t 1867-1 956 ROBERT RUSSELL BENSLEY ROBERT RUSSELL BENSLEY 1867-1956 To all of us who received our training from Bensley, one of his sayings was impressed most deeply on our consciousness (and conscience). Whenever we criticized a paper arrogantly, or an investigator’s misguided approach to a problem, or conclusions we considered unjustified, or particularly if the question of priority came up, we were apt to get a small sermon. Bensley would say: “It doesn’t matter who is right; the important question is what is true.” ?Ve could not help but grow a little more in stature in being reminded that the search for truth is the privileged duty of a scientist, and is t o be pursued with singleness of purpose. We also gained in tolerance. Doctor Bensley died on June 11, 1956. With his passing we have lost one of the great leaders of anatomy, one who helped to bring about the twentieth century evolution of Anatomy from a science of purely descriptive morphology to one in which the emphasis is on the functional aspects of structure. His research has left its impress upon investigations all over the world in fields f a r beyond anatomy and has helped to bring anatomy to its present day eminence in biology and medicine. But his warm personality, his interest in and encouragement of young investigators and students, all have had a significance more far-reaching than we can estimate. From 1901 when he moved to the University of Chicago from Toronto, almost until the day of his death, his studies, his ideas and ideals, and the example he set played a n im‘Memoir prepared by Normand L. Hoerr, with the help of Sylvia H. Bensley, Gordon Scott and others, for the American Association of Anatomists. 3 THg ANATOMICAL RECORD, V O L 128, NO. YAY 1957 1 4 ROBERT RUSSELL BENSLEY portant part, not only in the development of the university, but in their influence on medical investigators and educators of the entire world. His outstanding characteristics were his invariable encouragement of any inquiring mind and his ability to stimulate the thinking of anyone who talked to him. These qualities made him a great teacher. Although Doctor Bensley early showed evidence of mental brilliance, it was not clear in the beginning that his contribution would be in the field of science. He was born on November 13, 1867, the third of six children, on a hilltop farm near Hamilton, Ontario. He always spoke with great warmth of his childhood days and of his family, but with a special love for his father and mother. His father, Robert Daniel Bensley, of English ancestry, was a well-to-do farmer. He served f o r a time as treasurer of the township of Barton, Ontario. His mother, Caroline Vandeleur, of Irish parentage, gave Doctor Bensley some of his love of the arts. She taught him songs in Gaelic, and undoubtedly awakened his interest in and love for languages as well as music. From his father he inherited a strong physique and mathematical ability, in addition to his joy in the outdoors. Bensley first attended a small country school south of Hamilton where he was an indifferent pupil. Just before his final examinations his teacher, who apparently understood him, threatened to punish him if he did not do better than he had been doing. As a consequence he passed all his examinations and took a prize in French. Bensley’s original plans had been to become a farmer, but his teachers somehow persuaded him to continue his education at the Collegiate Institute in Hamilton. For two years he walked every day down and up the mountain two and a half miles each way, and also did his share of the farm chores. He graduated from the Institute when he was sixteen. Although he himself never took formal music lessons, he shared his sister’s assignments in piano and thus learned to play not only the piano but also the organ and later the MEMOIR 5 violin. He developed a remarkable dexterity on both the piano and violin and kept all his life a great love of music. I n the fall of 1884 he left for Toronto to enter University College in the class of '88. He entered into collegiate life with zest and took part in the lively pranks and practical jokes which were customary at this time. He registered for all the courses in arts and sciences that were given. Obviously he did not attend all the classes, but passed the courses by examination. He retained all his life considerable fluency in French and German, and a reading knowledge of Italian and Spanish. In his third year of college at the age of twenty he suffered a severe gunshot wound in his left leg when alone on a hunting expedition near Hamilton. When he realized that a large artery had been sevcred and that he was bleeding to death, he had the presence of mind to apply a tourniquet and call f o r help. His feeble calls were eventually heard and he was carried home to the farmhouse. A noted surgeon of the day amputated the shattered leg below the knee, performing the operation on the dining-room table in Bensley 's home. Following the operation gangrene set in and another amputation high in the thigh had to be performed. Subsequent to this second operation, thrombophlebitis, septicemia, and bacterial endocarditis developed. The surgeon left a pound of opium with the family with directions to keep the patient comfortable, and left the young man to die. Bensley, however, regained consciousness and found a fierce determination to live. After about two weeks he recognized his dependence upon the opium and suspected that many of his symptoms were due to the drug. With typically firm resolution, he refused to take any more. He suffered about four days of torture after the withdrawal of the opium, but after this time his recovery was progressive and complete. His convalescence took almost a year. During this time at home he did a prodigious amount of reading and acquired his life-long interest in cellular biology. He persuaded his 6 ROBERT RUSSELL BENSLEP father to buy a microscope f o r him with the argument that he might be able to solve the problem of the wheat rust that was then damaging Canadian crops. He never did solve this problem, but he learned how to make dyes out of bark and fruit and how to stain tissues with them. He mounted his preparations with balsam collected from the blisters of a Canada balsam tree in the front yard. It was during this year that a whole new world was opened to him and the central theme of all his future investigations had its origin, the structure and function of the cell. I n the autumn of 1888 he returned to University College and at the end of a crowded academic year graduated with honors in 1889. He won the Governor General’s medal in both arts and sciences and prizes in both chemistry and languages. Characteristically he spent the prize money on four books. It was also characteristic of him that he had been able to take two courses simultaneously in the college and win honors in both. I n 1892 he received the degree of M.B. from the University of Toronto. The same year he married Cariella May, and also was appointed to the teaching staff of the department of biology. At that time his salary was insufficient to live on and he practiced medicine part time. Bensley taught as an assistant demonstrator in biology from 1892 to 1899 and then as demonstrator from 1899 to 1901. He taught botany, comparative anatomy, human gross anatomy, and histology. During this period he worked out the microchemical reaction for masked iron under the direction of A. B. McCallum. I n 1896 Bensley published his paper on the gastric glands. This classic paper is still the foundation of our knowledge of the replenishment of cells in the mucous membrane of the stomach. He often spoke of a letter which he received from Sir William Osler congratulating him on this paper and mentioned how encouraging these kind words were to him as a young scientist. During this period in addition to his regular duties he served as assistant to the provincial pathologist and through this connection obtained from a hanging MEMOIR 7 excellently fixed normal human material for histological study. He also served as assistant to the head of the Public Health Department, J. J. MacKenzie. During epidemics of typhoid, diphtheria, and scarlet fever he was kept busy collecting and diagnosing cultures. During the summers before 1900 Bensley rented a cottage at the Muskoka lakes and later bought an island there. He had a rare ability of combining work and play. During these summers at the Muskoka lakes he collected botanical and zoological specimens, fished, hunted, explored, corrected examination papers, worked on manuscripts, and enjoyed the company of his family. One summer he attended the opening of the Marine Biological Laboratory at Woods Hole, Massachusetts, as the Canadian representative. In the summer of 1898 he founded the Marine Biological Laboratory at St. Andrews, New Brunswick, for the Canadian government. In 1901 the Canadian government asked him to design, build, and equip a marine biological laboratory on Georgian Bay. That summer he sold his island in the Muskoka lake district and bought an island at Gohome Bay, built a summer cottage for his family, and supervised the building and equipping of the marine biological laboratory there. In the fa11 he resigned from the University of Toronto and moved to the University of Chicago. His younger brother, Arthur, who had been living with him since their father’s death, remained on the staff of the department of biology at the University of Toronto, later becoming chairman of the department. When Bensley joined the staff of the anatomy department at the University of Chicago, the head of the department was Llewellys Barker. TVhcn Barker left to join the faculty of Johns Hopkins in 1905, Bensley was made acting head of the department and in 1907 became director of the laboratories of anatomy. His appointment as director of the laboratories of anatomy rather than head of the department reflects the result of one of Bensley’s firm convictions. All his life he believed that 8 ROBERT RUSSELL BENSLEY a university could make the best progress if it consisted of departments with a high degree of autonomy and if the direction of each department was held by one person. During the time that he was acting head of the department, the administrators of the University of Chicago had decided that in the future chairmen of departments rather than heads with full authority would be appointed. Bensley refused a chairmanship and remained unofficially the director of the department for the next 26 years. No one ever disputed his authority or judgment because his leadership brought the department to a position of preeminence. During these 26 years he brought to the department C. Judson Herrick, George Bartelmee, Alexander Maximow, Charles H. Swift, and William Bloom. Basil Harvey and Preston Kyes had already been on the staff when Bensley came to Chicago. During this period he served as president of the American Association of Anatomists in 1918 and 1919 and as editor of the Internationale Monatschrift. During these years also, Bensley trained a number of graduate students. He seldom had more than one or two graduate students working with him at a time, so that the total number, about 30, is not impressive, but they received a measure of personal attention such as is seldom given to graduate students. He had small patience with formal regulations, and his graduate students often followed what seemed to be a haphazard course of study, but each acquired from Bensley broad interests both in anatomy and outside anatomy and a respect for scholarship of a high order. It was during the early years in Chicago that he became intrigued by the problems of mitochondria. I n the preceding decade Altmann and others had described these small particles in the cell. Fisher and Hardy had claimed that they were fixation artifacts. Micliaelis had found that they could be stained supravitally with Janus green. Bensley, who all his life was a great believer in looking at living tissues and fresh tissue wherever possible, as well as in studying pre- MEMOIR 9 pared tissues, found that these particles existed in the living tissue. His interest in mitochondria throughout his life finally led in 1934 to his separation of these particles from macerated liver cells by a method which has enabled us to analyze them directly. For many years Bensley had been dissatisfied with the indirect methods of the histologist who tried by selective staining methods to acquire some information on mitochondria in intact cells. He had been equally dissatisfied with the method of the chemist who applies analytic techniques to homogenized tissues. Bensley often said that if one wanted to know how a watch was put together, to break it into little bits with a hammer and then analyze the metals in it was only a partial approach. He said that he would be more interested in taking the watch apart bit by bit to learn as much as possible about each component part. The technique he devised was the method of partial maceration of tissues followed by differential centrifugation and isolation of the morphological constituents of a cell. This method as applied to the analysis of cellular components has won enthusiastic acceptance from the biochemists in the past fifteen years, and now the chemical study of mitochondria and submicroscopic particulates forms a large part of current literature. Beginning in 1906 he investigated the pancreas and in a series of studies established the islets of Langerhans as entities distinct from the acinar cells. He devised, with his student Lane, staining methods which distinguished the alpha and beta cells. As everyone knows, these investigations led eventually to the discovery of insulin. Before Bensley’s studies, the literature on the islets of the pancreas was confusing and contradictory. Although Laguesse had early suggested that the islets had an endocrine function, many prominent investigators determinedly maintained that these cell groups simply represented the exhausted state of the secretory acini of the pancreas and that the size and number of the islets varied with the secretory activity of the pancreas. 10 ROBERT RUSSELL BENSLEY Bensley approached the problem by developing supravital staining methods which stained all the islets in the pancreas and made possible actual counts of the islets. He also worked out a method for staining the ducts by vital dyes. These various methods made it possible to study the functional changes of the islets, to determine their relation to the ducts, and to ascertain the precise effects of duct ligation. I n his Harvey lecture, delivered in 1915, Bensley stated as follows: ‘(We can now state with assurance that the islets of Langerhans are specialized elements of the pancreas, having secretory powers differing from those of the acinous tissue, developing in embryonic life from the undifferentiated epithelium of the pancreatic anlagen and in post-fetal life from the epithelium of the ducts. The islets have a peculiar blood supply characterized by its direct arterial source, by the larger calibre of its capillary vessels, and by the close association of the latter with the epithelial cells. These experimental morphological studies give us no information as to the nature of the internal secreting function which is obviously indicated by their structure. For enlightenment on this topic we must look to further experimental work.” Since Bensley’s work had left no doubt that the islets of the pancreas were independent structures which had an independent function, Banting was stimulated to carry out extraction studies on a duct-ligated pancreas. By choosing the proper time following duct ligation when the acini were degenerated, but while the islet tissue was preserved, Banting was able to recover active insulin. In 1952 the American Diabetes Association honored Doctor Bensley with the award of the Banting Medal because of the great service he rendered in the field of diabetes. In 1912 Bensley reported with B. C. H. Harvey studies on the mechanism of hydrochloric acid secretion in the stomach. I n this paper they formulated a theory which has not been successfully refuted, although it failed to conform to a number of theoretical biochemical deductions. This work was followed in 1915 by the histological demonstration of the secretion MEMOIR 11 antecedent in the thyroid gland, together with a hypothesis of the mode of secretion which is still accepted. Bensley’s lifelong interest in the structure of the cell kept him interested in the puzzles presented by the organoid most frequently known as the Golgi apparatus. In 1910 in a paper to which little attention has been paid, Bensley established the identity of the Golgi apparatus of animal cells and the vacuolar system of plant cells, an observation which was amply confirmed by a group of French cytologists in the twenties. In 1929 Bensley wrote a paper with his student Owens on the variable results obtained in staining the Golgi apparatus with osmic acid, and one of his last papers was entitled “Golgi Apparatus : Fact Versus Artifact.” I n this paper he called attention to earlier discoveries made in his laboratory that the Golgi apparatus of pancreatic islet cells can be recognized in living tissue and that the classical structure can be demonstrated in frozen-dehydrated tissues. Bensley ’s interest in the chromidial substance, the nucleoprotein constituent of the cytoplasm which is stained by basic dyes, stimulated him to investigate the microchemical tests for phosphorus, to discover the microchemical test for masked iron mentioned above as one of his earliest studies, and to develop a microchemical test for protein using a modification of Millon’s reagent, a test stated by Lison to be the most keliable of all. His interest in the chromidial substance led to a number of studies by his students on the Nissl substance of the nerve cell, to his own further work with Gersh on the Nissl substance in 1933, and, most recently, to a paper on the same subject by Gersh and Bodian. Many of these investigations necessitated or resulted from the development of new methods of research. His elaboration of supravital staining methods and of the maceration technique have been mentioned. He set Gersh to the task of developing with modern means the technique of frozen dehydration of tissues originally suggested by Altmann. These various methods resulted in his discovery of the protein responsible for sol-gel transformations in the cytoplasm and 12 ROBERT RUSSELL BENSLEY for the organization of the cytoplasm, as well as in the separation of mitochondria mentioned before. The method of differential centrifugation made possible the localizing of lipid fractions in the cell and the discovery of submicroscopic particulates, again a field of active investigation at the present time. But to give even a brief catalog of some of Bensley's accomplishments does not give a picture of the warmth and full generosity of the man. He made the department of anatomy at the University of Chicago one of the great departments of the world by his wise selection of colleagues and his loyal support of them. The door to his laboratory was always open, and f o r everyone who walked in, whether he was a medical student, a graduate student, a colleague or a visiting scientist, he always had a friendly word. Throughout his long career, men even in fields other than anatomy were accustomed to drop in on him for advice. From his fund of knowledge and with his imaginative ability, he was always able to stimulate them to a line of thought which later proved fruitful. Those of us who had the privilege of working closely with him in our investigations soon discovered his enormous powers of concentration. On the days when he was working out the answer to a question in which he was deeply interested, we sometimes found that, although we had stopped in to talk with him about a different topic, in a very short time the conversation revolved around the question he was mulling over in his own mind. At such times we rapidly saw that it would not be courteous to bother him further and quickly left. However, at lunch or later that evening we might find him more relaxed, and then he would come back to the topic with which we had been concerned and offer most valuable help. Visitors were often amazed by Doctor Bensley 's apparent power of total recall. His ability to remember accurately information acquired fifty years before was, however, not one of photostatic memory, but a power of recall by association. The fruitfulness of his recollections of investigations of the MEMOIR 13 19th century is well known to his students. He once explained that when he read the early literature he never doubted the accuracy or honesty of the investigator’s observations, but tried then to reinterpret the findings in the light of modern knowledge. We who worked with him were often puzzled by Bensley’s detailed knowledge of current literature. We could not see how anyone who was so busy in the laboratory found time to read papers in so many different fields. For example, although he had never concerned himself directly with experimental work in cancer, he knew a great deal about the current work on cancer. His retentive memory and his vivid imagination enabled him to point the way to the diagnosis of an islet cell tumor. I n 1930 he was visiting professor at Washington University in St. Louis and, in talking to one of the residents who was puzzled by the symptoms and findings in one of his patients, suggested that the patient might have a tumor of the pancreas. The patient was operated upon and an islet cell tumor was discovered in the tail of the pancreas. Many more examples of Bensley ’s helpfulness to people outside his own laboratory could be mentioned. Bensley often made the remark that people might think of him as being mainly interested in devising new methods. I think that he used to say this to his students so that he could present his defense against this theoretical accusation. He would continue by saying that he had never been interested in methods as such but in the approach to fundamental biological problems. It is quite true that through the years, in trying to answer cytological problems, he was forced, as he used to say, to develop new methods of approach. For example, in his early years he took a great interest in staining methods and with his knowledge of chemistry thereby became an authority in dye chemistry. I n 1917, when our supply of German stains was cut off, Bensley was able to advise some of the American chemists on the manufacture of dyes. Now, thirty-five years later, we are no longer dependent on foreign sources for the dyes used in biology and medicine. 14 ROBERT RUSSELL BENSLEY Doctor Bensley’s interest in dyes, in photography, and in color theory enabled him to join his son in mutual photographic research. I n recent years he was undoubtedly of considerable help in his son’s discovery of a new method of true color photography and his development of fine grain emulsions which permit extremely great enlargements. I have mentioned the stimulation that Bensley ’s graduate students and visitors received from him. In the routine teaching of undergraduate medical courses he was completely unsystematic, and many of his colleagues considered him a poor teacher. He had no interest in surveying an organized body of knowledge for the student. His main interest in teaching was to stimulate the student into reading and thinking and learning for himself. The student who wanted to be spoon-fed found no sympathy from Bensley. Although Bensley lost his leg at the age of twenty, he never allowed this handicap to keep him from any of the interests of an active, full-blooded person. He not only continued hunting but took a great interest in fishing and all other outdoor pursuits. No one was ever a more rollicking companion in the field, and at the same time more knowledgeable of nature lore. During World War I he organized a rifle club which held its meetings under the old stadium, and he taught many of the students how to shoot. With some of his colleagues in anatomy and botany he organized a rifle team which placed second at a national meet in the middle twenties. Bensley retired twenty years ago, and with each passing year fewer men were able to meet him and receive the inspiration of his acquaintancc. His increasing feebleness in the past five years gradually reduced the frequency of his visits t o the laboratory. However, he continued working to the very end, and his last paper, an address delivered to the Histochemical Society in April, 1953, was received with great enthusiasm by the audience. As usual with Bensley, his talk was not only of the accomplishments in science in the preceding fifty years but was full of optimism for progress MEMOIR 15 in the years to come. Even though he was almost eighty-nine when he died, he died young. A biography of Doctor Bensley would be incomplete without mention of Mrs. Bensley. She was the finest companion and helpmate Bensley could have had; she was always amiable and cheerful about his every need. One example of her wonderful disposition was her unfailingly friendly hospitality to the students and guests whom Bensley without warning so often brought home at mealtime. To sit at her table was always a heart warming experience. Mrs. Bensley predeceased him by five years, and one could always sense the depth of his loss. He is survived by his daughter, Caroline May, and his son, Robert Daniel, his daughter-in-law, Sylvia Holton Bensley, and his twin grandsons, Edward and Russell. He derived the utmost pleasure and pride from the two boys, one of whom, Russell, was living with him and his daughter in recent years and helped her nurse him tenderly in his terminal illness. Reqzliescat in pace. His memory will always live in our hearts. BIBLIOGRAPHY O F ROBERT RUSSELL BEWSLEY 1896 1898 The histology and physiology of the gastric glands. Proc. Can. Inst., 2: 11-16. The structure of the mammalian gastric glands. Quart. Jour. Micr. 800.) 4 1 : 361-389. 1900 The eosophageal glands of Urodela. Biol. Bull., 8 : 87-104. 1902 The cardiac glands of mammals. Amer. Jour. Anat., 2 : 105-156. The cardiac glands of the mammalian stomach. Anat. Rec., 4: 375-390. 1903 The structure of the glands of Brunner. Decennial Publications of the University of Chicago, 10: 279-326. On the histology of the glands of Brunner. Proe. Assoc. Amer. Anat. The differentiation of the specific elements of the gastric glands of the pig. Ibid. Concerning the glands of Brunner. Anat. Anz., 83: 497-507. Stomach. Ref. Handbook Yed. Sei., 7.- 461-473. 16 ROBERT RUSSELL BENSLEY 1906 An examination of the methods for the microchemical detection of phosphorus compounds other than phosphates in the tissues of animals and plants. Biol. Bull., 10: 49-65. 1908 Professor Prenant’s theory on the nature of the granule cells of Paneth. Anat. Rec., 3: 92-95. Observations on the salivary glands of mammals. Ibid., 105-107. 1910 On the nature of the canalicular apparatus of animal cells. Biol. Bull., 19: 179-194. On the so-called Altmann granules in normal and pathological tissues. Trans. Chicago Path. Soc., 8: 78-83. 1911 Studies of the pancreas of the guinea pig. Amer. Jour. Anat., 12: 297-388. 1912 Upon the formation of hydrochlonc acid in the foveolae and on the surface of the gastric mucous membrane and the non-acid character of the contents of gland cells and lumina. Biol. Bull., 29: 225-249 (with B. C. H. Harvey). 1913 The formation of hydrochloric acid on the free surface and not in the glands of the gastric mucous membrane. Trans. Chicago Path. Soc., 19: 14-16 (with B. C. H. Harvey). 1914 The thyroid gland of the opossum. Anat. Rec., 8: 431-440. 1915 Structure and relationship of the islets of Langerhans: criteria of histological control in experiments on the pancreas. Harvey Lecture, 10: 250-289. 1916 The normal mode of secretion in the thyroid gland. Amer. J. Anat., f9: 37-54. The influence of diet and iodides on the hyperplasia of the thyroid gland of opossums in captivity. Ibid., 57-65. 1923 Irruption of injection masses into the hepatic cell. Anat. Rec., 25: 120. 1928 Functions of differentiated segments of uriniferous tubules. Amer. Jour. Anat., 41: 75-96 (with W.B. Steen). On the nature of Rouget cells of capillaries. Anat. Rec., 89: 37-55 (with B. Vimtrup). The gastric glands, in Cowdry’s Special Cytology, Paul B. Hoeber, New York City. 1928 Unders@gelser over de Rouget’ske cellers funktion og struktur. Det Kgl. Danske Videnskabernes Selskab. Biologiske Meddelelser. Bind 7, no. 4. pp. 1-26 (with B. J. Vimtrop). 1929 On osmic acid as a microchemical reagent, with special reference to the reticular apparatus of Golgi. Amer. Jour. Anat., 4 4 : 79-109 (with H.B. Owens). 1930 The structure of the renal corpuscle. Anat. Rec., 47: 147-165 (with R. D. Bensley) . MEMOIR 17 1931 The functions of the differentiated parts of the uriniferous tubule in the mammal. Amer. Jour. Anat., 47 : 241-275 (with S. H. Bensley). Pathological Report to paper by Carr, A. D., Parker, R., Grove, E., Fischer, A. O., and Larrimore, J. W. on Hyperinsulinism from B-cell adenoma of the pancreas. J. A. M. A., 96: 1362-67. Pathological Report to paper by Womack, N. A., Gnaji, W. B., Jr., and Graham, E. A. on Adenoma of the islands of Langerhans with hypoglycemia. J. A. M. A., 97: 831-38. 1931 Nethods and Problems of Medical Education. University of Chicago Division of Biological Sciences Department of Anatomy. Nineteenth. The Rockefeller Foundation, New Pork. 1933 Studies on cell structure by the freezing-drying method. I. Introduction. 11. The nature of mitochondria in the hepatic cell of Amblystoma. Anat. Rec., 67: 205-235 (with I. Gersh). Studies on cell structure by the freezingdrying method. 111. The distribution in cells of the basophil substances, in particular the Nissl substance of the nerve cell. Ibid., 369-385 (with I. Gersh). Studies on cell structure by the freezing-drying method. IV. The structure of the interkinetic and resting nuclei. Anat. Rec., 58: 1-15. 1934 Studies on cell structure by the freezingdrying method. V. The chemical basis of the organization of the cell. Anat. Rec., 60: 251-266 (with N. L. Hoerr). Studies on cell structure by the freezing-drying method. VI. The preparation and properties of mitochondria. Ibid., 449-455 (with N. L. Hoerr). 1936 Introduction to Cytological studies with the freezing-drying method. 11. Section of HCI in the stomach. Anat. Rec., 65: 417-435 (article by N. L. Hoerr). 1937 On the f a t distribution in mitochondria of the guinea pig liver. Anat. Rec., 69: 341-353. 1938 Plasmosin. The gel-and-fiber-forming constituent of the protoplasm of the hepatic cell. Anat. Rec., 72: 351-369. Handbook of histological and cytological technique. Univ. of Chicago Press (with S. H. Bensley) . 1942 Chemical structure of cytoplasm. Science, 96 : 389-393. 1943 The localization of lipoids in cytoplasm, in Essays in Biology in honor of Herbert M. Evans. pp. 95-107. Univ. of California Press. 1945 Simon Henry Gage. 1851-1944. Anat. Rec., 92: 3-5 (with B. F. Kingsbury and George L. Streeter). 1946 Book Review. Physical Chemistry of Cells and Tissues. By Rudolph Hober. Anat. Rec., 95: 75, 76. 18 ROBERT RUSSELL BENSLEY 1947 On the nature of the pigment of mitochondria and of submicroscopic particles in the hepatic cell of guinea pig. Anat. Rec., 98.-609-619. 1951 Facts versus artifacts in cytology: the Golgi apparatus. Exper: Cell Res., 3: 1-9. 1953 Symposium : The structure and biochemistry of mitochondria: Introduction and greetings. Jour. of Histochem. and Cytochem., 1 : 179-182.