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Robert Russell Bensley. 1867 У1956

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1867-1 956
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
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.
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
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
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
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
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-
pared tissues, found that these particles existed in the living
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.
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
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
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
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
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
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
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.
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
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
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
The histology and physiology of the gastric glands. Proc. Can. Inst., 2:
The structure of the mammalian gastric glands. Quart. Jour. Micr. 800.)
4 1 : 361-389.
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.
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:
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.
1930 The structure of the renal corpuscle. Anat. Rec., 47: 147-165 (with R. D.
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.
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.
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.
Studies on cell structure by the freezing-drying method. VI. The preparation and properties of mitochondria. Ibid., 449-455 (with N. L. Hoerr).
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.
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) .
Chemical structure of cytoplasm. Science, 96 : 389-393.
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).
Book Review. Physical Chemistry of Cells and Tissues. By Rudolph Hober.
Anat. Rec., 95: 75, 76.
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.
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