George Hoyt Whipple

In 1934 George Hoyt Whipple, George Richards Minot (1885-1950) and William Parry Murphy (1892-1987) received the Nobel Prize in Physiology or Medicine "for their discoveries concerning liver therapy in cases of anaemia".

George Hoyt Whipple was born in Ashland, New Hampshire. His grandfather, Solomon Mason Whipple (1820-1884), had been a practitioner and an active proponent of better medical education. Also his father, Ashley Cooper Whipple (1852-1880), was educated as a physician but died of pneumonia at the early age of 28 years, when George was only two years old.. His wife, Frances Anna Hoyt Whipple (1857-1904), was then pregnant and in July 1880 gave birth to a daughter who was named Ashley after her father.

The father’s premature death left the family in a difficult economical situation. The two children were brought up by their mother and their maternal grandmother Frances Moody Hoyt. Thanks to the efforts of the two women and their own hard work, George and Ashley were able to continue their education and preparation for college at Phillips Andover Academy. After graduating from Phillips Academy, Whipple entered Yale College, where he took his A.B. degree in 1900. As an undergraduate there, Whipple distinguished himself not only as an outstanding student of the sciences, but also as a prize-winning gymnast and oarsman

In order to improve the family economy and raise money to pay for his planned medical education, Whipple subsequently worked for one year as a teacher of the natural sciences at Dr. Holbrook's Military School in Ossining, New York. Here he also served as an athletic coach.

A student at Johns Hopkins
From early childhood George Whipple had been convinced that he was going to devote his life to medicine. With his thorough basic education and good marks he could expect to be admitted anywhere. The most reputable insitution in medical education at the time was the Johns Hopkins Medical School in Baltimore. This faculty had been established as recently as in 1893, but soon had attracted distinguished teachers and created a fruitful research climate. In this he was significantly influenced by his mother, who had learned about the university's outstanding teachers. Hopkins was the only medical school in the United States that requiered a bachelor's degree and a knowledge of Greek, Latin, French and German for admission.

The dean of the university was William Henry Welch (1850-1934), professor of pathology and at that time considered the foremost medical researcher in the U.S. Other great names at the faculty were William Osler (1840-1919) and Harvey Williams Cushing (1869-1939). In his first year at Hopkins, Whipple's training in physiological chemistry at Yale Qualified him to apply for a student teaching assistantship in John Jacob Abel's (1857-1938) Depeartment of Physiological Chemistry. His performance in his first year anatomy course was outstanding enough to win him a second-year appointment as a student instructor in anatomy. During that year Whipple's training was dominated by his introduction to pathology.

Young physician goes to science
Whipple graduated in medicine from Johns Hopkins in 1905. Standing fourth in his class of fifty-four students. At that time he had planned to specialise in pediatrics, but was offered a position as assistant under professor Welch. He was soon fascinated by pathology and sought a second year's appointment from Welch. Professor Welch’s saying : "...... another year of pathology, and he is anchored to it", was proved right.

In 1907, after two years of assistantship, Whipple embarked on his career as a pathologist and was encouraged to undertake research in descriptive and experimental pathology. Thus he never fulfilled his mother’s wish of seeing her son a general practitioner. On the other hand, however, she had ample reeason to enjoy her son’s brilliant career in research.

Doctor in Panama - researcher in Germany
In 1907, with the encouragement of Welch, Whipple went to the Gorgas Hospital in Panama to work for a year as a pathologist with Samuel Taylor Darling (1872-1925), the resident pathologist, and General William Crawford Gorgas (1854-1920). Whipple's main researches were concerned with anaemia and the physiology and pathology of the liver. He worked on anaemia caused by parasitic infections and especially on the lesions found in the intestinal tract in people suffering from these infections. He also studied the histology of the tissues in patients suffering from blackwater fever.

The salary for the work in Panama allowed him to travel to Europe before returning to Hopkins. He spent several months at Heidelberg in the laboratories of Ludolf Krehl (1861-1937) and Paul Morawitz (1879-1939). There he saw a first-class European laboratory in action, and participated briefly in some studies involving the experimental production of anaemia in rabbits.

A prize winner in Austria
Upon his return to Hopkins in 1909, Whipple began work on the pathologic disturbance of function such as that associated with acute chloroform poisoning and liver injury in the dog. Early interest in the pathogenesis of jaundice led him to submit an Essay on the Pathogenesis of Icterus in the blind competition for the Warren Triennial Prize of the Massachusetts General Hospital. Whipple was declared the winner of the prize in April 1910. This immensely pleased his department chairman Welch and added considerably to Whipple's growing renown. Soon thereafter he was offered professorships at the University of Pennsylvania and the University of California schools of medicine. He chose to turn them down and in 1911 was appointed an associate professor of pathology at Hopkins.

Whipple spent the spring and summer of 1911 in Vienna in the laboratory of Professor Hans Horst Meyer (1853-1939). There he learned how to produce the experimental porto-caval shunt in the dog known as the Eck fistula. The Russian physiologist Nikolai Vladimirovich Eck (1849–1917) introduced this technique in 1877 in order to study liver function. Using this technique in later years Whipple was able to study the effects of totally diverting the portal vein of blood flow on a number of hepatic functions in the dog.

Liver studies with Hooper
During the period 1907 to 1914 Whipple's research interests shifted from studies primarily concerned with histopathologic anatomy to problems in which altered functions could be studied with the tools of biochemistry and physiology.

During the last several years at Hopkins, he collaborated with Charles W. Hooper on a long series of studies in dogs on the origin and excretion of bile pigment and on the icterus as one manifestation of impaired hepatic function. They found that the liver cells had an almost limitless power of regeneration. He then became interested in jaundice, which is always associated with chloroform poisoning and injury to the liver. He studied the route by which the bile pigments pass into the blood and thus produce jaundice of various parts of the body and he found that the lympathic system was of little importance in transporting them.

Bile and blood
Whipples studies of bile pigments led to his interest in the body’s production of the oxygen-carrying haemoglobin, which also plays an important part in the production of bile pigment.

He later showed that histiocytes broke red cells down into bilirubin and bile pigments and by exclusion of the liver that histiocytes in other organs could also do this. He demonstrated that fibrinogen was made in the liver and proposed the term Thalassaemia for Cooley's anaemia. He also conducted experiments on artificial anaemia (1923-1925) demonstrating that iron is the strongest inorganic factor involved in the production of red blood cells. This work was done in collaboration with Charles W. Hooper.

His studies were briefly interrupted in the spring of 1914 when Whipple, at age thirty-four, married Katherine Ball Waring of Charleston, South Carolina. They had one son George Hoyt (born 1917), and one daughter Barbara (born 1921) and got seven grandchildren. In 1914 he also accepted an offer to become professor of experimental medicine and head of the newly established George Williams Hooper Foundation for Medical Research at the University of California School of Medicine in San Francisco. This Hooper family had no connection to the family of Whipple's colleague Charles W. Hooper

Over to California
Charles W. Hooper accompanied Whipple in the move from Hopkins to San Francisco. In spite of the many difficulties, they managed to establish a totally new laboratory where Whipple continued his research on bile pigment metabolism. This work culminated in a series of twelve publications between 1915 and 1917.

In the course of studying bile pigment metabolism and recognizing that blood red cells were the major normal source of bilirubin, Whipple and Hooper studied the effects of acute haemorrhagic anaemia and diet composition on bilirubin excretion and shifted the emphasis of their research to the study of the regeneration of red cells in simple anaemia.
Whipple was dean of the medical school in San Francisco during the years 1920 and 1921.

Over to Rochester
In 1921, Whipple, was offered the position of professor and dean of the School of Medicine and Dentistry at a university then planned in Rochester, New York. One of the men behind the initiative was George Eastman (1854-1932), the film magnate who had the headquarter if his corporation in this city.

With his research program in full swing, Whipple was reluctant to leave California, but he was finally won over as the offer was a rare opportunity to create a medical school from the ground up, with a full-time faculty in a physical setting conductive to easy exchange between clinical and preclinical disciplines.

Due to Eastman’s enormous contribution the university was able to receive its first students in 1925. Whipple, now also proving himself a skilled administrator, built the university into a reputable school, contributing importantly with his own lectures and seminaries. He served as dean and professor of pathology until his retirement in 1955, aged 77. However, he continued lecturing and giving seminaries until the beginning of the 1960s, when he was more than 80 years old. In the midst of his busy retirement he also found time to write a short autobiography in which he wrote "I would be remembered as a teacher".

Whipple's disease
The work leading to his description of the disease that bears his name, commenced in the pathological institute at Johns Hopkins University Medical School on May 9, 1907. Whipple conducted an autopsy on a 36 year old physician who had been a missionary in Turkey, domiciled in Constantinople. He had returned home to the U.S. due to a disease characterised by painful arthritis in multiple joints, gradual loss of weight and strength, abdominal pains and diarrhoea.

Based on his findings, Whipple incorrectly concluded that the condition arose from an abnormality of fat metabolism; hence, he coined the term “intestinal lipodystrophy”. Similar pathological findings were reported by William Henry Allchin (1846-1922) and Richard Grainger Hebb (1848-1918) at the Westminster Hospital in London in 1895 under the name “lymphangiectasis intestini.” This similarity went unnoticed until 1961, during an organising of the collection of unusual pathological findings at the Westminster Hospital. A. D. Morgan then reviewed the original tissue blocks, restained the sections, and demonstrated PAS (para-aminosalicylic acid)-positive macrophages.

Liver to the Nobel banquet
In 1920 Whipple began studies on the effects of various foodstuffs on blood regeneration. His research technician Frieda Robscheit-Robbins arrived in Rochester in December 1922 with forty of her special strain of dogs. Their experimental model was to bleed dogs to make them anaemic, and then to feed them diets that were restricted to food from one particular organ, for instance kidney, liver, brain, etc. Using this approach he found that liver was the most effective, followed by kidney and then muscle.

In 1925 they published the first of what was to become a series of eighteen papers on "Blood Regeneration in Severe Anemia". That report, clearly establishing the superior potency of fed liver in promoting the regeneration of haemoglobin of the anaemic dog, caught the attention of George Richards Minot (1885-1950) in Boston; he and William Parry Murphy (1892-1987) were preoccupied with the treatment of humans afflicted with pernicious anaemia for which there was at the time no cure. In a relatively short time they were able to demonstrate conclusively that a diet containing large amounts of raw or cooked beef liver produced phenomenal sustained remissions of pernicious anaemia. The effectiveness of liver feeding in the successful treatment of pernicious anaemia was soon widely confirmed and recognized internationally.

Between 1925 and 1930 Whipple and Robscheit-Robbins published a total of twenty-one papers describing the use of the standard anaemic dog to test a lengthy array of foods of animal and vegetable origin.

Their work culminated in Whipple, Minot, and Murphy sharing the Nobel prize in physiology or medicine in 1934. Although a modest man who did not enjoy being the centre of attention, it is obvious that the journey to Stockholm with his wife Katharine was one of the great moments in his life. His mother, Anna Whipple, cancelled her subscription to the reputable newspaper Boston Transcript, because she meant that it gave too much of the credit to Minot. During the Nobel dinner in Stockholm, Katharina Whipple was King Gustaf VI Adolph's partner. Upon his returning home Whipple showed his generosity by sharing a part of the prize money with his assistant over many years, Frieda Saur Robscheit-Robbins (1893-1973). His mother, too, received a substantial slump of money in recognition for her sacrifice.

Whipple's later years in Rochester were dominated to total commitment and devotion to his work as dean, as department head, as teacher of pathology and as medical researcher. He continued to use the standard anaemic dog to explore several questions bearing on the metabolism and synthesis of haemoglobin. In 1934 William B. Hawkins and Whipple used bile fistula dogs to determine the average life span of the red blood cell in the dog. They did that by timing the large increase in bile pigment that occurred at about 124 days after the acute massive regeneration of red cells in response to the massive haemolysis induced by the administration of phenylhydrazine.

A man of honours
Among the many honours and distinctions Whipple received are honorary doctorates of several American Universities as well as of the Universities of Athens and Glasgow; the Popular Science Monthly Gold Medal and Annual Award in 1930 (with Dr. Minot), and the William Wood Gerhard Gold Medal of the Pathological Society of Philadelphia, in 1934.

He was a Trustee of the Rockefeller Foundation, a Corresponding Member of the Association of Physicians in Vienna and of the Royal Society of Physicians in Budapest, and of the European Society of Haematology, as well as a Foreign Corresponding Member of the British Medical Association. He was an Honorary Member of the Pathological Society of Great Britain and Ireland, and of the American Philosophical Society and the Society of Experimental Biology and Medicine. He was, from 1936-1953, a member of the Board of Scientific Directors of the Rockefeller Institute, a member of the Board of Trustees of this Foundation from 1939-1953, Vice-Chairman of its Board of Trustees from 1953-1960, and in 1960 he was appointed Trustee Emeritus.

The old man and the sea
Whipple resigned as professor of pathology and dean of the school of medicine and retired two years later, aged 77. He had grown up in a landscape of wild nature that early woke the interest in hunting and fishing in the young George. He maintained these interests all through his life and, due to his vitality, was able to enjoy them even at very advanced ages. As a retiree he spent sustained periods of time every year in Florida with big game fishing, a pleasure he reluctantly gave up.

However, in 1962 two invitations to Washington forced him to cancel his trip to Florida. First he was to receive a high medical honour, second he and his wife with other Nobel prize laureates were invited to a gala dinner at the White House by president George F. Kennedy. Whipple immediately made it clear that he could not refuse the award, but said no thanks to the White House dinner, as it would cost one week of fishing, and that was out of the question.

He died in Rochester on February 1, 1976, aged 97.

"I would be remembered as a teacher"

I cannot recall a single instance where Whipple spoke in praise of his own work or ideas.
Leon L. Miller

"This report with its unequivocal emphasis on liver feeding is the most important single paper as regards George H. Whipple's world reputation as a scientist, in the whole of his immense lifetime list of more than 300 publications."
G. W. Corner, Whipple's biographer, on the second paper (1925) by Whipple and Robscheit-Robbins on "Blood Regeneration in Severe Anemia".

"Of the three prize winners, it was Whipple who first occupied himself with the investigations for which the prize is now awarded . . . Whipple's experiments were planned exceedingly well, and carried out very accurately, and consequently their results can lay claim to absolute reliability. These investigations and results of Whipple's gave Minot and Murphy the idea to see whether favourable resuots might also be obtained in the case of pernicious anaemia, an anaemia of quite different type, by making use of the foods of the kind that Whipple had found to yield favourable results in his experiments regarding anaemia from loss of blood."
Professor I. Holmgren, speaking for the Nobel Prize award committee.

Biographical
• Isidor Fischer (1869-1943), publisher:
  Biographisches Lexikon der hervorragenden Ärzte der letzten fünfzig Jahre.
  Berlin – Wien, Urban & Schwarzenberg, 1932.
• George H. Whipple. The Nobel Prize in Physiology or Medicine 1934.
  The Nobel Foundation 1934. On the Official Web Site of the Nobel Foundation.
• G. W. Corner:
  George Hoyt Whipple and his friends: The life story of a Nobel Prize pathologist.
  Philadelphia, J. B. Lippincott Company, 1963.
  Includes information on Frieda S. Robscheit-Robbins, Whipple's chief assistant
• Obituary notices. British Medical Journal, 1976, February 21, 1 (6007): 468.
• Barry G. Firkin and Judith A. Whitworth:
  Dictionary of Medical Eponyms.
  The Parthenon Publishing Group. 1989. New edition in 2002.
• Magnus Carlsson and Jan Wiberg:
  George Hoyt Whipple. Först med teorin om infektiös orsak till sällsynt systemsjukdom.
  Läkartidningen, Stockholm, 1989, 14: 99-102.
  In the series: Mannen bakom syndromet [The Man Behind the Syndrome].
• Leon L. Miller:
  George Hoyt Whipple. August 28, 1878–February 2, 1976.
  Biographical Memoirs. National Academy of Science. 1995, 66: 371-393. Works by George Hoyt Whipple: 1902
• The nucleoproteid of the suprarenal gland.
  Walter Jones and G. H. Whipple
  American Journal of Physiology, September 1, 1902, 7 (6): 423-434. 1907
• A hitherto undescribed disease characterized anatomically by deposits of fat and fatty acids in the intestinal and mesenteric lymphatic tissues.
  G. H. Whipple. The Johns Hopkins Hospital Bulletin, Baltimore, 1907, 18: 382-391.
  
  1909
• The presence of a weak hemolysin in the hook worm and its relation to the anemia of uncinariasis.
  G. H. Whipple. The Journal of Experimental Medicine, 1909, 11: 331-343. 1910
• The Cammidge test in experimental pancreatitis and other conditions.
  G. H. Whipple, B. S. Chaffee, B.S., and R. F. Fisher.
  Bulletin of The Johns Hopkins Hospital, 1910, 21: 339. 1911
• The pathogenesis of icterus. G. H. Whipple and J. H. King.
  The Journal of Experimental Medicine, 1911, 13: 115-135.
• Fibrinogen of the blood as influenced by the liver necrosis of chloroform poisoning.
  G. H. Whipple and S. H. Hurwitz.
  The Journal of Experimental Medicine, New York, 1911, 13: 136-161. 1912
• Reflex albuminuria. Renal albuminuria secondary to irritation of the urinary bladder.
  F. A. Evans, H. M. N. Wynne, and G. H. Whipple.
  Bulletin of The Johns Hopkins Hospital, 1912, 23: 311.
• Chloroform poisoning. Resistance of the pigeon, frog and terrapin to late chloroform poisoning.
  R. E. Mosiman and G. H. Whipple.
  Bulletin of The Johns Hopkins Hospital, 1912, 23: 323
• Pregnancy and chloroform anesthesia : A study of the maternal, placental, and fetal tissues.
  G. H. Whipple. The Journal of Experimental Medicine, 1912, 15: 246-258.
• Insusceptibility of pups to chloroform poisoning during the first three weeks of life.
  G. H. Whipple. The Journal of Experimental Medicine, 1912, 15: 259-269. 1913
• Tests for hepatic function and disease under experimental conditions.
  G. H. Whipple, V. R. Mason, and T. C. Peightal.
  Bulletin of The Johns Hopkins Hospital, 1913, 24: 207
• Tests for hepatic function and disease under experimental conditions. Phenoltetrachlorphthalein.
  G. H. Whipple, T. C. Peightal, and A. H. Clark.
  Bulletin of The Johns Hopkins Hospital 1913, 24: 343
• A test for hepatic injury: blood lipase. G. H. Whipple.
  Bulletin of The Johns Hopkins Hospital , 1913, 24: 357
• Intestinal obstruction : I. A study of a toxic substance produced in closed duodenal loops.
  G. H. Whipple, H. B. Stone, and B. M. Bernheim.
  The Journal of Experimental Medicine, 1913, 17, 286-306.
• Intestinal obstruction :II. A study of a toxic substance produced in closed duodenal loops.
  G. H. Whipple, H. B. Stone, and B. M. Bernheim.
  The Journal of Experimental Medicine, 1913, 17: 307-323.
• Icterus. A rapid change of hemoglobin to bile pigment in the circulation outside the liver.
  G. H. Whipple and C. W. Hooper.
  The Journal of Experimental Medicine, 1913, 17: 612-635. 1914
• Fibrinigen: I. An Investigation Concerning Its Origin and Destruction in the Body.
  G. H. Whipple. American Journal of Physiology, January 1, 1914, 33 (1): 50-69.
• Intestinal obstruction : VIII. The Experimental Study of Intestinal Obstruction.
  Harvey B. Stone, B. M. Bernheim, and G. H. Whipple
  Annals of Surgery, Philadelphia, May 1914, 59 (5): 714-726.
• Intestinal obstruction : III. The defensive mechanism of the immunized animal against duodenal loop poison.
  G. H. Whipple, H. B. Stone, and B. M. Bernheim.
  The Journal of Experimental Medicine, 1914, 19: 144-165.
• Intestinal obstruction : IV. The mechanism of absorption from the mucosa of closed duodenal loops.
  G. H. Whipple, H. B. Stone, and B. M. Bernheim.
  The Journal of Experimental Medicine, 1914, 19: 166-180.
• The adrenalin index of the suprarenal glands in health and disease.
  V. P. W. Sydenstricker, B. J. Delatour, and G. H. Whipple.
  The Journal of Experimental Medicine, 1914, 19: 536-551.
• Liver function as influenced by the ductless glands.
  G. H. Whipple and P. W. Christman.
  The Journal of Experimental Medicine, 1914, 20: 297-319. 1915
• Liver function as influenced by anesthetics and narcotics.
  G. H. Whipple and J. S. Speed.
  The Journal of Experimental Medicine, 1915, 21: 203-212. 1916
• Intestinal obstruction : V. Proteose intoxication.
  G. H. Whipple, F. H. Rodenbaugh, and A. R. Kilgore.
  The Journal of Experimental Medicine, 1916, 23: 123-135
• Icterus : A rapid change of hemoglobin to bile pigment in the pleural and peritoneal cavities.
  C. W. Hooper and G. H. Whipple.
  The Journal of Experimental Medicine, 1916, 23: 137-147.
• Intestinal obstruction : VI. A study of non-coagulable nitrogen of the blood.
  J. V. Cooke, F. H. Rodenbaugh, and G. H. Whipple.
  The Journal of Experimental Medicine, 1916, 23: 717-738
• Bile pigment metabolism: I. Bile Pigment Output and Diet Studies.
  C. W. Hooper and G. H. Whipple
  American Journal of Physiology, April 1, 1916, 40 (2): 332-348.
• Bile pigment metabolism: II. Bile Pigment Output Influenced by Diet.
  G. H. Whipple and C. W. Hooper.
  American Journal of Physiology, April 1, 1916, 40 (2): 349-359. 1917
• Bile pigment metabolism: III. Bile Pigment Output and Blood Feeding.
  G. H. Whipple and C. W. Hooper.
  American Journal of Physiology, January 1, 1817, 42 (2): 256-263.
• Bile pigment metabolism: IV. Influence of Fresh Bile Feeding upon Whole Bile and Bile Pigment Secretion.
  C. W. Hooper and G. H. Whipple.
  American Journal of Physiology, January 1, 1917, 42 (2): 264-279.
• Bile pigment metabolism: VI. Bile Pigment Output Influenced by the Eck Fistula.
  G. H. Whipple and C. W. Hooper.
  American Journal of Physiology, March 1, 1917, 42 (4): 544-557.
• Bile pigment metabolism: VII. Bile Pigment Output Influenced by Hemoglobin Injections, Anemia and Blood Regeneration.
  G. H. Whipple and C. W. Hooper
  American Journal of Physiology, May 1, 1917, 43 (2): 258-274.
• Bile pigment metabolism: VIII. Bile Pigment Output Influenced by Hemoglobin Injection; Splenectomy and Anemia.
  C. W. Hooper and G. H. Whipple
  American Journal of Physiology, May 1, 1917, 43 (2): 275-289.
• Bile pigment metabolism: IX. Bile Pigment Output Influenced by Hemoglobin Injection in the Combined Eck-Bile Fistula Dog.
  C. W. Hooper and G. H. Whipple
  American Journal of Physiology, May 1, 1917, 43 (2): 290-297.
• Animal experimentation and medical progress.—An argument in support of a bill now before the state legislature. G. H. Whipple.
  California State Journal of Medicine, San Francisco, March 1917, 15 (3): 68-70.
• Studies on the blood proteins: II. The albumin-globulin ratio in experimental intoxications and infections.
  S. H. Hurwitz and G. H. Whipple.
  The Journal of Experimental Medicine, 1917, 25: 231-253.
• Proteose intoxications and injury of body protein: I. The metabolism of fasting dogs following proteose injections.
  G. H. Whipple and J. V. Cooke.
  The Journal of Experimental Medicine, 1917, 25: 461-477.
• Proteose intoxications and injury of body protein : II. The metabolism of dogs with duodenal obstruction and isolated loops of intestine.
  G. H. Whipple, J. V. Cooke and T. Stearns.
  The Journal of Experimental Medicine, 1917, 25: 479-494. 1918
• Blood regeneration after simple anaemia. I. Curve of regeneration influenced by dietary factors.
  G. H. Whipple and C. W. Hooper.
  American Journal of Physiology, Washington DC, 1918, 45: 573-575.
• Regeneration of blood serum proteins. I. Influence of fasting upon curve of protein regeneration following plasma depletion.
  G. H. Whipple, W. J. Kerr and S. H. Hurwitz.
  American Journal of Physiology, 1918, 47: 356-369.
• Regeneration of blood serum protein: II. Influence of Diet upon Curve of Protein Regeneration Following Plasma Depletion.
  Wm. J. Kerr, S. H. Hurwitz, and G. H. Whipple
  American Journal of Physiology, December 1, 1918, 47 (3): 370-378.
• Regeneration of blood serum proteins: III. Liver Injury Alone: Liver Injury and Plasma Depletion: The Eck Fistula Combined with Plasma Depletion.
  Wm. J. Kerr, S. H. Hurwitz, and G. H. Whipple.
  American Journal of Physiology, December 1, 1918, 47 (3): 379-392.
• Proteose intoxications and injury of body protein : III. Toxic protein catabolism and its influence upon the non-protein nitrogen partition of the blood.
  G. H. Whipple and Donald D. Van Slyke.
  The Journal of Experimental Medicine, 1918, 28: 213-221.
• Proteose intoxications and injury of body protein : IV. The metabolism of dogs with sterile abscess, pancreatitis, and pleuritis.
  J. V. Cooke and G. H. Whipple.
  The Journal of Experimental Medicine, 1918, 28: 223-241.
• Proteose intoxications and injury of body protein : V. The increase in non-protein nitrogen of the blood in acute inflammatory processes and acute intoxications.
  J. V. Cooke and G. H. Whipple.
  The Journal of Experimental Medicine, 1918, 28: 243-252. 1919
• The rapid construction of liver cell protein on a strict carbohydrate diet contrasted with fasting mechanism of protein sparing action of carbohydrate. III.
  G. H. Whipple, N. C. Davis and C. C. Hall.
  Archives of Internal Medicine, Chicago, 1919, 23: 689-710.
• The metabolism of bile acids.II. Normal fluctuations in healthy bile fistula dogs.
  M. G. Foster, C. W. Hooper, and G. H. Whipple
  The Journal of Biological Chemistry, Baltimore, 1919, 38: 367-377.
• The metabolism of bile acids.III.. Administration by stomach of bile, bile acids, taurine, and cholic acid to show the influence upon bile acid elimination.
  M. G. Foster, C. W. Hooper, and G. H. Whipple
  The Journal of Biological Chemistry, 1919, 38: 379-392.
• The metabolism of bile acids. IV. Endogenous and exogenous factors.
  M. G. Foster, C. W. Hooper, and G. H. Whipple
  The Journal of Biological Chemistry, 1919, 38: 393-411.
• The metabolism of bile acids. V. Control of bile ingestion and food factors.
  M. G. Foster, C. W. Hooper, and G. H. Whipple.
  The Journal of Biological Chemistry, 1919, 38: 413-420
• The metabolism of bile acids. VI. Origin of taurocholic acid.
  M. G. Foster, C. W. Hooper, and G. H. Whipple
  The Journal of Biological Chemistry, 1919, 38: 421-433.
• I. Renal function influenced by intestinal obstruction.
  Irvine McQuarrie and G. H. Whipple.
  The Journal of Experimental Medicine, 1919, 29: 397-419.
• II. Renal function influenced by proteose intoxication.
  Irvine McQuarrie and G. H. Whipple.
  The Journal of Experimental Medicine, 1919, 29: 421-444. 1920
• III. Factors concerned in the perfusion of living organs and tissues: Artificial Solutions Substituted for Blood Serum and the Resulting Injury to Parenchyma Cells.
  E. Belt, H. P. Smith, and G. H. Whipple
  American Journal of Physiology, May 1, 1920, 52 (1): 101-120.
• Blood volume studies. I. Experimental control of a dye blood volume method.
  W. Hooper, H. P. Smith, A. E. Belt, G. H. Whipple.
  American Journal of Physiology, 1920, 51: 205-220.
• Blood volume studies: III. Behavior of Large Series of Dyes Introduced into the Circulating Blood.
  B. Dawson, H. M. Evans, and G. H. Whipple
  American Journal of Physiology, March 1, 1920, 51 (2): 232-256.
• Rapid blood plasma protein depletion and the curve of regeneration.
  H. P. Smith, A. E. Belt, and G. H. Whipple.
  American Journal of Physiology, May 1, 1920, 52 (1): 54-71.
• II. Shock as a manifestation of tissue injury following rapid plasma protein depletion: The Stabilizing Value of Plasma Proteins.
  G. H. Whipple, H. P. Smith, and A. E. Belt
  American Journal of Physiology, May 1, 1920, 52 (2): 72-100.
• Blood regeneration following simple anemia: I. Mixed diet Reaction.
  G. H. Whipple, C. W. Hooper, and F. S. Robscheit.
  American Journal of Physiology, 1920, 53: 151-166.
• Blood regeneration following simple anemia: II. Fasting Compared with Sugar Feeding.
  G. H. Whipple, C. W. Hooper, and F. S. Robscheit
  American Journal of Physiology, 1920, 53: 167-205.
• Blood regeneration following simple anemia: III. Influence of Bread and Milk, Crackermeal, Rice and Potato, Casein and Gliadin in Varying Amounts and Combinations.
  C. W. Hooper, F. S. Robscheit, and G. H. Whipple
  American Journal of Physiology, September 1, 1920, 53 (2): 206-235.
• Blood regeneration following simple anemia. IV. Influence of meat, liver and various extractives, alone or combined with standard diets.
  G. H. Whipple, F. S. Robscheit and C. W. Hooper.
  American Journal of Physiology, 1920, 53: 236-262.
• Blood regeneration following simple anemia: V. The Influence of Blaud's Pills and Hemoglobin.
  W. Hooper, F. S. Robscheit and G. H. Whipple.
  American Journal of Physiology, September 1, 1920, 53 (2): 263-282. 1921
• Studies of liver function. Benzoate administration and hippuric acid synthesis.
  G. D. Delprat and G. H. Whipple.
  The Journal of Biological Chemistry, 1921, 49 (1): 229-246.
• Value of animal experimentation to mankind. George Hoyt Whipple.
  American Journal of Public Health, New York, February 1921, 11 (2): 105-107.
• Blood volume studies: V. The Carbon Monoxide Method—Its Accuracy and Limitations.
  H. R. Arnold, E. B. Carrier, H. P. Smith, and G. H. Whipple.
  American Journal of Physiology, June 1, 56 (2): 313-327.
• Blood volume studies: VI. Plasma Volume as Determined by Hemoglobin Injection.
  F. W. Lee and G. H. Whipple.
  American Journal of Physiology, June 1, 1921, 56 (2): 328-335.
• Blood volume studies: VII. Comparative Values of Welcker, Carbon monoxide and Dye Methods for Blood Volume Determinations. Accurate Estimation of Absolute Blood Volume.
  H. P. Smith, H. R. Arnold, and G. H. Whipple
  American Journal of Physiology, June 1, 1921, 56 (2).336-360. 1922
• Blood fibrin studies: I. An Accurate Method for the Quantitative Analysis of Blood Fibrin in Small Amounts of Blood.
  D. P. Foster and G. H. Whipple.
  American Journal of Physiology, January 1, 1922, 58 (3): 365-378.
• Blood fibrin studies: II. Normal Fibrin Values and the Influence of Diet.
  P. Foster and G. H. Whipple.
  American Journal of Physiology, January 1, 1922, 58 (3): 379-392.
• Blood fibrin studies: IV. Fibrin Values Influenced by Cell Injury, Inflammation, Intoxication, Liver Injury and the Eck Fistula. Notes concerning the origin of fibrinogen in the body.
  P. Foster and G. H. Whipple
  American Journal of Physiology, January 1, 1922, 58 (3): 407-431.
• The physiology of the phenols. II. Absorption, Conjugation, and excretion.
  K. F. Pelkan and G. H. Whipple.
  The Journal of Biological Chemistry, 1922, 50 (2): 499-511.
• Studies of liver function. III. Phenol Conjugation as influenced by liver injury and insufficiency.
  K. F. Pelkan anf G. H. Whipple.
  The Journal of Biological Chemistry, 1922, 50: 513-526
• Roentgen ray intoxication : I. Unit dose over thorax negative—over abdomen lethal. epithelium of small intestine sensitive to x-rays.
  S. L. Warren and G. H. Whipple.
  The Journal of Experimental Medicine, 1922, 35: 187-202.
• Roentgen ray intoxication: II. A study of the sequence of clinical, anatomical, and histological changes following a unit dose of x-rays.
  S. L. Warren and G. H. Whipple.
  The Journal of Experimental Medicine, 1922, 35: 203-211.
• Roentgen ray intoxication: III. Speed of autolysis of various body tissues after lethal x-ray exposures. the remarkable disturbance in the epithelium of the small intestine.
  S. L. Warren and G. H. Whipple.
  The Journal of Experimental Medicine, 1922, 35: 213-224.
• Study of renal function in roentgen ray intoxication: resistance of renal epithelium to direct radiation. Irvine McQuarrie and G. H. Whipple.
  The Journal of Experimental Medicine, 1922, 35: 225-242.
• Variations in output of bile salts and pigments during 24-hour periods: Observations on Standard Bile Fistula Dogs.
  F. P. Wisner and G. H. Whipple.
  American Journal of Physiology, March 1, 1922, 60 (1): 119-133.
• The origin and significance of the constituents of the bile. G. H. Whipple.
  Physiological Reviews, Washington, July 1, 1922, 2 (3): 440-459.
• Determination of plasma and hemoglobin volumes after unit hemorrhages under controlled experimental conditions.
  B. Carrier, F. W. Lee, and G. H. Whipple.
  American Journal of Physiology, June 1, 61 (1): 138-148.
• II. Simultaneous determinations of plasma and hemoglobin volumes: Influence of Fluids by Mouth and Vigorous Exercise.
  W. Lee, E. B. Carrier, and G. H. Whipple.
  American Journal of Physiology, June 1, 1922, 61 (1): 149-158. 1923
• Roentgen ray intoxication: I. Bacterial invasion of the blood stream as influenced by x-ray destruction of the mucosal epithelium of the small intestine.
  S. L. Warren and G. H. Whipple.
  The Journal of Experimental Medicine, 1923, 38: 713-723.
• Roentgen ray intoxication:II. the cumulative effect or summation of x-ray exposures given at varying intervals.
  S. L. Warren and G. H. Whipple.
  The Journal of Experimental Medicine, 1923, 38: 725-730.
• Roentgen ray intoxication: III. The path of a beam of hard rays in the living organism.
  S. L. Warren and G. H. Whipple.
  The Journal of Experimental Medicine, 1923, 38: 731-739.
• Roentgen ray intoxication: IV. Intestinal lesions and acute intoxication produced by radiation in a variety of animals.
  S. L. Warren and G. H. Whipple
  The Journal of Experimental Medicine, 1923, 38: 741-752. 1924
• Bile salt metabolism. I. Influence of chloroform and phosphorus on bile fistula in dogs.
  Francis S. Smyth and G. H. Whipple.
  The Journal of Biological Chemistry, 1924, 59 (3): 623-636.
• Bile salt metabolism.II. Proteose and x-ray intoxication, thyroid and thyroxin.
  Francis S. Smyth and G. H. Whipple.
  The Journal of Biological Chemistry, 1924, 59 (3): 637-646.
• Bile salt metabolism. III. Gelatin, fish, yeast,cod liver, and meat extracts.
  Francis S. Smyth and G. H. Whipple.
  The Journal of Biological Chemistry, 1924, 59 (3): 647.
• Bile salt metabolism. IV. Negative influence of drugs, atropine, pilocarpine, phlorhizin, quinine, etc. Francis S. Smyth and G. H. Whipple.
  The Journal of Biological Chemistry, 1924, 59: 655-660. 1925
• Blood regeneration in severe anemia. I. Standard basal ration bread and experimental methods.
  H. Whipple and F. S. Robscheit-Robbins.
  American Journal of Physiology, 1925, 72: 395-407.
• Blood regeneration in severe anemia. II. Favorable influence of liver, heart and skeletal muscle in diet.
  F. S. Robscheit-Robbins and G. H. Whipple.
  American Journal of Physiology, May 1, 1925, 72 (3): 408-418.
• Blood regeneration in severe anemia: III. Iron Reaction Favorable—Arsenic and Germanium Dioxide Almost Inert.
  H. Whipple and F. S. Robscheit-Robbins
  American Journal of Physiology, May 1, 1925: 72 (3): 419-430.
• Blood regeneration in severe anemia: IV. Green Vegetable Feeding.
  F. S. Robscheit-Robbins and G. H. Whipple
  American Journal of Physiology, May 1, 1925, 72 (3): 431-435. 1926
• The identity of muscle hemoglobin and blood hemoglobin.
  R. P. Kennedy and G. H. Whipple.
  American Journal of Physiology, May 1, 1926, 76 (3): 685-692.
• The hemoglobin of striated muscle: I. Variations Due to Age and Exercise.
  G. H. Whipple.
  American Journal of Physiology, May 1, 1926, 76 (3): 693-707.
• The hemoglobin of striated muscle: II. Variations Due to Anemia and Paralysis.
  G. H. Whipple.
  American Journal of Physiology, May 1, 1926, 76 (3): 708-714.
• III. Muscle hemoglobin as a source of bile pigment.
  G. H. Whipple and F. S. Robscheit-Robbins
  American Journal of Physiology, November 1, 1926, 78 (3): 675-682. 1927
• Blood regeneration in severe anemia: V. Influence of Striated and Smooth Muscle Feeding.
  G. H. Whipple and F. S. Robscheit-Robbins.
  American Journal of Physiology, January 1, 1927, 79 (2): 260-270.
• Blood regeneration in severe anemia: VI. Influence of Kidney, Chicken and Fish Livers, and Whole Fish. F. S. Robscheit-Robbins and G. H. Whipple.
  American Journal of Physiology, January 1, 1927, 79 (2): 271-279.
• Blood regeneration in severe anemia: VII. Influence of Dairy Products on Hemoglobin Production.
  G. H. Whipple and F. S. Robscheit-Robbins.
  American Journal of Physiology, January 1, 1927, 79 (2): 280-288.
• Blood hemoglobin production and conservation in severe experimental anemia.
  F. S. Robscheit-Robbins and G. H. Whipple.
  Proceedings of The American Society of Experimental Pathology, April 1927.
• Blood regeneration in severe anemia: VIII. Influence of Bone Marrow, Spleen, Brains and Pancreas Feeding. The Question of Organic Iron in the Diet.
  G. H. Whipple and F. S. Robscheit-Robbins.
  American Journal of Physiology, April 1, 1927, 80 (2): 391-399.
• Blood regeneration in severe anemia: IX. Influence of Fresh and Dried Fruits.
  F. S. Robscheit-Robbins and G. H. Whipple.
  American Journal of Physiology, April 1, 1927, 80 (2): 400-410.
• Blood regeneration in severe anemia: X. Assimilation and Conservation of Bile Pigment, Blood Hemoglobin and Muscle Hemoglobin.
  G. H. Whipple and F. S. Robscheit-Robbins.
  American Journal of Physiology, December 1, 1927, 83 (1): 60-75.
• Blood regeneration in severe anemia: XI. Iron Effect Separated From Organ Effect in Diet.
  F. S. Robscheit-Robbins and G. H. Whipple.
  American Journal of Physiology, December 1, 1927, 83 (1): 76-83. 1928
• Blood regeneration in severe anemia. XII. Potent influence of inorganic ash of apricots, liver, kidney, and pineapple.
  Frieda S. Robscheit-Robbins, C. A. Elden, Warren M. Sperry, and G. H. Whipple.
  The Journal of Biological Chemistry, Baltimore, 1928, 79: 563-576.
• Blood regeneration in severe anemia. XIII. Influence of certain copper saltsupon hemoglobin output. C. A. Elden, Warren M. Sperry, Frieda S. Robscheit-Robbins, and G. H,. Whipple.
  The Journal of Biological Chemistry, 1928:79: 577-586
• Bile salt metabolism. I. Control diets, methods and fasting output.
  P. Smith, A. H. Groth, and G. H. Whipple.
  The Journal of Biological Chemistry, 1928: 80: 659-669.
• Bile salt metabolism. II. Influence of meat and meat extractives, liver and kidney, egg yolkand yeast in the diet.
  G. P. Smith and G. H. Whipple.
  The Journal of Biological Chemistry, 1928, 80 : 671-684.
• Bile salt metabolism. III. Tryptophane, tyrosine, and related substances as influencing bile salt output.
  G. H. Whipple and H. P. Smith
  The Journal of Biological Chemistry, 1928, 80: 685-695.
• Bile salt metabolism. IV. How much bile salt circulates in the body.
  G. H. Whipple and H. P. Smith.
  The Journal of Biological Chemistry, 1928, 80 : 697-707
• Muscle hemoglobin concentration during growth as influenced by diet factors.
  G. H. Whipple, A. H. Groth, and F. S. Robscheit-Robbins
  American Journal of Physiology, November 1, 1928, 87 (1): 185-191.
• The hemoglobin of smooth and striated muscle of the fowl.
  R. P. Kennedy and G. H. Whipple.
  American Journal of Physiology, November 1, 1928, 87 (1): 192-195. 1929
• Blood regeneration in severe anemia : XIV. A liver fraction potent in pernicious anemia fed alone and combined with whole liver, liver ash and fresh bile.
  F. S. Robscheit-Robbins and G. H. Whipple.
  The Journal of Experimental Medicine, 1929, 49: 215-227.
• Blood regeneration in severe anemia.XV. Liver fractions and potent factors.
  Warren M. Sperry, C. A. Elden, Frieda S. Robscheit-Robbins, and G. H,. Whipple.
  The Journal of Biological Chemistry, 1929, 81 (2): 251-265
• Various hemoglobins and their renal thresholds in the dog.
  E. J. Manwell and G. H. Whipple
  American Journal of Physiology, April 1, 1929, 88 (3): 420-431. 1930
• Blood regeneration in severe anemia: XVII. Influence of Manganese, Zinc, Copper, Aluminum, Iodine and Phosphates.
  F. S. Robscheit-Robbins and G. H. Whipple
  American Journal of Physiology, March 1, 1930, 92 (2): 378-387.
• Blood regeneration in severe anemia. XXI. A liver fraction potent in anemia due to hemorrhage.
  With F. S. Robscheit-Robbins and G. B. Walden.
  The American Journal of the Medical Sciences, Philadelphia, 1930, 179: 628-643.
• Bile salt metabolism. V. Casein, egg albumin, egg yolk, blood, and meat proteins as diet factors.
  H. P. Smith and G. H. Whipple.
  The Journal of Biological Chemistry, 1930, 89: 689-704.
• Bile salt metabolism. VI. Proline, tryptophane, and glycine in diet.
  G. H. Whipple and H. P. Smith.
  The Journal of Biological Chemistry, 1930. 89: 705-717.
• Bile salt metabolism. VII. Indene, hydrindene, and isatin.
  H. P. Smith and G. H. Whipple.
  The Journal of Biological Chemistry, 1930, 89: 719-725.
• Bile salt metabolism. VIII. Liver injury and liver stimulation.
  G. H. Whipple and H. P. Smith.
  The Journal of Biological Chemistry, 1930, 89: 727-738.
• Bile salt metabolism. IX. Eck fistula modifies bile salt output.
  H. P. Smith and G. H. Whipple.
  The Journal of Biological Chemistry, 1930, 89: 739-751.
• Blood regeneration in severe anemia: XVI. Optimum iron therapy and salt effect.
  G. H. Whipple and F. S. Robscheit-Robbins.
  American Journal of Physiology, March 1, 1930, 92 (2): 362-377.
• Blood regeneration in severe anemia: XVII. Influence of manganese, zinc, copper, aluminum, iodine and phosphates.
  F. S. Robscheit-Robbins and G. H. Whipple.
  American Journal of Physiology, 1930, 92: 378-387.
• Blood regeneration in severe anemia: XVIII. Influence of liver and blood sausage, veal, eggs, chicken and gelatin.
  G. H. Whipple and F. S. Robscheit-Robbins.
  American Journal of Physiology, March 1, 1930, 92 (2): 388-399.
• Blood regeneration in severe anemia: XIX. Influence of spinach, cabbage, onions and orange juice.
  F. S. Robscheit-Robbins and G. H. Whipple.
  American Journal of Physiology, March 1, 1930, 92 (2): 400-407.
• Blood regeneration in severe anemia: XX. Conservation of sheep and goose hemoglobin given intravenously to form dog hemoglobin.
  G. B. Taylor, E. J. Manwell, F. S. Robscheit-Robbins, and G. H. Whipple.
  American Journal of Physiology, 1930, 92: 408-413. 1931
• Effects of the intravenous injection of colloidal silver upon the hematopoietic system in dogs.
  Samuel S. Shouse and George H. Whipple.
  The Journal of Experimental Medicine, 1931, 53: 413-420.
• II. Aplasia of marrow and fatal intoxication in dogs produced by Roentgen radiation of all bones.
  With S. S. Shouse and S. L. Warren.
  The Journal of Experimental Medicine, 1931, 53: 421-435.
• Bile pigment and hemoglobin interrelation in normal dogs.
  Khun Sribhishaj, William B. Hawkins, and George H. Whipple.
  American Journal of Physiology, February 1, 96 (2): 449-462.
• II. Bile pigment and hemoglobin interrelation in anemic dogs.
  W. B. Hawkins, K. Sribhishaj, F. S. Robscheit-Robbins, and G. H. Whipple.
  American Journal of Physiology, February 1, 1931, 96 (2): 463-476. 1932
• Renal thresholds for hemoglobin in dogs : depression of threshold due to frequent hemoglobin injections and recovery during rest periods.
  John A. Lichty Jr., William H. Havill, and George H. Whipple.
  The Journal of Experimental Medicine, 1932, 55: 603-615.
• II. Renal threshold for hemoglobin in dogs uninfluenced by mercury poisoning.
  William H. Havill, John A. Lichty Jr, Gordon B. Taylor, and George H. Whipple.
  The Journal of Experimental Medicine, 1932, 55: 617-625.
• III. Tolerance for mercury poisoning increased by frequent hemoglobin injections.
  William H. Havill, John A. Lichty Jr., and George H. Whipple.
  The Journal of Experimental Medicine, 1932, 55: 627-635.
• IV. Hemoglobin injections and conservation of pigment by kidney, liver and spleen : the influence of diet and bleeding.
  William V. Newman and George H. Whipple.
  The Journal of Experimental Medicine, 1932, 55: 637-652.
• V. The iron content of blood free tissues and viscera : variations due to diet, anemia and hemoglobin injections.
  Robert P. Bogniard and George H. Whipple.
  The Journal of Experimental Medicine, 1932, 55: 653-665.
• Marrow hyperplasia and hemoglobin reserve in experimental anemia due to bleeding.
  L. W. F. Oehlbeck, F. S. Robscheit-Robbins, and G. H. Whipple.
  The Journal of Experimental Medicine, 1932, 56: 425-448. 1933
• Bone marrow volume in adult dogs.
  Edna Fairman and George H. Whipple
  American Journal of Physiology, April 30, 1933, 104 (2): 352-357.
• New formed hemoglobin and protein catabolism. Conservation of intermediates in the anemic dog on a protein-free diet.
  Floyd Shelton Daft, Frieda S. Robscheit-Robbins and G. H. Whipple.
  The Journal of Biological Chemistry, 1933, 103 (2): 495-510.
• Splenectomy in bile fistula dogs : bile pigment overproduction, anemia and intoxication.
  F. B. Queen, W. B. Hawkins, and G. H. Whipple.
  The Journal of Experimental Medicine, 1933, 57: 399-418.
• Hemoglobin production factors in the human liver : I. Normal, infection and intoxication.
  G. H. Whipple and F. S. Robscheit-Robbins.
  The Journal of Experimental Medicine, 1933, 57: 637-651.
• Hemoglobin production factors in the human liver : II. Liver degeneration, cancer, cirrhosis and hepatic insufficiency. F. S. Robscheit-Robbins and G. H. Whipple.
  The Journal of Experimental Medicine, 1933, 57: 653-670.
• Hemoglobin production factors in the human liver : III. Anemias—primary, aplastic and secondary—leukemias.
  G. H. Whipple and F. S. Robscheit-Robbins.
  The Journal of Experimental Medicine, 1933, 57: 671-687. 1934
• Blood plasma protein regeneration controlled by diet. I. Liver and casein and potent diet factors.
  With R. L. Holman and E. B. Mahoney.
  The Journal of Experimental Medicine, 1934, 59: 251-267.
• Blood plasma protein given by vein utilized in body metabolism. II. A dynamic equilibrium between plasma and tissue proteins.
  Russell L. Holman, Earle B. Mahoney, and G. H. Whipple.
  The Journal of Experimental Medicine, 1934, 59: 269-282.
• II. Bile cholesterol : fluctuations due to diet factors, bile salt, liver injury and hemolysis.
  Angus Wright and George H. Whipple.
  The Journal of Experimental Medicine, 1934, 59: 411-425.
• Hemoglobin production factors in the anemic horse liver.
  G. H. Whipple and F. S. Robscheit-Robbins.
  American Journal of Physiology, April 30, 1934, 108 (2): 270-278.
• Hemoglobin production factors in normal livers of domestic animals: Horse Liver Rates High and Beef Low. F. S. Robscheit-Robbins and G. H. Whipple.
  American Journal of Physiology, April 30, 1934, 108 (2): 279-284.
• Hemoglobin regeneration as influenced by diet and other factors.
  Nobel Lecture, December 12, 1934. 1935
• Reserve store of hemoglobin producing substances in growing dogs as influenced by diet.
  F. S. Robscheit-Robbins and G. H. Whipple
  American Journal of Physiology, April 30, 1935, 112 (1): 27-32.
• II. Hemoglobin and bile pigment overproduction in the splenectomized bile fistula dog.
  R. E. Knutti, W. B. Hawkins, and G. H. Whipple.
  The Journal of Experimental Medicine, 1935, 61: 127-138.
• Blood plasma protein regeneration controlled by diet : systematic standardization of food proteins for potency in protein regeneration. fasting and iron feeding.
  W. T. Pommerenke, H. B. Slavin, D. H. Kariher, and G. H. Whipple.
  The Journal of Experimental Medicine, 1935, 61: 261-282.
• Dog plasma protein given by vein utilized in body metabolism of dog : horse plasma and dog hemoglobin not similarly utilized.
  W. T. Pommerenke, H. B. Slavin, D. H. Kariher, and G. H. Whipple.
  The Journal of Experimental Medicine, 1935, 61: 283-297.
• New formed hemoglobin and protein catabolism in the anemic dog.
  Floyd Shelton Daft, Frieda S. Robscheit-Robbins and G. H. Whipple.
  The Journal of Biological Chemistry, 1935, 108: 487-496.
• Liver injury by chloroform, nitrogen metabolism, and conservation. Liver function and hemoglobin production in anemia.
  Floyd Shelton Daft, Frieda S. Robscheit-Robbins, and G. H. Whipple.
  The Journal of Biological Chemistry, 1935, 113 (2): 391-404.
• Anemias, Experimental and Clinical. George H. Whipple.
  Address given at the second general session of the California Medical Association, at its sixty-fourth annual session, Yosemite National Park, May 13 to 16, 1935.
  California and Western Medicine, San Francisco, July 1935, 43 (1): 1-4.
• Letters of appreciation from annual session Guest Speakers.
  George H. Whipple
  California and Western Medicine, July 1935, 43 (1): 103.
• Blood regeneration in severe anemia: Fractions of Kidney, Spleen and Heart Compared with Standared Liver Fractions.
  F. S. Robscheit-Robbins, G. B. Walden, and G. H. Whipple
  American Journal of Physiology, September 30, 1935, 113 (2): 467-475.
• Bile fistulas and related abnormalities : bleeding, osteoporosis, cholelithiasis and duodenal ulcers.
  W. B. Hawkins and G. H. Whipple.
  The Journal of Experimental Medicine, 1935, 62: 599-620. 1936
• Blood plasma protein regeneration controlled by diet. Effects of plant proteins compared with animal proteins. The influence of fasting and infection.
  With J. B. McNaught, V. C. Scott, and F. M. Woods.
  The Journal of Experimental Medicine, 1936, 64: 277-301.
• Iron metabolism: its absorption, storage and utilization in experimental anemia.
  P. F. Hahn and G. H. Whipple.
  The American Journal of the Medical Sciences, 1936, 191: 27-41.
• Control basal diets in anemic dogs: Method Factors and Hemoglobin Production.
  G. H. Whipple and F. S. Robscheit-Robbins.
  American Journal of Physiology, April 30, 1936, 115 (3): 651-661.
• Infection and intoxication : their influence upon hemoglobin production in experimental anemia.
  F. S. Robscheit-Robbins and G. H. Whipple.
  The Journal of Experimental Medicine, 1936, 63: 767-787. 1937
• Plasma protein given by vein and its influence upon body metabolism.
  Floyd Shelton Daft, Frieda S. Robscheit-Robbins, and G. H. Whipple.
  The Journal of Biological hemistry, 1938, 123 (1): 87-98.
• Blood plasma protein regeneration as influenced by infection, digestive disturbances, thyroid, and food proteins : a deficiency state related to protein depletion.
  S. C. Madden, P. M. Winslow, J. W. Rowland, and G. H. Whipple.
  The Journal of Experimental Medicine, 1937, 65: 431-454.
• Abscess nitrogen metabolism in anemicand non-anemic dog.
  Floyd Shelton Daft, Frieda S. Robscheit-Robbins, and G. H. Whipple.
  The Journal of Biological hemistry, 1937, 21 (1): 45-59.
• Liver function and blood plasma protein formation : normal and eck fistula dogs.
  R. E. Knutti, C. C. Erickson, S. C. Madden, P. E. Rekers, and G. H. Whipple.
  The Journal of Experimental Medicine, 1937, 65: 455-467.
• Globin utilization by the anemic dog to form new hemoglobin.
  F. S. Robscheit-Robbins and G. H. Whipple.
  The Journal of Experimental Medicine, 1937, 66: 565-578. 1938
• Lactoflavin (riboflavin) increases hemoglobin production in the anemic dog.
  Paul György, F. S. Robscheit-Robbins, and G. H. Whipple
  American Journal of Physiology, March 31, 1938, 122 (1): 154-159.
• The life cycle of the red blood cell in the dog.
  W. B. Hawkins and G. H. Whipple.
  merican Journal of Physiology, April 30, 1938, 122 (2): 418-427.
• Protein Production and Exchange in the Body including Hemoglobin Plasma Protein and Cell Protein.
  G. H. Whipple. The American Journal of the Medical Sciences, 1938, 196: 609.
• Hemoglobin production in anemia as influenced by the bile fistula.
  W. B. Hawkins, F. S. Robscheit-Robbins, and G. H. Whipple.
  The Journal of Experimental Medicine, 1938, 67: 89-110.
• Iron metabolism in experimental anemia : "availability of iron".
  P. F. Hahn and G. H. Whipple.
  The Journal of Experimental Medicine, 1938, 67: 259-265. 1939
• Hemoglobin production factors in the liver: Fish, Frog and Turtle Compared with Domestic Animals.
  F. S. Robscheit-Robbins and G. H. Whipple.
  American Journal of Physiology, April 30, 1939, 126 (1): 142-147.
• Hemoglobin production in anemia limited by low protein intake : influence of iron intake, protein supplements and fasting.
  P. F. Hahn and G. H. Whipple. M.D.
  The Journal of Experimental Medicine, 1939, 69: 315-326.
• Nephritis and its influence upon hemoglobin production in experimental anemia.
  G. H. Whipple and F. S. Robscheit-Robbins.
  The Journal of Experimental Medicine, 1939, 69: 485-498.
• Blood plasma protein production as influenced by amino acids : cystine emerges as a key amino acid under fixed conditions. S. C. Madden, W. A. Noehren, G. S. Waraich, and G. H. Whipple.
  The Journal of Experimental Medicine, 1939, 69: 721-738.
• Radioactive iron and its metabolism in anemia. Its absorption, transportation, and utilization.
  P. F. Hahn, W. F. Bale, E. O. Lawrence, and G. H. Whipple.
  The Journal of Experimental Medicine, 1939, 69: 739-753.
• Radioactive iron and its excretion in urine, bile, and feces.
  P. F. Hahn, W. F. Bale, R. A. Hettig, M. D. Kamen, and G. H. Whipple.
  The Journal of Experimental Medicine, 1939, 70: 443-451.
• Chloroform liver injury increases as protein stores decrease. Studies in nitrogen metabolism in these dogs.
  L. L. Miller and G. H. Whipple.
  The American Journal of the Medical Sciences, 1940, 199: 204-216.
• Plasma proteins: Their source, production and utilization.
  S. C. Madden and G. H. Whipple. Physiological Reviews, 1940, 20: 194-217.
• Methionine and cystine, specific protein factors preventing chloroform liver injury in protein-depleted dogs.
  L. L. Miller, J. F. Ross, and G. H. Whipple.
  The American Journal of the Medical Sciences, 1940, 200: 739-756.
• Blood plasma protein production and utilization : the influence of amino acids and of sterile abscesses.
  S. C. Madden, C. A. Finch, W. G. Swalbach, and G. H. Whipple.
  The Journal of Experimental Medicine, 1940, 71: 283-297.
• Amino acids and hemoglobin production in anemia.
  G. H. Whipple and F. S. Robscheit-Robbins:
  The Journal of Experimental Medicine, New York, 1940, 71: 569-583.
• The utilization of iron and the rapidity of hemoglobin formation in anemia due to blood loss.
  P. F. Hahn, J. F. Ross, W. F. Bale, and G. H. Whipple.
  The Journal of Experimental Medicine, 1940, 71: 731-736.
• Hemoglobin and plasma protein : simultaneous production during continued bleeding as influenced by diet protein and other factors.
  F. S. Robscheit-Robbins, S. C. Madden, A. P. Rowe, A. P. Turner, and G. H. Whipple.
  The Journal of Experimental Medicine, 1940, 72: 479-497. 1941
• Red Cell Volume Circulating and Total as Determined by Radio Iron.
  P. F. Hahn, W. M. Balfour, J. F. Ross, W. F. Bale, G. H. Whipple.
  Science, 1941, 93 (2404): 87-88.
• Blood plasma protein production as influenced by various degrees of hypoproteinemia and by amino acids.
  S. C. Madden, A. P. Turner, A. P. Rowe, and G. H. Whipple.
  The Journal of Experimental Medicine, 1941, 73: 571-580.
• Casein digests parenterally utilized to form blood plasma protein.
  S. C. Madden, L. J. Zeldis, A. D. Hengerer, L. L. Miller, A. P. Rowe, A. P. Turner, and G. H. Whipple.
  The Journal of Experimental Medicine, 1941, 73: 727-743.
• Hemoglobin production increases with severity of anemia.
  F. S. Robscheit-Robbins and G. H. Whipple.
  American Journal of Physiology, August 31, 1941, 134 (2): 263-267. 1942
• Copper and cobalt related hemoglobin production in experimental anemia.
  F. S. Robscheit-Robbins, and G. H. Whipple.
  The Journal of Experimental Medicine, 1942, 75: 481-487.
• Radioactive iron absorption in clinical conditions: normal, pregnancy, anemia, and hemochromatosis.
  W. M. Balfour, P. F. Hahn, W. F. Bale, W. T. Pommerenke, G. H. Whipple.
  The Journal of Experimental Medicine, 1942, 76: 15-30.
• Hemoglobin production factors in the human liver : anemias, hypoproteinemia, cirrhosis, pigment abnormalities, and pregancy.
  G. H. Whipple and F. S. Robscheit-Robbins.
  The Journal of Experimental Medicine, 1942, 76: 283-298 1943
• Ten amino acids essential for plasma protein production effective orally or intravenously.
  S. C. Madden, J. R. Carter, A. A. Kattus Jr., L. L. Miller, and G. H. Whipple.
  The Journal of Experimental Medicine, 1943, 77: 277-295.
• Hemoglobin and plasma protein. Ssimultaneous production during continued bleeding as influenced by amino acids, plasma, hemoglobin, and digests of serum, hemoglobin, and casein.
  F. S. Robscheit-Robbins, L. L. Miller, and G. H. Whipple.
  The Journal of Experimental Medicine, 1943, 77: 375-396.
• Radioactive iron absorption by the gastrointestinal tract: influence of anemia, anoxia, and antecedent feeding distribution in growing dogs.
  P. F. Hahn, W. F. Bale, J. F. Ross, W. M. Balfour, and G. H. Whipple. 1943.
  The Journal of Experimental Medicine, 1943, 78: 169-188. 1944
• Amino acid mixtures effective parenterally for long continued plasma protein production. Casein digests compared.
  With S. C. Madden, R. R. Woods, and F. W. Shull.
  The Journal of Experimental Medicine, 1944, 79: 607-624.
• Peritoneal absorption : red cells labeled by radio-iron hemoglobin move promptly from peritoneal cavity into the circulation.
  P. F. Hahn, L. L. Miller, F. S. Robscheit-Robbins, W. F. Bale, and G. H. Whipple.
  The Journal of Experimental Medicine, 1944, 80: 77-82.
• Gelatin—its usefulness and toxicity : blood protein production impaired by continued gelatin by vein.
  F. S. Robscheit-Robbins, L. L. Miller, and G. H. Whipple.
  The Journal of Experimental Medicine, 1944, 80:145-164.
• Plasma protein metabolism—normal and associated with shock : observations using protein labeled by heavy nitrogen in lysine.
  R. M. Fink, T. Enns, C. P. Kimball, H. E. Silberstein, W. F. Bale, S. C. Madden, and G. H. Whipple.
  The Journal of Experimental Medicine, 1944, 80: 455-475. 1945
• Eck fistula liver subnormal in producing hemoglobin and plasma proteins on diets rich in liver and iron.
  G. H. Whipple, F. S. Robscheit-Robbins, and W. B. Hawkins.
  The Journal of Experimental Medicine, 1945, 81. 171-191.
• Hemoglobin and plasma protein : their relation to internal body protein metabolism. L. L. Miller, F. S. Robscheit-Robbins, and G. H. Whipple.
  The Journal of Experimental Medicine, 1945, 81: 405-422.
• Tolerance to amino acid mixtures and casein digests given intravenously : glutamic acid responsible for reactions.
  S. C. Madden, R. R. Woods, F. W. Shull, J. H. Remington, and G. H. Whipple.
  The Journal of Experimental Medicine, 1945, 81: 439-448.
• Plasma protein production influenced by amino acid mixtures and lack of essential amino acids : A deficiency state related to unknown factors.
  S. C. Madden, F. W. Anderson, J. C. Donovan, and G. H. Whipple.
  The Journal of Experimental Medicine, 1945, 82: 77-92.
• Plasma protein production as influenced by parenteral protein digests, very high protein feeding, and red blood cell catabolism.
  S. C. Madden, A. A. Kattus Jr., J. R. Carter, L. L. Miller, and G. H. Whipple.
  The Journal of Experimental Medicine, 1945, 82 (3): 181-191.
• Maximal hemoglobin and plasma protein production under the stimulus of depletion. F. S. Robscheit-Robbins, L. L. Miller, and G. H. Whipple.
  The Journal of Experimental Medicine, 1945, 82: 311-316. 1946
• Effects of inflammation (turpentine abcess) on iron absorption.
  P. F. Hahn, W. F. Bale, and G. H. Whipple.
  Proceedings of the Society for Experimental Biology and Medicine, 1946, 61: 405.
• Plasma substitutes : human and animal globin related to the production of hemoglobin and plasma protein dog hemoglobin utilization improved by methionine but not by isoleucine.
  F. S. Robscheit-Robbins, L. L. Miller, E. L. Alling, and G. H. Whipple.
  The Journal of Experimental Medicine, 1946, 83: 355-371.
• Hemoglobin and plasma protein production : various proteins, concentrates, and digests influence blood protein production in anemia and hypoproteinemia.
  F. S. Robscheit-Robbins, L. L. Miller, and G. H. Whipple.
  The Journal of Experimental Medicine, 1946, 83: 463-475. 1947
• Plasma protein and hemoglobin production : deletion of individual amino acids from growth mixture of ten essential amino acids. significant changes in urinary nitrogen. F. S. Robscheit-Robbins, L. L. Miller, and G. H. Whipple.
  The Journal of Experimental Medicine, 1947, 85: 243-265.
• Anemia and hypoproteinemia : weight maintenance effected by food proteins but not by mixtures of pure amino acids.
  L. L. Miller, F. S. Robscheit-Robbins, and G. H. Whipple.
  The Journal of Experimental Medicine, 1947, 85: 267-275.
• Raiding of body tissue protein to form plasma protein and hemoglobin : what is premortal rise of urinary nitrogen?
  G. H. Whipple, L. L. Miller, and F. S. Robscheit-Robbins.
  The Journal of Experimental Medicine, 1947, 85: 277-286. 1948
• Protein metabolism and exchange as influenced by constriction of the vena cava : experimental ascites an internal plasmapheresis: sodium chloride and protein intake predominant factors.
  Frank W. McKee, Paul R. Schloerb, John A. Schilling, Garson H. Tishkoff , and George H. Whipple.
  The Journal of Experimental Medicine, 1948, 87: 457-471.
• Proteinuria related to hyperproteinemia in dogs following plasma given parenterally : a renal threshold for plasma proteins.
  Roger Terry, David R. Hawkins, Edwin H. Church, and G. H. Whipple.
  The Journal of Experimental Medicine, 1948, 87: 561-573.
• Parenteral plasma protein maintains nitrogen equilibrium over long periods.
  Roger Terry, William E. Sandrock, Robert E. Nye Jr., and G. H. Whipple.
  The Journal of Experimental Medicine, 1948, 87: 547-559. 1949
• Dietary Effects on Anemia plus Hypoproteinemia in Dogs. I. Some Proteins Further the Production of Hemoglobin and Other Plasma Protein Production.
  F. S. Robscheit-Robbin and G. H. Whipple.
  The Journal of Experimental Medicine, 1949, 89: 339-358.
• Plasma and red cell radioiron following intravenous injection : turpentine abscesses in normal and anemic dogs.
  L. Yuile, C. G. Bly, W. B. Stewart, A. J. Izzo, J. C. Wells, and G. H. Whipple.
  The Journal of Experimental Medicine, 1949, 90: 273-282.
• The use of radioactive lysine in studies of protein metabolism. Synthesis and utilization of plasma proteins.
  L. L. Miller, W. F. Bale, Charls L. Yuile, R. E. Masters, G. H. Tishkoff, and G. H. Whipple.
  The Journal of Experimental Medicine, 1949, 90: 297-313.
• Hemoglobin labeled by radioactive lysine : erythrocyte life cycle.
  W. F. Bale, C. L. Yuile, L. DeLaVergne, L. L. Miller, and G. H. Whipple.
  The Journal of Experimental Medicine, 1949, 90: 315-320.
• Protein metabolism and exchange as influenced by constriction of the vena cava : II. Effects of parenterally administered plasma, amino acid mixture, and ascitic fluid, and of orally administered ascitic fluid in the experimental ascitic dog.
  Frank W. McKee, Robert E. Hyatt, William G. Wilt Jr., Garson H. Tishkoff, and George H. Whipple.
  The Journal of Experimental Medicine, 1949, 90: 447-459. 1950
• The circulation of ascitic fluid : interchange of plasma and ascitic fluid protein as studied by means of c14-labeled lysine in dogs with constriction of the vena cava.
  Frank W. McKee, William G. Wilt Jr., Robert E. Hyatt, and George H. Whipple.
  The Journal of Experimental Medicine, 1950, 91: 115-122.
• Radioiron absorption in anemic dogs; fluctuations in the mucosal block and evidence for a gradient of absorption in the gastrointestinal tract.
  Wellington B. Stewart, Charles L. Yuile, Herbert A. Clairborne, Richard T. Snowman, and George H. Whipple.
  The Journal of Experimental Medicine, October 1, 1950, 92 (4): 375-382. 1951
• Conversion of plasma protein to tissue protein without evidence of protein breakdown : results of giving plasma protein labeled with carbon14 parenterally to dogs.
  L. Yuile, B. G. Lamson, L. L. Miller, and G. H. Whipple.
  The Journal of Experimental Medicine, 1951, 93: 539-557.
• Anemia plus hypoproteinemia in dogs; various proteins in diet show various patterns in blood protein production; beef muscle,. egg, lactalbumin, fibrin, viscera, and supplements.
  G. H. Whipple, F. S. Robscheit-Robbins.
  The Journal of Experimental Medicine, September 1951, 94 (3): 223-242. 1952
• Albumin and globulin circulation in experimental ascites; relative rates of interchange between plasma and ascitic fluid studied with C14-labeled proteins.
  Frank W. McKee, C. L. Yule, B. G. Lamson, G. H. Whipple.
  The Journal of Experimental Medicine, February 1952, 95 (2): 161-172.
• Plasma protein labeled with lysine-epsilon-C14; its oral feeding and related protein metabolism in the dog.
  C. L. Yuile, A. E. O'Dea, F. V. Lucas, G. H. Whipple.
  The Journal of Experimental Medicine, September 1952, 96 (3): 247-254. 1953
• Red Cell Stroma in Dogs. Variations in the Stroma Protein and Lipid Fractions Related to Experimental Conditions.
  G. H. Tishkoff, F. S. Robscheit-Robbins, and G. H. Whipple.
  Blood, May 1953, 8 (5): 459-468.
• Inflammation and protein metabolism studies of carbon-14-labeled proteins in dogs with sterile abscesses.
  C. L. Yuile, F. V. Lucas, C. K. Jones, S. J. Chapin, G. H. Whipple.
  The Journal of Experimental Medicine, August, 1953, 98 (2): 173-194. 1954
• Luminous bacteria as a radiobiological test organism. G. H. Whipple.
  Journal of Cellular Physiology, Philadelphia, June 1954, 43 (3): 415-423. 1955
• The placenta and protein metabolism; transfer studies using carbon 14-labeled proteins in dogs.
  G. H. Whipple, R. B. Hill Jr., R. Terry, F. V. Lucas, and C. L. Yuile.
  The Journal of Experimental Medicine, June 1, 1955, 101 (6): 617-626.
• Red cell stroma in dogs: stroma protein and stroma lipides vary in different types of anemia.
  S. Robscheit-Robbins and G. H. Whipple.
  The Journal of Experimental Medicine, 1955, 102: 705-711.
• Red cell stroma and hemoglobin metabolism in anemic dogs: regeneration of red cell proteins labeled with c14 lysine.
  With G. H. Tishkoff, C. L. Yuile, and F. S. Robscheit-Robbins
  The Journal of Experimental Medicine, December 1, 1955, 102 (6): 713-723.
• Red cell stroma protein rich in vitamin b12 during active regeneration: anemia studies using radioactive cobalt b12 in dogs. G. H. Whipple, F. S. Robscheit-Robbins and W. F. Bale.
  The Journal of Experimental Medicine, December 1, 1955, 102 (6): 725-731. 1958
• Vitamin B12Co60 distribution in dog tissue during many months; red cell stroma with labeled B12 in hemolytic anemia.
  W. D. Woods, W. B. Hawkins, and G. H. Whipple.
  The Journal of Experimental Medicine, July 1, 1958, 108 (1): 1-8. 1959
• Autobiographical sketch.
  Perspectives in Biology and Medicine, Baltimore, 1959, 2 (3): 253-289.
• Depletion of reserve protein from extravascular extracellular fluid; C14 labeling of plasma proteins in dogs after plasmapheresis.
  C. L. Yuile, F. V. Lucas, R. D. Neubecker, C. G. Cochrane, G. H. Whipple.
  The Journal of Experimental Medicine, February 1, 1959, 109 (2): 165-171. 1960
• Vitamin B12 Co-60 readily passes the placenta into fetal organs and nursing provides B12 from mother to pup. A Record of its distribution.
  W. D. Woods, W. B. Hawkins, and G. H. Whipple.
  The Journal of Experimental Medicine, September 1, 1960, 112: 431-444.
• Rate of formation of red blood cells: the standard anemic dog. G. H. Whipple.
  Methods in Medical Research, Chicago, 1960, 8: 55-58.