy.

Kölliker, of course, was the recipient of many honors. In 1897 there was conferred on him the title "Excellenz" by Prince Luitpold of Bavaria. Only once, however, was he rector of his university, and he did not take any very active part in the inner academic life.

In early youth Kölliker was a great gymnast and indulged largely in manly sports. He was a keen sportsman, especially as a hunter. He also was a great climber, and he records that in 1837 there was not an inn in Zermatt. Sic tempora mutantur.

He also traveled much and was a splendid linguist. English he spoke with great fluency, and he was a great admirer of English life and English ways. French and Italian he knew well, and he published papers in all three languages. In 1840 he visited Heligoland and worked up the fauna of the surrounding sea, for most of his

visits had a scientific object. Naples and Sicily were visited in 1842. In 1845 he first visited London, and on his way stopped at Louvain to make the acquaintance of Schwann and Van Beneden. In London he made the acquaintance of Sharpey, who was then professor in University College. A close friendship sprang up between them, and Kölliker had the highest regard for Sharpey, who was greatly interested in the young histologist, who demonstrated to Sharpey the termination of nerves in Pacinian corpuscles. He also became the friend of Owen, Bowman, Todd, Kiernan, Wharton Jones, and Ed. Forbes. Spain was visited in 1849, Holland, England, and Scotland in 1850. To London he was accompanied by Czermak. In Edinburgh he was the guest of Goodsir and Simpson.

In his letters he gives a charming account of the Edinburgh professors of those days. He says he knows only three anatomists and physiologists in all England who do not practice medicine for a profession, namely, Owen, Sharpey, and Grant. He placed Sharpey and Bowman at the head of English microscopists. When he and Czermak were the guests of Goodsir in Edinburgh, Goodsir gave him Tom Jones to read in bed. When he was tired he got up and blew out the gas, but fortunately he did not fall asleep just at once. The smell of the gas aroused him, else the career of the young histologist might probably have been short. He was present at the meeting of the British Association in Glasgow in 1855 as the guest of Allen Thomson, and on that occasion read several papers. He made another visit to Scotland in 1857 and again in 1861. He delivered the Croonian lecture in 1862. He visited Manchester to see the histological work of the late Professor Williamson on fossil plants. In 1887 he visited Pavia to study under Golgi his method of preparing sections of the nervous system, though he was then already 70 years of age. He published about twenty papers on the results he obtained on the nervous system by the Golgi and Cajal methods.

To give a detailed account of Kölliker's work would be to write a treatise on comparative histology. His Gewebelehre was published from 1850 to 1854. His Handbuch der Gewebelehre was first published in 1852 and reached its sixth edition in 1893-1899. The Manual of Human Histology was translated by Busk and Huxley for the Sydenham Society in 1853-54. Even as late as 1851 Virchow held to Schwann's doctrine of free cell formation." Before this Kölliker had rejected this theory. His studies on the eggs of cephalopods had shown him the untenable nature of Schwann's views. He attached great importance to the study of protozoa and the simplest animals. Kölliker published but little on ordinary naked-eye anatomy. His work lay chiefly in microscopical studies. He was an excellent lecturer, good draftsman, and a most methodical teacher. For eighteen

years he taught physiology, and admits that the physical and chemical side of this subject were left somewhat in the background. His microscopical course was largely attended and was given in the evening; each meeting lasted two hours. He regarded the incisures of Lantermann, the funnels and spirals of Golgi, and the EwaldKühne networks in nerve fibers as artificial and produced by the action of reagents. He supported Ranvier's view of the outgrowths of nerve fibers from a nerve cell.

That he did not do more work in naked-eye anatomy, and, in fact, produced only one large work on this subject, On the Position of the Internal Female Sexual Organs (1882), he explains by the fact that his scientific activity fell at a time when microscopical anatomy and embryology were in their infancy. Comparative anatomy he was fond of, partly because it stood in such intimate relation to the Darwinian theory, a theory which he subjected to keen criticism.

Kölliker made many important observations in physiology. He studied the action of poisons such as curare, strychnine, veratrin, upas antiar, and conium. He was specially interested in curare, for in 1856 he showed that there were poisons that, although they paralyzed intramuscular nerves, left the excitability of the muscle tissue itself intact. He published his researches on curare before those of Bernard were published. He also extended the study of poisons to other muscular tissues, such as the heart. The emission of light by Lampyris and Noctiluca was carefully studied. He also made some experiments on the electrical condition of the heart and on the secondary contraction resulting when a nerve of a nerve-muscle preparation is laid on a beating heart.

On embryology he published his great work in 1861, of which a second edition appeared in 1879. His Grundriss, on the same subject, reached a second edition in 1884.

Many papers were published on bone; the most elaborate and the best illustrated is " Die normale Resorption des Knochengewebes und ihre Bedeutung für die Entstehung der typischen Knochen formen," 1873. In 1872 he coined the word " osteoclast "in lieu of Robin's word "myeloplaque," and linked up their function with the production of Howship's lacunæ.

To the subject of Darwinism he contributed several papers, and he gives in his Erinnerungen a broad and excellent account of the doctrine of descent as it affects the cellular and other elements of the body.

Classifying his published papers, we find that on histology he published 108, including in this number his various text-books and their editions; on anatomy, 2; physiology, 16; embryology, 52; Darwinism, 5; comparative anatomy and zoology, 19, including his

Icones Histologicæ and 5 papers on unclassified subjects. This gives a measure of the variety of his work, but not of its importance. He continued to produce good scientific work until a very short time before his death.

When the writer first made the acquaintance of Kölliker his hair was already white-not so much with age, for it appears that his hair changed its color rapidly. This gave to his suave and noble face a pleasant and reverend appearance, which, coupled with a charming manner, made Kölliker an attractive personality, while his great knowledge, keen interest in all that was new, and his vast experience made him a veritable Gamaliel at whose feet it was a pleasure to sit and easy to gain inspiration and profit.

Dr. J. Dulberg (Manchester) writes:

When I was at Würzburg some sixteen years ago or so, Kölliker was still, in spite of his advanced age, one of the most indefatigable workers in the medical school, and it was a real pleasure to listen to him when either lecturing or demonstrating. It was touching to see the solicitude with which he used to arrange his microscopical specimens with his own hand and the pride he took in them. On one occasion, particularly, I remember he kept me for over a quarter of an hour discussing the merits of a section of the spinal cord which he told me was the finest he had ever seen. He examined me in anatomy for the " doctor examen," but instead of asking me questions he spent nearly the whole of the time allotted to me in giving me a miniature lecture on some anatomical detail. Even in his ordinary conversation he would not forget that he was an anatomist. When I went to say good-by to him before leaving Würzburg he presented me with his photograph, which stands before me as I write, and in doing so pointed out to me that the photographer had exaggerated the activity of his corrugator supercilii. As a teacher Kölliker was kind, patient, and helpful, far more than the generality of German professors, and to foreigners he was particularly attentive. What an army of doctors all over the world there must be to mourn the loss of one who has been a teacher for well-nigh three generations!