A Selected Chronological Bibliography of Biology and Medicine


Part 4A


1925 — 1936



Compiled by James Southworth Steen, Ph.D.

Delta State University


Dedicated to my loving family


This document celebrates those secondary authors and laboratory technicians without whom most of this great labor of discovery would have proved impossible.


Please forward any editorial comments to: James S. Steen, Ph.D., Professor Emeritus, jsteen08@bellsouth.net



"The key to every biological problem must finally be sought in the cell." Edmund Beecher Wilson {Wilson, 1925 #8391}.


“When we consider what religion is for mankind, and what science is, it is no exaggeration to say that the future course of history depends upon the decision of this generation as to the relations between them.” Alfred North Whitehead {Whitehead, 1925 #14608}.


“Almost all new ideas have a certain aspect of foolishness when they are first proposed.” Alfred North Whitehead {Whitehead, 1925 #14608}.


Richard Adolf Zsigmondy (AT-DE) received the Nobel Prize in Chemistry for his demonstration of the heterogeneous nature of colloid solutions and for the methods he used, which have since become fundamental in modern colloid chemistry. He developed a dark field ultra-microscope that could detect particles with diameters below the wavelengths of visible light.


Ronald Aylmer Fisher (GB-AU) invented the statistical method known as analysis of variance {Fisher, 1925 #8532}.


Walter S. Abbott (US) developed a method for computing the effectiveness of insecticides {Abbott, 1925 #9320}.


William S. Hoffman (US) isolated adenine nucleotide in crystalline form from pig blood {Hoffman, 1925 #17088}.


John Mason Gulland (GB) and Robert Robinson (GB) worked out the structure of morphine {Gulland, 1925 #12109}.


Treat Baldwin Johnson (US) and Robert D. Cogshill (US) discovered the occurrence of 5-methylcytosine in nature (tubercle bacteria) {Johnson, 1925 #12112}.

Rollin Douglas Hotchkiss (US) and Gerard R. Wyatt (CA) discovered 5-methylcytosine in plant and animal tissues {Hotchkiss, 1948 #12113;Wyatt, 1951 #12114}.


Gottfried Koller (DE), Earle Bryant Perkins (US), Theodore Snook (US), and Benjamin Kropp (US) found that there is produced in the eyestalks of crustaceans a hormone which, when carried in the blood stream, is effective in inducing chromatophore changes. Koller named this hormone contractin {Koller, 1925 #16847;Koller, 1927 #17563;Koller, 1928 #19445;Perkins, 1928 #14588;Perkins, 1931 #19448;Perkins, 1932 #19449}.


Bernhard Zondek (DE-IL) and Benno Brahn (DE) isolated an estrogenic hormone from the ovarian follicle {Zondek, 1926 #3384;Zondek, 1925 #3383}.


Edouard Chatton (FR) first used the terms prokaryote and eukaryote in his 1925 paper when he suggested that the most significant taxonomic distinction among life forms today is not between plants and animals but between cells with nuclei, eukaryotes, and cells without them, prokaryotes. Pro means before and eu means true, karyote refers to a nut or kernel {Chatton, 1925 #1958;Chatton, 1938 #6321}.“Protozoologists agree today in considering the flagellated autotrophs the most primitive of the Protozoa possessing a true nucleus, Eukaryotes (a group which also includes the plants and the Metazoans), because they alone have the power to completely synthesize their protoplasm from a mineral milieu. Heterotrophic organisms are therefore dependent on them for their existence as well as on chemotrophic Prokaryotes and autotrophs (nitrifying and sulphurous bacteria, Cyanophyceae.” This is a translation by Jan Sapp (CA) from the 1938 reference {Sapp, 2005 #19875}.

Eukaryotes are now placed in the domain Eucarya while prokaryotes are placed in either the domain Bacteria or Archaea.

André Michel Lwoff (FR) used the word eukaryote {Lwoff, 1938 #16381}.

Ellsworth C. Dougherty (US) proposed the prokaryotic-eukaryotic dichotomy. He used the words eukaryon (Greek: true kernel) for the nucleus of “higher organisms” and prokaryon (Greek: before kernel) for the moneran nucleus. {Dougherty, 1957 #19872}.

Roger Yate Stanier (CA) and Cornelis Bernardus Kees van Niel (NL-US) later defined bacteria by conceptualizing two new biological entities, "prokaryotes" and "eukaryotes." They argued that the cells of all living things were either prokaryotic or eukaryotic, depending on their pattern of cellular organization. They defined eukaryotes as cells containing membrane-bound structures called organelles, the most important of which was the nucleus. By this definition, all cells of multi-cellular plants and animals were eukaryotes. Cells that lacked membrane-bound cell nuclei, like bacteria and blue-green algae, were designated prokaryotes. The term prokaryote implicitly elevated bacteria to equivalent biological status with all other organisms, to be known as eukaryotes {Stanier, 1962 #12332}. This represents the resurrection and embellishment of an idea first conceived by Edouard Chatton (FR).

Robert George Everitt Murray (CA) proposed Procaryotae as a taxon “at the highest level” and described it as “a kingdom of microbes…characterized by the possession of nucleoplasm devoid of basic protein and not bounded from cytoplasm by a nuclear membrane.” He suggested Eucaryotae as a possible taxon at the same level to include other protists, plants, and animals {Murray, 1968 #19874}.


Erwin Broun Fred (US), Ira Lawrence Baldwin (US) and Elizabeth McCoy (US) published the definitive text on nitrogen fixation {Fred, 1932 #19855}.


Agnes Robertson Arber (GB) authored Monocotyledons; a Morphological Study, which developed the thesis, first suggested by Augustin Pyramus de Candolle (CH), that the monocot leaf is derived from a dicot petiole (phyllode theory) {Arber, 1925 #7465}.


Thurlow Nelson (US) implicated the ctenophore Mnemiopsis as a predator on oyster larvae in inland coastal waterways along the New Jersey coast {Nelson, 1925 #26750}.


Emil-Karl Frey (DE) observed in 1925 a considerable reduction in arterial blood pressure when he injected human urine into dogs. Unlike many other contemporary scientists, he did not attribute this effect to a toxic action of urine, but rather as the specific activity of an unknown substance with potential biological functions {Frey, 1926a #19806;Frey, 1926b #19807}.

Heinrich Kraut (DE), Emil-Karl Frey (DE) and Eugen Werle (DE) reasoned that, “It is a substance that probably originates from several organs, is eliminated by the kidneys and has a pronounced cardioactive and vasoactive effect: a substance that is assigned the role of a hormone in the organism”. This F-substance was then called kallikrein, since it was considered to have originated in the pancreas (Greek synonym: kallikreas) {Kraut, 1930 #19808}.

Eugen Werle (DE), W. Götze (DE), A. Keppler (DE), and M. Grunz (DE) identified kallikrein as a proteolytic enzyme (‘ferment’) that liberates the biologically highly active, basic polypeptide ‘DK’ or kallidin (i.e lys-bradykinin) from a blood plasma protein called kallidinogen or kininogen (H- and L-kininogen) {Werle, 1937 #2080;Werle, 1939 #2081}. Its name was later changed to kallidin and that of the precursor to kallidinogen. This work laid the foundation for understanding the system that we refer to today as the kallikrein-kinin system (cascade).


George Edward Briggs (GB) and John Burdon Sanderson Haldane (GB-IN) made important refinements in the theory of enzyme kinetics with their steady-state derivation of the single-substrate enzyme saturation curve {Briggs, 1925 #5527}.


Earl Perry Cark (US) and James Bertram Collip (CA) reported an important improvement in the methodology for the determination of blood serum calcium {Clark, 1925 #9321}.


Cyrus Hartwell Fiske (US) and Yellapragada Subbarow (IN-US) developed a colorimetric method useful for the detection of phosphorus in organic material {Fiske, 1925 #9322}.


Evert Gorter (NL) and Francois Grendel (NL) determined that the area of the monomolecular film formed on a Langmuir trough by the membrane lipids was double the surface area of the erythrocytes from which the lipids had been extracted, suggesting that the cell membrane is bimolecular {Gorter, 1925 #11048}. This was the first evidence that cell membranes are bi-layered.


Hans Molisch (CZ), in 1925, obtained the evolution of oxygen by illuminating preparations of dried leaves {Molisch, 1938 #6485}.


Gottfried Samuel Fraenkel (DE-US) discovered that statocysts on jellyfish medussa respond to gravity {Fraenkel, 1925 #26681}.


William Rowan (CH-GB-CA) tested the effect of photoperiodism on migratory drive and reproductive histology in birds. He proved that the annual cycle of changing day-length is the major stimulus for bird migration and is associated with profound physiological change within birds {Rowan, 1925 #12152;Rowan, 1926 #8935;Rowan, 1929 #26646;Rowan, 1931 #5184;Rowan, 1932 #26648}. Note: Before Rowan, it was unknown what environmental cues animals used to time the seasonal changes in their biology. See, Piersma, 1998.


Robert Kyle Burns, Jr. (US) established the adequacy of the hormonal theory of sex determination as a general mechanism for vertebrates {Burns, 1925 #10572;Burns, 1939a #10573;Burns, 1939b #10574;Burns, 1939c #10575;Burns, 1942 #10576;Burns, 1955 #10577}. See Frank Rattray Lillie, 1916.


Carl Ferdinand Cori (CZ-US) described the rapid absorption of galactose in the intestine, galactose being the most active sugar in this respect {Cori, 1925a #23141}.


Josef von Halban (AT) and Robert Koehler (AT) proved that cessation of internal secretion of the corpus luteum is responsible for the menstrual bleeding in humans {von Halban, 1925 #23289}.


Ernest Henry Starling (GB) and Ernest Basil Verney (GB) isolated dog kidneys, which they supplied with oxygenated blood at controlled flow rates, temperature, and pressure. They found they could produce regular flow rates of normal urine. From their analysis of the urine they determined that the glomeruli generate a protein-free filtrate. They artificially blocked tubular metabolic activity with hydrocyanic acid, collected ureter filtrate, and found that water, chloride, bicarbonate, and glucose are normally reabsorbed from the glomerular filtrate by the tubular cells. They also found that pituitrin (a proprietary preparation of the posterior lobe of the pituitary gland) influenced the re-absorption mechanisms for water and chloride and that without it the mammalian kidney reverted to a urine characteristic of fish and amphibian classes {Starling, 1925 #3364}. Note: Vasopressin would later be isolated as the active ingredient in pituitrin.


Edwin B. Hart (US), Harry Steenbock (US), Conrad Arnold Elvehjem (US), and James Waddell (US) demonstrated that when rabbits were induced into a state of anemia by milk diet, the addition of ferric salts alone was insufficient to bring about recovery. They discovered that the additional presence of copper is necessary for the animals to synthesize the pyrrol nucleus of hemoglobin {Hart, 1925 #3063;Hart, 1928 #3064}.


Louis Sigurd Fridericia (DK) and Eiler Holm (DK) demonstrated that vitamin A (retinol) is necessary for normal night vision in rats. They also found that both the retinas of A-deficient rats and of rats whose eyes had been strongly illuminated were depleted of visual purple {Fridericia, 1925 #2934}.


Gaston Ramon (FR) and Pierre A. Descombey (FR) produced a tetanus vaccine from formaldehyde-treated tetanus toxin {Ramon, 1925b #8239}.


Simeon Burt Wolbach (US), Otto A. Bessey (US), and Percy R. Howe (US) concluded that vitamin A (retinol) deficiency in diets led to improper development of teeth and retardation of the growth of the skeleton and that vitamin C deficiency led to a deficiency of formation of intercellular cement substance leading to fragility of blood capillaries {Wolbach, 1925 #2944;Wolbach, 1926 #2980;Wolbach, 1928 #2945;Wolbach, 1933a #2946;Wolbach, 1933b #2947;Wolbach, 1941 #2943}.


J.H.C. Ruyter (NL) was the first to call attention to the structural peculiarities of unique cells lining the afferent arteriole as it approaches the glomerulus in mice and rats. He suggested that by swelling these cells could occlude the lumen of the afferent arterioles and thereby regulate blood flow to the glomerular capillaries {Ruyter, 1925 #6213}.

Charles Oberling (FR) also reported a distinct group of cells at the same site, this time in the human kidney {Oberling, 1927 #25170}.

Norbert Oscar Jean Goormaghtigh (BE) named the juxtaglomerular apparatus and suggested that it might have an endocrine function {Goormaghtigh, 1932 #6214}.

Norbert Oscar Jean Goormaghtigh (BE) and Keith S. Grimson (US) confirmed that the modified cells in the afferent arteriolar wall of the juxtaglomerular apparatus are endocrine in nature {Goormaghtigh, 1937 #19785;Goormaghtigh, 1939a #16849;Goormaghtigh, 1939b #19787;Goormaghtigh, 1940a #16850;Goormaghtigh, 1940b #19786;Goormaghtigh, 1944 #16851;Goormaghtigh, 1945 #6215;Goormaghtigh, 1951 #19788}.

George White Pickering (US), Myron Prinzmetal (US), Juan Carlos Fasciolo (AR), Bernardo Alberto Houssay (AR), and Alberto C. Taquini (AR) rediscovered renin from the kidney as an inducer of hypertension {Pickering, 1938 #6209;Fasciolo, 1938 #7801}. See Tigerstedt, 1898.

Eduardo Braun-Menéndez (AR), Juan Carlos Fasciolo (AR), Luis Federico Leloir (AR), Juan M. Muñoz (AR), and Irvine Heinly Page (US) determined that renin is an enzyme which acts on a substrate (hypertensinogen) in the plasma converting it to a decapeptide (angiotensin I) {Braun-Menéndez, 1939 #6210;Page, 1939 #6211}.

Leonard T. Skeggs, Jr. (US), Kenneth E. Lentz (US), Joseph R. Kahn (US), Norman P. Shumway (US), and Kenneth R. Woods (US) determined that a serum enzyme (angiotensinase) converts the inactive decapeptide (angiotensin I) to the active octapeptide (angiotensin II) {Skeggs, 1956 #6212}. This activity occurs primarily in the lungs.

William Stanley Peart (GB) demonstrated that the juxtaglomerular apparatus cells produce renin within the kidney {Peart, 1977 #16567}.


Hugo Fricke (US) measured the electrical capacitance of the surface membrane of erythrocytes, using a high-frequency alternating current bridge {Fricke, 1925a #11049;Fricke, 1925b #11050}.


Edward M. East (US) and Albert J. Mangelsdorf (US), using Nicotiana tabacum as their research material, laid the foundation for understanding gametophytic self-incompatibility {East, 1925 #1985}.


Thomas Hunt Morgan (US), Calvin Blackman Bridges (US) and Alfred Henry Sturtevant (US) showed that chromosome imbalance could produce intersex types in Drosophila.

They also observed that occasionally crosses produced female-biased sex ratios, a clear violation of Mendel’s first law. Daughters from the distorted brood produced an equal number of males and females, but half of their sons produced female-biased sex ratios, whatever the origin of their mates. The remaining grandsons produced normal sex ratios and were shown to have lost the trait {Morgan, 1925 #1213}.

David Policansky (US) and John Ellison (US) showed that in the Drosophila males producing biased sex ratio offspring there is a high mortality among sperm bearing the Y chromosome. They concluded that in some way a gene on the X chromosome kills sperm bearing a Y chromosome {Policansky, 1970 #6084}.


Felix Bernstein (DE) proposed that the human ABO blood group is inherited in a multiple allele pattern {Bernstein, 1925 #14662}.


Hermann Joseph Muller, Jr. (US) reported on the mental traits and heredity as studied in a case of identical twins reared apart {Muller, 1925 #12347}.

Horatio H. Newman (US), Frank N. Freeman (US), and Karl J. Holzinger (US) presented a large study of heredity and environment in the development of 100 pairs of twins {Newman, 1937 #12348}.


Sears P. Doolittle (US) and Lewis Ralph Jones (US) demonstrated that Macrosiphum pisi Harris (pea aphid) acts as a vector for pea mosaic virus {Doolittle, 1925 #1616}.


Henri Vallée (FR), Henri Carré (FR), Paul Rinjard (FR), Otto Waldmann (DE), K. Kobe (DE), Gottfried Pyl (DE), Karl Otto Hobohm (DE), Hubert Moehlmann (DE), and H.S. Frenkel (NL) produced vaccines for foot-and-mouth disease {Frenkel, 1947 #25969;Frenkel, 1951 #25970;Vallée, 1925 #25967;Vallée, 1926 #26999;Waldmann, 1937 #25968;Waldmann, 1941 #25971}.


Carl Hamilton Browning (GB) defined substances that incite the formation of antibodies and react with them as antigens {Browning, 1925 #97}.


A smallpox epidemic emerged in Milwaukee, Wisconsin resulting in 386 cases, of whom 87 people died (about 23%). Of the 386 cases, 327 had never been vaccinated, 46 had been vaccinated 10 or more years previously, 11 had been vaccinated more than 5 and less than 10 years previously, 2 had an uncertain vaccination history, and none had been vaccinated within the previous 5 years. Health workers performed about 400,000 vaccinations during the epidemic, and officials enforced quarantine measures and isolation of those infected {Members,  #26005}.


John Cunningham (GB), working in India, found that the organism, Borrelia carteri, occurs in two antigenic varieties in human infections of relapsing fever {Cunningham, 1925 #98}. John Cunningham (GB), J.H. Theodore (GB), and A.G.L. Fraser (GB) extended the number of antigenic varieties to nine {Cunningham, 1934 #19791}.


William George Savage (GB) and Philip Bruce White (GB) laid the groundwork for antigenic analysis of the Salmonella group {Savage, 1925 #9086}.

Fritz Kaufmann (DK) greatly extended and refined Savage and White’s work {Kaufmann, 1954 #17828}.


Gaston Ramon (FR) demonstrated that it was possible to augment the antitoxin response to diphtheria and tetanus by administering vaccines with pyogenic bacteria or with various additional compounds (his favorite substance was tapioca). This represents the first occasion on which immunological adjuvants were used. Ramon called them adjuvantes et stimulantes de l’immunite {Ramon, 1925a #2187}. Note: This discovery indicated a need to involve an inflammatory reaction at the antigen‐injection site to enhance the immune response.

Jules T. Freund (AT-HU-US), Jordi Casals-Ariet (ES-US), and Elizabeth Page Hosmer (US) proposed to add inactivated Mycobacterium tuberculosis to an oil-water emulsion, leading to the popular Freund’s adjuvant {Freund, 1937 #23052}.

Jules T. Freund (AT-HU-US) and Katherine McDermott (US) made a variation in oil adjuvants, which allowed them to be used with any antigen. They emulsified water in oil with the assistance of a water-miscible lanolin-like material to incorporate both dried inactivated tubercle bacilli and the target antigen in a single aggregate preparation. This was the original Freund’s complete adjuvant that provided a method for inducing antibody formation and cellular responses to substances that are weakly antigenic {Freund, 1942 #7307}.

Porter Warren Anderson, Jr. (US) and David Hamilton Smith (US), Richard B. Johnston, Jr. (US), Michael E. Pichichero (US), Richard A. Insel (US), Robert Frank Betts (US), and Ronald J. Eby (US) made the first polysaccharide-conjugate vaccines against Haemophilus influenzae to be tested in adults and infants {Anderson, 1972 #9669;Anderson, 1986 #9668}. They went on to develop a vaccine, which was a commercial success.

Louis P. Rodriques (US), John B. Robbins (US) Rachel Schneerson (US), James C. Parke, Jr. (US), Clara Bell (US), James J. Schlesselman (US), Ann Sutton (US), Zhen Wang (US), Gerald Schiffman (US), Arthur Karpas (US), and Joseph Shiloach (US) studied the antigenicity of the Haemophilus influenzae type b (Hib) polysaccharide then developed a clinically acceptable method of binding this polysaccharide to a medically useful protein, tetanus toxoid to form a conjugate vaccine. In sequential studies, their Hib-tetanus conjugate elicited protective levels in mice, rabbits, young rhesus monkeys, and then in human adults, children, and infants. Their achievement opened the door to their and others' development of conjugate vaccines for other bacterial pathogens whose surface polysaccharide could serve as a protective antigen {Rodrigues, 1971 #9670;Schneerson, 1971 #9671;Schneerson, 1986 #9672}.

Francoise Audibert (FR), Louis A. Chedid (FR), Pierre Lefrancier (FR), Jean Choay (FR), and Edgar Lederer (FR) revealed the smallest active adjuvant structure derived from the bacterial peptidoglycan, namely, the muramyl dipeptide {Audibert, 1977 #23053}.

Charles A. Janeway, Jr. (US), in light of the function of adjuvants, redefined the immune response. What used to be defined as nonspecific immunity was renamed innate immunity. Innate immunity and inflammation are more or less two names for a similar mechanism, namely, the defense of the host against infectious or sterile insults {Janeway, 1989 #23051}.


William C. Boeck (US) and Jaroslav Drbohlav (US) were the first to develop in vitro cultures of Endamoeba histolytica. They grew it in a diphasic egg slant medium they developed for isolation of intestinal flagellates {Boeck, 1925 #2303}.


Harold Robert Dew (AU) reported the histogenesis of the hydatid parasite (Taenia echinococcus) in the pig. He gave a very complete account of the earliest stages of development of the hydatid and of the tissue changes set up in the liver {Dew, 1925 #25140}.


Alfred Stock (DE) wrote about the danger of mercury vapor and concluded that mercury is a poison, which can accumulate in the tissues over a considerable period of time {Stock, 1926 #14585}.


Martha Schmidtmann (DE) by following the spread of dye in cardiac cells presented the first evidence for a direct intercellular pathway permeable to molecules {Schmidtmann, 1925 #10339}.


Walther Vogt (DE) prepared a fate map that showed where the cells on the surface of an early gastrula would be in the later embryo. That is, he determined the destiny, or fate, of the gastrula cells {Vogt, 1925 #1103;Vogt, 1929 #16479}.

Nicole Le Douarin (FR), Claude Le Lièvre (FR), and Marie-Aimée Teillet (FR) used chick-quail chimeras to track the fate of neural crest cells and found that many of them migrate to form a variety of structures {Le Douarin, 1969 #15486;Le Douarin, 1970 #15487;Le Douarin, 1973 #15485}.


Carl Ferdinand Cori (CZ -US) and Gerty Theresa Cori, née Radnitz (CZ -US) were the first to discover that tumors in the living animal show an abnormally high formation of lactic acid from glucose {Cori, 1925b #23142;Cori, 1925d #23143;Cori, 1925c #23144}. Note: Otto Heinrich Warburg (DE) had previously established that high aerobic and anaerobic glycolysis occurs in tumors in an in vitro environment.


John Thomas Scopes (US) was the defendant in Tennessee v. John Thomas Scopes, the so-called monkey trial held in Dayton, Tennessee in July 1925, in which a science teacher was arrested for teaching evolution in violation of the state laws at that time. Two-time presidential candidate William Jennings Bryan led the prosecution. Labor lawyer Clarence Darrow led the defense and goaded Bryan into declaring that humans are not mammals {Tompkins, 1965 #6488}.


George Hoyt Whipple (US), Frieda Saur Robscheit-Robbins (US), C. Arthur Elden (US), Warren M. Sperry (US), and George B. Walden (US) found that liver, heart, skeletal muscle, ash of liver, ash of kidney, and ash of pineapple promotes regeneration of hemoglobin following severe anemia {Whipple, 1925 #10566;Robscheit-Robbins, 1925 #10567;Robscheit-Robbins, 1927 #3060;Robscheit-Robbins, 1928 #10568;Robscheit-Robbins, 1929 #3061;Whipple, 1930 #10569}. These results suggested that iron in the diet is important for hemoglobin formation.


Thomas Benton Cooley (US) and Pearl Lee (US) reported on two Italian children with symptoms of severe anemia, enlarged spleens and livers, discolored skin and bone alterations {Cooley, 1925 #12164}. Thalassaemia or thalassemia

George Hoyt Whipple (US) and William Leslie Bradford (US) coined the word thalassaemia deriving it from two Greek words - Thalassa meaning the sea and haima meaning blood, literally "sea water in the blood" {Whipple, 1936 #12165}.


Gustav Riehl (DE) described a therapy for deep burn wounds {Riehl, 1925 #11765}.


Arnold Rice Rich (US) concluded that hemoglobin, derived from destroyed erythrocytes, is the sole source of bile pigment; its normal site of origin being in reticuloendothelial cells alone, especially the Kupffer cells, and the epithelial liver cells having no role in the formation, but only in the excretion of the bile pigment {Rich, 1925 #10103}.


Sanford M. Rosenthal (CA) and Edwin C. White (CA) introduced the bromsulphalein test for hepatic function with its attendant superiority over other tests for liver function {Rosenthal, 1925 #26577}.


Percival Bailey (US) and Harvey Williams Cushing (US) postulated that medulloblastomas (Bailey coined this name) are derived from embryonic undifferentiated cells in the ependymal lining of the fourth ventricle. They described the characteristic gross and microscopic structure of this tumor, its reddish-gray color, extremely cellular nature, numerous mitotic figures, and the formation of pseudo rosettes. The tendency of this tumor to spread to the leptomeninges, which proved valuable in terms of treatment considerations, was noted. They recommended radiotherapy in the post-operative care of patients with medulloblastomas {Bailey, 1925 #16880;Bailey, 1948 #16881}.


Walter Edward Dandy (US) reported a case in which he totally removed an acoustic neuroma. He used a unilateral sub-occipital approach during which, following gutting of the tumor, he gently drew the capsule away from the brainstem {Dandy, 1925 #12215}. Dandy's success was due in part to his innovation called ventriculography, which involved x-rays and injecting a gas into the brain's cerebral ventricles for visualizing the tumor. See, Dandy 1918.


Percy Sargent (GB) was the first to remove a suprarenal tumor thus reversing virilism in the patient. Gordon Morgan Holmes (GB) described the operation {Holmes, 1925 #13321}.


Henry Sessions Souttar (GB) performed a "digital" mitral commissurotomy (closed mitral valvulotomy) {Souttar, 1925 #7905}.


Geoffrey Jefferson (GB) performed the first successful embolectomy in Britain {Jefferson, 1925 #14311}.


Konstantine Michaelovich Bykov (RU) and Alexei Dmitriev Speranski (RU) were the first to demonstrate experimentally that the bilateral synergic activity of the hemispheres of the brain may be dependent on the corpus callosum and indicated the part the corpus callosum plays in the development of symmetrical reproduction of function in the hemispheres. He severed the corpus callosum in dogs and then used Ivan Pavlov's classical conditioning methods to assess the effects of his surgeries. His experiments showed the importance of the corpus callosum for interhemispheric communication {Bykov, 1924-1925 #6613;Bykov, 1924 #6614}.

Howard J. Curtis (US) and Archibald Philip Bard (US) used electrical stimulation to prove the point-to-point connection of one side of the cortex to the other by way of the corpus callosum {Curtis, 1939 #6615}.

Andreas Vesalius (Flemish) gave the corpus callosum its name {Vesalius, 1543 #1044}.


Howard Christian Naffziger (US) described the phenomenon of the pineal shift on skull x-rays, an important sign for brain tumor localization {Naffziger, 1925 #18232}.


John Silas Lundy (US), in 1925, developed the concept of balanced anesthesia. He introduced the idea of combining more than one anesthetic technique—for example, using morphine preoperatively, a regional block to the surgical site, alongside an inhalational anesthetic of ethylene {Lundy, 1926 #16853}.


Hermann Ludwig Blumgart (US) and Otto Christian Yens (US), in 1925, performed one of the most famous of all radiotracer experiments on humans, the well-known study of arm-to-arm circulation time {Blumgart, 1927 #20340}. This ushered in the field of nuclear medicine.


Marius Nygaard Smith-Petersen (NO-US) introduced the three-flanged steel nail for insertion across the fracture site in hip fractures, an innovation that considerably improved recovery and mortality rates from hip fractures {Smith-Petersen, 1936 #25317}.


Wolfgang Köhler (DE) showed that instead of learning simply by trial and error chimpanzees can solve problems by grasping the relations between means and ends, i.e., they displayed insight {Köhler, 1925 #16568}.


Merritt Lyndon Fernald (US) proved that in the northeastern United States some species of plants and animals persisted in areas that escaped glaciation during the Pleistocene {Fernald, 1925 #20630}.


Fritz Berckhemer (DE) found a fossil skull of a young female in the Sigrist gravel pit north of Stuttgart, Germany and gave it to Karl Sigrist {Berckhemer, 1925 #17130;Berckhemer, 1933 #17131}. This, so called, Steinheim skull can be considered a Homo erectus/Homo sapiens transitional form from the Middle Pleistocene, c. 1.3 M BP.


Raymond Arthur Dart (AU-ZA) found in material from a limestone quarry at Taung (place of the lion), South Africa a fossil cast of the inside of a primate skull, which fitted into another lump of stone which possibly contained a face. It took Dart about a month to remove enough stone to reveal the face and jaw of a young fossil primate, which would be nicknamed the Taung baby. Dart considered the fossil “an extinct race of apes intermediate between living anthropoids and man.” He described it and named it Australopithecus africanus (Southern ape from Africa) and dated it to between 3 and 2.3 M {Dart, 1925 #6487}. Note: It was placed in the early Pleistocene or late Pliocene. This finding revolutionized ideas about early human evolution after human ancestors and apes split on the evolutionary tree.

Australopithecus africanus existed between 3 and 2 million years ago. It is like A. afarensis, and was also bipedal, but body size was slightly greater. Brain size may also have been slightly larger, ranging between 420 and 500 cc. This is a little larger than chimp brains (despite a similar body size), but still not advanced in the areas necessary for speech. The back teeth were a little bigger than in A. afarensis. Although the teeth and jaws of A. africanus are much larger than those of humans, they are far more like human teeth than to those of apes. The shape of the jaw is now fully parabolic, like that of humans, and the size of the canine teeth is further reduced compared to A. afarensis {Johanson, 1981 #7990;Coffing, 1994 #8748}.


The American Type Culture Collection was founded. It is a repository and source of known strains of viruses, bacteria, fungi, algae, and eukaryotic cells.


Archivos de Neurobiologia, Psichologia, Fisiologia, Histologia, Neurologia y Psiquiatria, now titled Archivos de Neurobiologia was founded.



“Men fear thought as they fear nothing else on earth—more than death. Thought is subversive, and revolutionary, destructive and terrible; thought is merciless to privilege, established institutions, and comfortable habits; thought is anarchic and lawless, indifferent to authority, careless to the well-tried wisdom of the ages. Thought looks into the pit of hell and is not afraid.…Thought is great and swift and free, the light of the world, and the chief glory of man.” Bertrand Russell {Russell, 1926 #6727}.


“Life is bottled sunshine” William Winwood Reade {Reade, 1926 #18984}.


“The striking similarity established by Meyerhof between the changes of carbohydrates in muscle and in the yeast cell is seen to be much closer than has been believed. The remarkable phenomena accompanying alcoholic fermentation are now duplicated in the case of lactic acid production, and it may reasonably be expected that most of the fermentative decompositions of sugars will be found to be initiated in a similar manner.” Arthur Harden {Harden, 1926 #9136}.


“The characters of the individual are referable to paired elements (genes) in the germinal matter that are held together in a definite number of linkage groups…. The members of each pair of genes separate when germ cells mature…. Each germ-cell comes to contain only one set…. These principles…enable us to handle problems of genetics in a strictly numerical basis and allow us to predict…what will occur…. In these respects, the theory [of the gene] fulfills the requirements of a scientific theory in the fullest sense.… It is difficult to resist the fascinating assumption that the gene is constant because it represents an organic chemical entity. This is the simplest assumption that one can make at present, and since this view is consistent with all that is known about the stability of the gene it seems, at least, a good working hypothesis.” Thomas Hunt Morgan {Morgan, 1926 #612}.


Harland W. Mossman (US) defined the placenta as "apposition or fusion of the fetal membranes to the uterine mucosa for physiological exchange." {Mossman, 1926 #26860}


Theodor Svedberg (SE) was awarded the Nobel Prize in Chemistry for his work on disperse systems.


Johannes Andreas Grib Fibiger (DK) was awarded the Nobel Prize in Physiology or Medicine for research indicating that a nematode, Spiroptera carcinoma, caused gastric cancer in rats {Fibiger, 1914 #16373}. The hypothesis was later rejected but, nonetheless, had an important role in the development of experimental research on cancer.


Gilbert Newton Lewis (US) coined the term photon for the smallest unit of radiant energy {Lewis, 1926 #16842}.


The Commission on Units and Measurements defined the röntgen at the Second International Congress of Radiology in Stockholm, Sweden. It was based on ionization of air.


Walter Norman Haworth (GB), William Charlton (GB), Stanley Peat (GB), John Vaughan Loach (GB), John Herbert Geoffrey Plant (GB), and Charles William Long (GB) had, by 1928, evolved and confirmed, among others, the structures of maltose, cellobiose, lactose, gentiobiose, melibiose, gentianose, raffinose and the glucoside ring structure of normal sugars {Charlton, 1926 #12207;Haworth, 1927a #12206;Haworth, 1927b #12208}.


Leopold Stefan Ruzicka (HR-CH) analyzed civetone and muscone, two active compounds in natural musk perfumes and found that they consist of very large rings of atoms. He showed that civetone is composed of a 17-membered, and muscone a 15-membered carbon ring. At this time, it was thought that rings with over 6 members were too unstable to exist for long {Ruzicka, 1926 #9144}. This paper was submitted in 1924.


Karl Lohmann (DE) and Lorand Jendrassik (HU) developed a colorimetric determination of phosphoric acid in muscle extracts {Lohmann, 1926 #9324}.


Francis Howard Carr (GB) and Ernest Arthur Price (GB) devised a test whereby color observed in samples of oils is a direct measure of vitamin A activity {Carr, 1926 #23364}.


Hans Karl August Simon von Euler-Chelpin (DE-SE), Christian Barthel (LU), and Karl David Reinhold Myrbäck (SE) found that dried yeast possesses practically the same power of fermentation as fresh yeast although they retain only 1% of the reproductive power {Barthel, 1929 #9134;von Euler-Chelpin, 1926 #9133}.


Julio Caesar Tello (PE) was inspired by the discovery in 1910 of the Paracas Textile (c. 100 BCE) at the site of Cabeza Larga on the Paracas Peninsula on the South coast of Peru. In his 1925 excavations Tello found coca leaves placed in an urn, which accompanied the burial of a Peruvian priest, noble, or king from the Nazca period. This is the earliest record of the use of cocoa leaves {Tello, 1926 #12549}.


Albert Jan Kluyver (NL) and Hendrick Jean Louis Donker (NL) wrote their treatise on the unity of biochemistry in which they stressed that biochemical mechanisms can be investigated using mutant strains of microorganisms and that hydrogen transfer is a basic feature of all metabolic processes {Kluyver, 1926 #99}.


Juda Hirsch Quastel (GB-CA) and Barnet Woolf (GB) published the first reported measurement of the equilibrium constant of an enzyme-catalyzed reaction. It was on the aspartate ammonia-lyase reaction {Quastel, 1926b #16112}.


James Batcheller Sumner (US) was the first to obtain an enzyme in pure crystalline form. This was the enzyme urease, which catalyzes the hydrolysis of urea to yield carbon dioxide and ammonia. Sumner had used a 32% solution of acetone in water to precipitate much of the organic material in a jack-bean extract rich in urease activity, while leaving most of the urease activity in solution. After filtering off this precipitate and letting the filtrate stand overnight in the cold, he found that crystals of protein had formed in the filtrate. Study of these protein crystals revealed that they are pure urease enzyme endowed with a specific enzymatic activity enormously greater than that of the original jack-bean extract. It was thus proven that the enzyme urease is a protein and that a polypeptide chain can act as a catalyst in the facilitation of a chemical reaction.

Sumner’s proof was an important milestone along the road to understanding the chemical basis of cell function {Sumner, 1926 #24263}.


John Jacob Abel (US) crystallized insulin {Abel, 1926 #12171}.


Anna L. Sommer (US) and Charles B. Lipman (US) experimentally demonstrated the essential nature of zinc, copper, and boron for plant growth {Sommer, 1926 #18503;Sommer, 1931 #18504}.


Alfred Henry Sturtevant (US) found that inversion of a section of the third chromosome in Drosophila melanogaster reduces crossover frequency {Sturtevant, 1926 #12353}.


Hugo W. Alberts (US) devised a method for calculating genetic linkage values {Alberts, 1926 #22108}.

Calvin Blackman Bridges (US) and Tomasz M. Olbrycht (PL) noted that to accurately construct genetic maps it is necessary to know the full number of crossovers (doubles counting as two crossovers, triples as three crossovers) that occur between the loci that are to be mapped. In constructing such maps, it is first necessary to determine experimentally the amounts by which the various crossing over values exceed the directly observed recombination percents. The experimental determination of the amount by which crossing over exceeds recombination is made through use of loci lying between the two loci in question. Bridges created the sc ec ct v g f multiple recessive and called it "Xple" (X-chromosome multiple). This organism was used to test many aspects of linkage mapping {Bridges, 1926 #22118}.


Edgar Altenburg (US) and Hermann J. Muller (US) demonstrated the nature of the mechanism of inheritance of the truncate character in Drosophila. They showed that it conformed regularly to the principles of chromosome heredity, factor constancy, etc., and disclosed the causes of the inconstancy of the inbred stock, and of the indefiniteness and variability of the ratios thrown in crosses. The truncate character, when it was analyzed, was found to depend on so many factors. 1) Two successive mutations were practically necessary, in the first place, before truncate became visible at all. 2) Since truncate depended on a lethal, and was, in addition, inconstant in somatic expression, it was subjected to selection, which perpetuated any new factors (balancing lethals and intensifiers) that still further differentiated truncate from normal. 3) Since the truncate character depended on an unstable developmental reaction, the initial appearance of such intensifiers was made more likely {Altenburg, 1920 #22109}.


Edgar Douglas Adrian (GB) and Yngve Zotterman (SE), using the afferent nerve of a stretch receptor in a frog muscle, showed that it is possible by amplification to record the impulses in single nerve cells to a natural stimulus (tension). They postulated the concept of adaptation of receptors to stimuli and predicted that other units of the nervous system would also exhibit adaptation {Adrian, 1926 #3286}.

Edgar Douglas Adrian (GB) and Yngve Zotterman (SE), established beyond doubt that the nerve impulse is invariant, that the intensity of sensation is conveyed by the frequency of impulses and the quality by the type of nerve fiber in action {Adrian, 1926 #3286}.


Eugene Markley Landis (US), Schack August Steenberg Krogh (DK), and Abbey H. Turner (US), in an elegant series of experiments, determined that the rate of net fluid movement across the capillary wall is proportional to the difference between capillary hydrostatic pressure and the osmotic pressure of the plasma proteins, thus providing the first experimental proof of Ernest Starling’s hypothesis of fluid exchange. Landis’s constant of proportionality was the first quantitative measurement of the hydrodynamic conductance of the capillary wall (cubic microns of fluid per second per centimeter water pressure difference per square micrometer of capillary wall). These papers included the first measurements of the pressure drops along the vascular tree and localization of the separate components of the peripheral resistance to blood flow, in mammals as well as frogs {Landis, 1926a #3233;Landis, 1926b #10615;Landis, 1927a #10617;Landis, 1927b #10618;Landis, 1927c #10620;Landis, 1928 #10621;Landis, 1930a #10622;Landis, 1930b #10623;Landis, 1930c #10624;Landis, 1931 #10625;Krogh, 1932 #10626;Landis, 1933a #10627;Landis, 1933b #10628;Landis, 1934 #10629}. See, Starling 1896.


John Belling (GB-US) perfected the aceto-carmine staining method by adding iron. This technique was especially important because it allowed a clear differentiation between chromosomes and cytoplasm {Belling, 1926 #1204}.


Walter Bradford Cannon (US) coined the word homeostasis: the maintenance of balance in the body’s state via chemical feedback mechanisms {Cannon, 1926 #3343;Cannon, 1932 #1435}. See, Claude Bernard (FR) for his concept called milieu intérieur, 1865. See, Alcmaèon (GR) for his concept called isonomy, 520 BCE


Gaston Ramon (FR) and Pierre A. Descombey (FR) described the flocculation reaction and observed that flocculation may occur in zones that do not correspond to the point at which the toxin and antitoxin neutralize each other {Ramon, 1926a #7078}.


Kenjiró Fujii (JP) observed the coiled structure of the chromosome for the first time noting that in certain stages of the cell cycle, two filaments appeared to be coiled around one another {Fujii, 1926 #19062}.


Frantisek Vejdovsky (CZ) showed that nuclear fission in an ovule is preceded by the splitting of the centrosome (which he called the periplast); he was apparently the first to observe the centriole in 1866 {Vejdovsky, 1926-1927 #7389}.


Félix Hubert d’Herelle (CA) described the three-step process for the life history of the bacteriophage virus: (1) attachment to the susceptible bacterium, (2) multiplication in the cell, and (3) disintegration of the cell to set free the progeny virus particles and attachment of the progeny to other susceptible bacteria, if such are present. These conclusions were based on the plaque-count, and dilution methods of assay that he had invented {d'Herelle, 1926 #101}.


Louis Otto Kunkel (US) demonstrated that Cicadula sexnotata (a leafhopper) could act as a vector of aster yellows virus {Kunkel, 1926 #1612}.


Thomas Clifford Vanterpool (CA) was the first to discover that mixed infections, the simultaneous occurrence of two viruses within a host plant acting together, could produce a distinct severe disease. He showed that tomato mosaic virus and potato mosaic virus acting together caused the severe disease of tomatoes called streak or winter blight while either virus acting alone was shown to cause mild symptoms {Vanterpool, 1926 #5718}.


Andre Paillot (FR) discovered granulosis virus (GV) infection in an insect, Pieris brassicae (cabbage butterfly). He was the first to describe a new group of diseases, the granuloses, which are characterized by the formation of virus inclusion bodies called granules within tissues of infected insects {Paillot, 1926 #5719}. These are now considered Baculovirus.


Frederik Kraneveld (NL) and Thomas M. Doyle (GB) discovered Newcastle disease virus {Doyle, 1927 #25963;Kraneveld, 1926 #25964}. Note: first paramyxovirus


William Edwin Cotton (US), Peter K. Olitsky (US), Jacob Traum (US), and Harry W. Schoening (US) discovered vesicular stomatitis viruses {Olitsky, 1926 #25965;Cotton, 1926 #25966}.


Patrick Playfair Laidlaw (GB) and George William Dunkin (GB) established the viral etiology of canine distemper {Laidlaw, 1926 #25988}.


Edson Sunderland Bastin (US) succeeded in culturing sulfate-reducing bacteria from groundwater samples extracted from an oil deposit that was hundreds of meters below the surface. Since this discovery over 9,000 strains of bacteria and fungi have been isolated from diverse subsurface environments {Bastin, 1926 #12172;Fredrickson, 1996 #681}.


Heinrich Bernward Prell (DE) reported an amoebic infection of an insect, Apis mellifera Linn. (honeybee). The etiological agent he described and named Malpighamoeba mellificae attacks the Malpighian tubules causing a dysentery {Prell, 1926a #16843;Prell, 1926b #7150}.


Walter H. Burkholder (US) observed that halo blight of beans is caused by Pseudomonas phaseolicola {Burkholder, 1926 #102}.


Everitt George Dunne Murray (CA), Robert Alexander Webb (GB), and Meredith Blake Robson Swann (GB) isolated and characterized Listeria monocytogenes as the etiological agent of an epizootic among laboratory rabbits and guinea pigs, which was characterized in part by monocytosis {Murray, 1926 #2248}.

Walter F. Schlech III (US), Pierre M. Lavigne (US), Robert A. Bortolussi (US), Alexander C. Allen (US), E. Vanora Haldane (US), A. John Wort (US), Allen W. Hightower (US), Scott E. Johnson (US), Stanley H. King (US), Eric S. Nicholls (US), and Clare V. Broome (US) established that human consumption of Listeria-contaminated foodstuffs causes a disease called listeriosis {Schlech, 1983 #8238}.


Louis Edmond den Dooren de Jong (NL) demonstrated that a strain of Pseudomonas putida can proliferate on a mineral medium to which any one of some eighty compounds was added as the sole organic substance available {de Jong, 1926 #12174}.


Maurice Lemoigne (FR) originally discovered poly-beta-hydroxybutyric acid (PHB) as a major component of the cells of the bacterium Bacillus megaterium {Lemoigne, 1926 #12178}.

Frederick P. Delafield (US), Michael Doudoroff (US), Norberto J. Palleroni (US), Carol Jean Lusty (US) and Rebecca Contopoulos (US) later honored Lemoigne by naming Pseudomonas lemoigne, an active oxidizer of extracellular PHB, for him {Delafield, 1965 #12179}.


Frank W. Tilley (US) and Jacob M. Schaffer (US) determined that the germicidal activity of the aliphatic alcohols increases regularly from methyl through octyl alcohol for each additional methyl group in the straight chain {Tilley, 1926 #8015;Schaffer, 1927 #12173}.


Samuel Ottmar Mast (US) proposed the tail contraction model to explain sol-gel amoeboid movement {Mast, 1926 #12166}.


John Nathaniel Couch (US) described for the first time the existence of physiologically distinct and separate male and female strains in an oomycete (Dictyuchus) {Couch, 1926 #10677}.


Friedrich Seidel (DE), using eggs of the dragonfly, Platycnemis pennipes, determined that embryonic development in insects is typified by the presence of a special kind of germ-band formation. Because of the distribution of cytoplasm and yolk in the egg the germ band is limited to a particular region. The ventral portion of the developing embryo is special because it is the carrier of the main system of organs. The ventral part of the embryo precedes the dorsal side in development {Seidel, 1926 #7228;Seidel, 1928 #7229;Seidel, 1971a #7227}.


Alden B. Dawson (US) carried out the first successful skeletal staining when he used alizarin red S {Dawson, 1926 #9323}.


Henry Allan Gleason (US) argued that every plant association is the unique product of the fluctuating environmental conditions of a particular time and place {Gleason, 1926 #8306}.


Gavin Rylands de Beer (GB) observed that certain cartilage and bone cells are derived from the outer ectodermal layer of the embryo; calling into question the germ-layer theory {de Beer, 1926 #8970}.


Lloyd R. Watson (US) devised instrumental methods to artificially inseminate the honeybee {Cale, 1926 #1890;Watson, 1927 #1891;Watson, 1928 #1892}. This instrument greatly improved genetics of the honeybee.


Sergei Sergeevich Chetverikov (RU) concluded that populations in nature maintain within themselves the variants, which arise within them by mutation. This would provide them with a supply of potential but hidden variability out of which the adaptiveness of the population to a changing environment could arise {Chetverikov, 1926 #5044;Chetverikov, 1961 #12175}. Note: Together with Ronald Aylmer Fisher, John Burdon Sanderson Haldane, and Sewall Green Wright, Chetverikov is now regarded as one of the founders of population genetics and modern evolutionary theory.


William Bloom (US) worked out the early stages of the embryogenesis of human bile capillaries {Bloom, 1926a #10667}.


William Bloom (US) described the transformation of small lymphocytes into myelocytes in germinal centers {Bloom, 1926b #10668}.


George Ellett Coghill (US), in his studies of the amphibian Amblystoma punctatum, established that innervation develops in a cephalo-caudal (head-tail) direction, and that limb movements emerge from a more general pattern of trunk movement. He then theorized that behavior develops as the expansion of a "total pattern," rather than simply as the combination or coordination of reflexes and suggested strongly that this might well be true for higher vertebrates, including man {Coghill, 1926 #12189;Coghill, 1929 #12188;Coghill, 1930 #12190;Coghill, 1931 #12191}.


Serguei Metalnikov (RU-FR) and Victor Chorine (FR) provided an important paper in psychoneuroimmunology when they adapted Pavlov’s procedures of stimulant conditioning to activate and enhance cellular and antibody immune responses to foreign substances, particularly to otherwise lethal doses of cholera and anthrax bacteria {Metalnikov, 1926 #7313}.


Ralph Milton Waters (US) authored a landmark article on carbon dioxide (CO2) absorption in anesthesiology {Waters, 1926 #12303}.


Karl Ernest Mason (US), working with the rat, was the first to associate testicular degeneration and hence male infertility with vitamin E deficiency {Mason, 1926 #23428}.

Harold S. Olcott (US) and Henry A. Mattill (US) fractionated the oil of lettuce and found that one fraction had high antioxidant power and high potency as a source of vitamin E {Olcott, 1931 #3069}.

Herbert McLean Evans (US), Oliver H. Emerson (US), and Gladys A. Emerson (US) isolated alpha-tocopherol from wheat germ oil and discovered that it is synonymous with vitamin E {Evans, 1936 #3068}.

Erhard Robert Fernholz (US) determined the structure of alpha-tocopherol (vitamin E) {Fernholz, 1938 #13690}.

Paul Karrer (RU-CH), Hans Heinrich Fritzsche (CH), Beat Heinrich Ringier (CH), and H. Salomon (CH) synthesized alpha-tocopherol (vitamin E) and proved its biological role as a vitamin {Karrer, 1938 #3070}.


Wells Phillips Eagleton (US) described cavernous sinus thrombophlebitis in relation to septicemia {Eagleton, 1926 #22634}.


Giovanni Di Guglielmo (IT) described a syndrome of unknown origin characterized by enormous numbers of nucleated erythrocytes appearing in the bone marrow and blood (acute erythroblastosis) {Di Guglielmo, 1926 #13294}.


Erik Adolf Willebrand (FI) described a previously unknown form of hemophilia with a prolonged bleeding time course as its most prominent sign. He named it pseudo-hemophilia {Willebrand, 1926 #13571;von Willebrand, 1931 #13572;Willebrand, 1939 #13573}. Note: Later called Willebrand’s disease I


Nikolai Mikhailovich Itsenko (RU) described glucocorticoid excess syndrome in which the hypersecretion of glucocorticoids is secondary to hypersecretion of adrenocorticotrophic hormone from the pituitary. It may also be caused by a basophilic adenoma of the pituitary {Itsenko, 1926 #13338}.

Harvey Williams Cushing (US) described this syndrome six years later {Cushing, 1932b #11778}. It is often called Cushing’s syndrome I but should be called Itsenko-Cushing syndrome.


Felix Mandl (AT) was the first to undertake parathyroid surgery, successfully removing a parathyroid adenoma in a patient with osteitis fibrosa cystica {Mandl, 1926 #12751}.


Henry Head (GB) presented his theory of aphasia as a condition with, “a disorder of symbolic formulation and expression” {Head, 1926 #13306}.


César Roux (CH) performed the first surgical resection of a pheochromocytoma, in 1926 {Manger, 1977 #13827}. Later the same year, Charles Horace Mayo (US) performed the first surgical resection in the United States and described medical and surgical management of pheochromocytoma {Mayo, 1927 #11548}.


Percival Bailey (US) and Harvey Williams Cushing (US) wrote a book, which formed the basis of modern day neurooncology. It completely revolutionized the understanding of neurooncology, and for first time the neurosurgical community was presented with an orderly classification of gliomas based on the tumor's natural history and clinical course. This work changed antiquated thinking by showing that the microscopic structure of a tumor is important for prognosis. It completely revamped the understanding of these tumors. In fact, the histopathological basis of brain tumors in relation to patient survival rate and outcome still influences present-day neurosurgical thought {Bailey, 1926 #16877}.

Percival Bailey (US) simplified, refined, and made the concepts presented in the 1926 book more practical {Bailey, 1927a #16878;Bailey, 1927b #16879}.


Harvey Williams Cushing (US) and William T. Bovie (US) conceived and introduced electrosurgery, which allowed the cutting of tissue with almost no bleeding. During 1927, Cushing removed a number of brain tumors previously considered inoperable {Cushing, 1928a #12094;Riedman, 1962 #7656}.

Maximilian Carl-Friedrich Nitze (DE), in 1896, developed an operating cystoscope fitted with an electric cautery {Nitze, 1907 #16845}.


Francis Bertody Sumner (US) made a thorough study of coat color among the mainland and Santa Rosa Island mice and concluded that both isolation and natural selection probably operated in the origin of species, but that environmental mechanisms could not be ruled out. Though still not conclusive, Sumner’s reading of nature’s experiments on Santa Rosa came closer than any other study at the time — field or lab — to providing empirical evidence of the mechanism of evolution. Santa Rosa is an island off the Florida panhandle {Sumner, 1926 #24263}.


Knud Haraldsen Krabbe (DK) founded the journal Acta Psychiatrica et Neurologica Scandinavica.



Heinrich Otto Wieland (DE) was awarded the Nobel Prize in Chemistry for his investigations of the constitution of the bile acids and related substances.


Julius Wagner-Jauregg (AT) was awarded the Nobel Prize in Physiology or Medicine for his discovery of fever therapy for paresis (dementia paralytica). He shared the honor with the pathologist Johannes Andreas Grib Fibiger (DK), who was awarded the prize "for his discovery of the Spiroptery carcinoma."


Georges Henri Joseph Édouard Lemaitre (BE) proposed the Big Bang theory of the origin of the universe. He visualized a primal atom, an incredibly dense egg containing all the material for the universe within a sphere about 30 times larger than the Sun. This primal atom exploded some 20-60 Ga scattering matter and energy in all direction {Lemaitre , 1927 #23561;Lemaitre, 1946 #8990}. This theory is now known popularly as the Big Bang Theory, a phrase coined by Fred Hoyle (GB) in a moment of facetiousness, during a radio broadcast {Hoyle, 1950 #16385}. Note: Alexander Friedmann (RU) proposed an expanding universe as early as 1922 {Belenkiy, 2012 #23562}. Note: Today, most physicists and cosmologists conceive of the primal atom as having been smaller than Lemaitre's!


Alexander Logie du Toit (ZA) compared the geology of South America and South Africa and found them to be similar in many ways {du Toit, 1927 #7390}.


Hans Busch (DE) theorized that magnetic fields could act as lenses by focusing electron beams to a point {Busch, 1927 #12168}. This was vital to the invention of the electron microscope.


Hermann Joseph Muller, Jr. (US) and Lewis John Stadler (US) discovered that x-rays induce mutations in animals and plants respectively. They found that the dose-frequency curve is linear {Muller, 1927 #539;Stadler, 1928a #540;Stadler, 1928b #14665}. Note: here there is emplied that somatic mutations initiate cancer


Albert Charles Chibnall (GB) and Harold John Channon (GB) discovered the exact structure of phosphatidic acid; described for the first time in living materials {Chibnall, 1927 #12110}.


Otto Knut Olof Folin (SE-US) and Vintila Ciocalteu (RO) designed the Folin-Ciocalteu reagent to detect phenolic and antioxidant polyphenols {Folin, 1927 #9325}. Note: This work used the now famous Folin-Ciocalteu reagent.


Rudolph John Anderson (SE-US) isolated the various lipoid and carbohydrate fractions from the tubercle bacillus. He was able to separate the lipoid constituents of tubercle bacilli into three groups consisting of wax, glycerides, and phosphatides {Anderson, 1927 #21462}.

Rudolph John Anderson (SE-US) described the constitution of phthiocol, a fat-soluble pigment he isolated from human tubercle bacillus. He determined that this previously unknown substance was in fact 2-methyl-3-hydroxy-1, 4-naphthaquinone {Anderson, 1933 #21463}.


Otto Fritz Meyerhof (DE-US) discovered that the first step in the fermentation of glucose (glucose → glucose-6-phosphate), catalyzed by an enzyme he called hexokinase, does not require inorganic phosphate but rather organic phosphate transferred from its terminal, or g position on adenosine triphosphate (ATP) {Meyerhof, 1927 #8347}.

Otto Fritz Meyerhof (DE-US) and Fritz Albert Lipmann (DE-US) discovered that sodium fluoride strongly inhibits the fermentation and phosphorylation of hexoses {Meyerhof, 1927 #8347;Lipmann, 1928 #8346}.

Hans Karl August Simon von Euler-Chelpin (DE-SE), Erich Adler (), Otto Fritz Meyerhof (DE-US), Sidney P. Colowick (US), and Herman Moritz Kalckar (DK-US) partially purified hexokinase then established the reaction which it catalyzes as: glucose + adenosine triphosphate (ATP) à glucose 6-phosphate + adenosine diphosphate (ADP) {Colowick, 1943 #11012;Meyerhof, 1935e #20835;von Euler-Chelpin, 1935d #20836}


Hans Karl August Simon von Euler-Chelpin (DE-SE) ws the first to show that the combination between an enzyme and a substrate can be traced back to a certain atomic group. He was experimenting with dipeptidases in which a carbonyl group is the site of combination {von Euler-Chelpin, 1927a #9132;von Euler-Chelpin, 1927b #23232}.


Juda Hirsch Quastel (GB-CA) and Walter Reginald Wooldridge (GB) studied dehydrogenases from Escherichia coli and further developed the concept of the active center, or site of activation, a term earlier coined by Quastel (1926). This work stressed the importance of using in vitro results to understand in vivo activities of enzymes {Quastel, 1927 #5547}. They also discovered that malonic acid is a powerful inhibitor of succinic dehydrogenase {Quastel, 1928 #5548}.


Philip Eggleton (GB) and Marion Grace Palmer Eggleton (GB) found an organic acid-labile phosphate in muscle tissue. They named it phosphagen (phosphocreatine) and established the fact that muscular contraction is accompanied by removal of phosphagen, and subsequent recovery in oxygen is characterized by a rapid restitution of the phosphagen—a phase of recovery apparently independent of the relatively slow oxidative removal of lactic acid {Eggleton, 1927 #3045}.

Cyrus Hartwell Fiske (US) and Yellapragada SubbaRow (IN-US) discovered the chemical nature of the phosphagen (phosphocreatine) present in muscles. They announced that voluntary muscle contains an unstable compound of creatine and phosphoric acid, which is hydrolyzed by stimulation, and resynthesized when the muscle is permitted to recover. They proposed a structure and pointed out some of the physiological properties of phosphocreatine (later changed to creatine phosphate) {Fiske, 1927 #1729;Fiske, 1928 #1730;Fiske, 1929b #1731}. Note: Michel-Eugène Chevreul (FR), in 1832, determined that muscle contains creatine {Chevreul, 1835 #19064}.


Bernhard Zondek (DE-IL) and Selmar Aschheim (DE) isolated estrogenic hormone from the urine of pregnant women in amounts averaging 12,000 mouse units per liter {Zondek, 1927 #3385}. They found that the urine of pregnant women, when injected into female animals, caused hyperemia of the ovaries, growth of the follicles, and, in some species ovulation {Aschheim, 1928b #13064}. This became the basis of the Ascheim-Zondek and Friedman-Lapham Tests for pregnancy.

Maurice Harold Friedman (US) and Maxwell Edward Lapham (US) developed the "Rabbit test" for early diagnosis of pregnancy. Two morning samples of urine were injected into the marginal ear vein of a virginal female rabbit that had been isolated from male rabbits. The result was available 48 hours after the first injection. Of all the biologic tests, Friedman's was the most accurate. This procedure tested for the presence of human chorionic gonadotropin (hCG; a hormone released from the implantation site of a blastocyst that prevents menstruation) in the urine of women. If hCG were present in the urine, the rabbit's ovaries would form corpora lutea (ovarian endocrine structures formed following ovulation) within 48 hours {Friedman, 1929 #19066;Friedman, 1931 #19065}.


Julius Moses Rogoff (US) and George Neil Stewart (US) demonstrated that a chemical extract of the adrenal glands of dogs greatly prolongs the lifespan of adrenalectomized dogs. They found that the active ingredient was not epinephrine (adrenaline) {Rogoff, 1927 #3402}. They experienced some success in using adrenal extract to treat human patients suffering from Addison’s disease {Rogoff, 1929 #3403}. Charles-Édouard Brown-Séquard (FR) had proved that removal of both suprarenal (adrenal) glands invariably proved fatal. See, Brown-Séquard, 1856.


Walter Bradford Cannon (US), Andries Querido (NL), Sydney W. Britton (US), and Elizabeth M. Bright (US) demonstrated that the adrenal glands produce more adrenaline (epinephrine) in animals exposed to the cold. The adrenaline (epinephrine) causes an increase in heat production {Cannon, 1927 #3349}.


Joseph Barcroft (GB) and John G. Stephens (GB) demonstrated the spleen's role as a blood reservoir in the dog {Barcroft, 1927 #12153}.


Thomas Milton Rivers (US) clearly distinguished between bacteria and viruses, thereby, giving rise to the field of virology {Rivers, 1927 #8809}.


Nicolaas Louis Söhngen (NL) was probably the first to report bacteria that lyse other bacteria, in some cases with a high degree of parasite-host specificity. Bacterium bacteriovorus, which lyses Bacillus danicus, he found to grow only in the presence of its host {Söhngen, 1927 #8653}.


Ronald Aylmer Fisher (GB-AU) explained the evolution of Batesian mimicry by a series of small evolutionary steps. He proposed that the phenotypic expression of genes could be modified by the action of other genes. Rare imperfect mimetic forms when they appeared in a population would vary due to the action of modifier genes. Selection would favor those modifier genes that produced an increase in the accuracy of the mimicry {Fisher, 1927 #6071}.

Cyril Astley Clarke (GB), Philip Macdonald Sheppard (GB), and Lawrence M. Cook (GB) provided experimental evidence to support Fisher’s explanation {Clarke, 1959a #6073;Clarke, 1959b #6074;Clarke, 1960a #6075;Clarke, 1960b #6076;Clarke, 1962 #6077;Clarke, 1963a #6072;Sheppard, 1962 #6078}.


John Charles Walker (US) pioneered research on genetic resistance in yellows disease of cabbage. He showed the scientific community that disease control through genetic resistance could be an effective and relatively inexpensive approach to solving plant disease problems {Walker, 1927 #11185;Walker, 1930 #11186}.


Bernard Ogilvie Dodge (US) worked out the life cycle of the pink bread mold, Neorospora crassa, an ascomycete {Dodge, 1927 #5031}. Cornelius Lott Shear (US) and Bernard Ogilvie Dodge (US) named the fungal genus Neurospora {Shear, 1927 #1687}.


Emil Bozler (DE-US) demonstrated that the nerve net of cnidarians is made up of separate cells connected by synaptic junctions. He also studied electrical aspects of muscle contraction and the role of calcium and magnesium in contraction and relaxation {Bozler, 1927 #12187}.


Nadine Dobrovolskaia-Zavadskaia (FR) and Nicolas Kobozieff (BE) identified t-haplotypes in mice because they contain the gene tct (t-complex tail interaction factor) that interacts with a spontaneous dominant mutation Brachyury (T). Brachyury produces short tails in T/+ mice but interacts with tct to produce a tailless phenotype in T/t mice {Dobrovolskaia-Zavadskaia, 1927 #16303;Dobrovolskaia-Zavadskaia, 1932 #16304}.

James R. Archer (GB), Steven J. Self (GB), and Bryan G. Winchester (GB) explained the t-complex in mice as a genetic entity which alters meiosis is such a way that its transmission into gametes is favored {Archer, 1978 #16302}. The t-complex is located on the proximal third of chromosome 17 in the house mouse. Naturally occurring variant forms of the t-complex, known as complete t-haplotypes, are found in wild mouse populations. The t-haplotypes contain at least four nonoverlapping inversions that suppress recombination with the wild-type chromosome, and lock into strong linkage disequilibrium.

Barbara C. Turner (US) and David D. Perkins (US), working with Neurospora, discovered spore killer, a chromosomal factor that kills meiotic products in which it is not contained {Turner, 1979 #16300}.

Larry Sandler (US) and Kent G. Golic (US) discovered segregation distorter (Sd), a meiotic drive system that operates in males of Drosophila melanogaster. Males heterozygous for Sd, a dominant neomorphic (gain-of-function) mutation on chromosome 2, can transmit their progeny in greater frequencies than expected on a Mendelian basis {Sandler, 1985 #16301}.


Evgenii Nikanorovich Pavlovsky (RU) outlined his theory of landscape epidemiology and initiated modern concepts of parasite epidemiology {Pavlovsky, 1966 #6564}.


Jesse Amos (GB), Ronald George Hatton (GB), R.C. Knight (GB), and Arthur Morel Massee (GB) suggested that eriophyid mites (gall mites) could have some connection with plant virus transmission {Amos, 1927 #8693}.

Arthur M. Massee (GB) was the first to demonstrate this to be the case {Massee, 1952 #8694}.


Erwin Stresemann (DE) wrote an important volume on bird biology as part of Kükenthal and Krumback’s Handbuch der Zoologie. It contains thorough discussions of avian physiology, anatomy, and other phases of avian biology {Stresemann, 1927-1934 #5198}.


Theophilus Shickel Painter (US) studied the Japanese Waltzer mouse and concluded that its phenotype results from a chromosomal deletion {Painter, 1927 #10776} This was the first cytological identification of a deletion producing a specific genetic effect and the first case in mammals of locating a definite gene on a definite chromosome.


Thomas Lewis (GB) originally described the wheal and flare reaction. He described how he believed histamine to be crucially involved in the central area of the weal, where it then stimulates an axon reflex or antidromic reflex, which transmits the signal to more distant parts, i.e., the flare {Lewis, 1927 #12192}.


Karl Landsteiner (AT-US) and Philip Levine (RU-US) characterized the M, N, P and Jay blood antigens {Landsteiner, 1927a #14664;Landsteiner, 1927b #7095;Landsteiner, 1928 #1679;Landsteiner, 1931 #1622}.


Florence R. Sabin (US) and Charles A. Doan (US) showed that bovine tubercle bacilli were able to stimulate the maturation of monocytes and were destroyed by clasmatocytes (histiocytes of the connective tissues) {Sabin, 1927 #23054}.


Matthew Walzer (US) and Sampson J. Wilson (US) demonstrated that the ingestion of foods would allow food antigens to penetrate the gastrointestinal barrier, which are then transported in the circulation to IgE-bearing mast cells in the skin {Walzer, 1927 #12226;Wilson, 1935 #12227}. The IgE class of immunoglobulin was not discovered until 1967.


Willem Storm van Leeuwen (NL) and Hermann Dekker (DE) suggested the possible significance of mites in house dust allergy {Dekker, 1928 #21047;van Leeuwen, 1927 #21052;van Leeuwen, 1929 #21053}.

Reindert Voorhorst (NL), Frederik Theodorus Maria Spieksma (NL), Marise I.M. Spieksma-Boezeman (NL), Hendrik Varekamp (NL), Maarten J. Leupen (NL), and Ankie W. Lyklema (NL) reported that house dust contains mites of the genus Dermatophagoides and suggested that these were the major source of allergens in house dust {Spieksma, 1967a #21051;Spieksma, 1967b #21050;Voorhorst, 1964 #21048;Voorhorst, 1967 #21049}.


David Marine (US) and Emil J. Baumann (US) found that administration of Ringer’s solution and isotonic solutions of sodium chloride and sodium acetate increased the life span of suprarenalectomized cats about three-fold {Marine, 1927 #16238}.


Soma Weiss (HU-US) developed the first practical method of measuring circulation in time {Weiss, 1927 #13556}.


Wallace Osgood Fenn (US) was the first to measure the quantity of oxygen required by a nerve to conduct an impulse {Fenn, 1927 #10867}.


Otto Heinrich Warburg (DE), Harry Goldblatt (US), Gladys Cameron (US), Fritz Kubowitz (DE), Karlfried Gawehn (DE), August-Wilhelm Geissler (DE), Detlev Kayser (DE) and Siegfried Lorenz (DE) provided the very unexpected and fundamental fact, that tissue culture is carcinogenic and that a too low oxygen pressure is the intrinsic cause. Anaerobiosis of cancer cells was an established fact by 1960 when methods were developed to measure the oxygen pressure inside of tumors in the living body {Goldblatt, 1953 #14829;Warburg, 1927b #14828;Warburg, 1956 #14825;Warburg, 1965 #14831;Warburg, 1966 #14830}.

Mark W. Woods (US), Katherine K. Sanford (US), Dean Burk (US), and Wilton R. Earle (US) found that cancer cells, descended in vitro from one single normal cell, were in vivo the more malignant, the higher their fermentation rate {Woods, 1959 #14826}.

Dean Burk (US), Mark W. Woods (US), and Jehu Hunter (US) found that when different carcinogens were used to induce in vivo hepatomas there was a direct correlation between malignancy and fermentation rate {Burk, 1967 #14827}.


Thomas Benton Cooley (US), E.R. Witwer (US), and O. Pearl Lee (US) described a disease, later known as Cooley erythroblastic anemia {Cooley, 1927 #11776}.


Guido Fanconi (CH) reported a family in which three brothers had died in childhood from a condition, which resembled pernicious anemia. He observed that these children had slight stature, hypogonadism and skin pigmentation. In subsequent reports defects of the thumb and radius were recognized as additional but variable syndromic components {Fanconi, 1927 #19309}. This became known as Fanconi’s anemia.


Leo Loeb (US) and Sewall Green Wright (US) demonstrated genetics of transplant specificity in mammals {Loeb, 1927 #14663}.


Albert Kuntz (US) described the gray ramus running from the second thoracic ganglion to the first thoracic nerve {Kuntz, 1927 #22781}.


Arthur Cecil Alport (ZA-GB) first published his description of a triad of symptoms in a family with hereditary familial congenital haemorrhagic nephritis, deafness and ocular changes (Alport’s syndrome) {Alport, 1927 #22884}.

David F. Barker (US), Sirkka Liisa Hostikka (US), Jing Zhou (US), Louise T. Chow (US), Arnold R. Oliphant (US), Steven C. Gerken (US), Martin C. Gregory (US), Mark H. Skolnick (US), Curtis L. Atkin (US), and Karl Tryggvason (US) reported that X-linked Alport syndrome is caused by mutations of the COL4A5 gene coding for a collagen chain normally found in the mature glomerular basement membrane {Barker, 1990 #25634}.


Philip Edward Smith (US) perfected the surgical production of hypophysectomized rats and described the symptoms resulting as inhibition of growth in the young animal, weight loss, atrophy of the genital system with loss of libido sexualis, cessation of the female sex cycle, atrophy of the thyroid, parathyroids and suprarenal (adrenal) cortex, lowered resistance to injury, loss of appetite, weakness, and flabbiness. Smith found that he could reverse the atrophied functions in these animals only by injection of fresh hypophyseal tissue from adults. These animals have been widely used in studies of the endocrine system {Smith, 1927 #3379}.


Alfred Washington Adson (US) and Jay R. Coffey (US) recommended the division of the scalenus anticus muscle for relief from symptoms of cervical rib syndrome {Adson, 1927 #22581;Adson, 1947 #22582;Adson, 1951 #22583}.


Manfred Joshua Sakel (PL-AT-US), in 1927, introduced hypoglycemic coma (insulin shock) produced by muscular injections of insulin as a treatment for psychotic disorders (e.g. schizophrenia) and drug addiction {Sakel, 1938 #12447;Sakel, 1994 #12449}. Note: In Berlin, between 1928 and 1931, Dr. Manfred Sakel used insulin to reduce the anxiety, nervousness, tremors, vomiting, weight loss, and agitation of patients undergoing opiate withdrawal. With insulin, they became calm, gained weight, and were much more cooperative. At times, when the dose of insulin was high, the patient went into stupor. After such events, the patients were less argumentative, less hostile, and less aggressive.

Ladislaus Joseph Meduna (HU) started seizure therapy by intravenous injection of cardiazol (in depressive states), a therapy that was abandoned in 1938 {Meduna, 1937 #12448;Meduna, 1938 #12453}.

Ugo Cerletti (IT) and Lucio Bini (IT) introduced electric convulsive therapy (E.C.T.) for severe mental states. This treatment was first used in schizophrenia, but severe depressive states very soon proved to be the main indication {Cerletti, 1938 #12452;Cerletti, 1940 #12450}. Note: people used electric eels and electric fish in ancient times to treat headaches and mental illness.

Jean Delay (FR) and Pierre Deniker (FR) demonstrated success with the first neuroleptic—coined by Delay— drug, the phenothiazine derivative chlorpromazine (Thorazine). Chlorpromazine has a remarkable effect on patients with schizophrenia, in particular highly agitated, anxious, and psychotic patients. It softens the effects of hallucinations and voices. Patients became quiet and much more manageable. It has effects on all types of patients {Delay, 1952a #12451}.


Otto Wuth (US) described the preparation of a stable colloidal gold solution for use in the Lange protein spinal fluid test. With the bromide assay, Wuth found that 21% of 238 admissions to the psychiatric unit at Johns Hopkins in a six-month period had detectable levels of this drug in their serum.

Wuth described the therapeutic and toxic serum levels found in patients on bromide therapy for epilepsy. Based on clinical information he established 15 mmol/L (1 200 mg/L) as the upper limit of therapeutic and results over 19 mmol/L (1 500 mg/L) as toxic. He further recommended that patients on bromide therapy for the control of epilepsy be routinely monitored for serum levels. Wuth demonstrated with his case studies that the blood levels of bromide correlate with therapeutic efficacy and not with the dose administered {Wuth, 1927 #23457}. Note: This fundamental principal in therapeutic drug monitoring (TDM) was advocated by Wuth almost 50 years before it became standard practice in clinical medicine.


Fritz Eichholtz (DE) and Otto Butzengeiger (DE) carried out the first experimental use of avertin (2,2,2-tribromoethanol) as an anesthetic in animals {Eichholtz, 1927 #17814}.

Otto Butzengeiger (DE) introduced avertin (2,2,2-tribromoethanol) as a rectal anesthetic in the clinic {Butzengeiger, 1927 #14592}.


Major Greenwood (GB) and Janet Elizabeth Lane-Claypon (GB) developed a key type of epidemiological investigation, the so-called "case-control study". They tracked down 500 women with a history of breast cancer- the “cases” - and compared them with 500 women who were free of the disease but otherwise broadly similar, known as “controls”. The detailed survey that emerged constituted, as far as is known, the first published epidemiological questionnaire. This yielded results that enabled them to identify many of the risk factors for breast cancer that are still considered valid today. Their conclusions (or their data reworked by later researchers) agreed with those of modern reviewers: breast cancer was associated with age at menopause, artificial menopause, age at first pregnancy (age at marriage used as a proxy), number of children, and lactation.

They published what is now considered the first “end results” study. It followed a large sample of women with pathologically confirmed breast cancer for up to 10 years after their surgery. The study confirmed that women who were surgically treated at an early stage of the disease had a much better chance of surviving three, five, or 10 years longer than those operated on at any later stage. They showed that breast cancer risk increased for childless women, women who married later than average, and women who did not breast feed. The overall breast cancer risk decreased according to the number of children. For all cases, rapid treatment held the key to survival among women with breast cancer. In reviewing the family histories of their cases, they anticipated the role that genes might play in the development of breast cancer. “There appear to be some families,” they wrote, “in which for reasons not certain at present, cancer plays havoc with the members, and there is (some) slight evidence in some instances that it attacks the same organs” {Greenwood, 1927 #20221}.


Charles Sutherland Elton (GB) redefined the species niche by emphasizing its functional role in the community: the niche is “the status of an animal in its community.” His concept idealized what the species does rather than where it lives—the functional niche concept {Elton, 1927 #8307}.


Erik Anderson Stensiö (SE) reconstructed a fossil Cephalaspis (an ostracoderm) and suggested its status as a vertebrate prototype replacing amphioxus {Stensiö, 1927 #14167}. Note: it is generally believed that living Agnatha (hagfishes and lampreys) and the ostracoderms are descended from a common ancestor.


Vladimir Mikhailovich Bekhterev (RU), in 1927, diagnosed Joseph Stalin with “grave paranoia.” Later that day, Bekhterev suddenly died causing speculation that Stalin had his revenge for the diagnosis. After Bekhterev’s death, Stalin had Bekhterev’s name and all his works removed from textbooks.


Arthur Keith (GB) discovered a human upper jaw in Southern England and initially diagnosed it as Upper Paleolithic modern human {Keith, 1927 #20359}.

Thomas F.G. Higham (GB), Tim Compton (GB), Chris Stringer (GB), Roger Jacobi (GB), Beth Shapiro (US), Erik Trinkaus (US), Barry Chandler (GB), Flora Gröning (GB), Chris Collins (GB), Simon Hillison (UK), Paul O'Higgins (GB), Charles FitzGerald (UK), and Michael Fagan (UK) tested the jaw Keith found using a Bayesian analysis of new ultra-filtered bone collagen. It was dated in an ordered stratigraphic sequence at the site to 44.2–41.5 K BP. This makes it older than any other equivalently dated modern human specimen and directly contemporary with the latest European Neanderthals {Higham, 2011 #20360}.


Arturo Palma di Cesnola (IT) and Borzatti von Löwenstern (IT), in 1964, found two deciduous molars in the so-called Uluzzian archaeological layers unearthed from the Grotta del Cavallo (Southern Italy). Classified as Neanderthal and dated to 45,000 years ago these fossils are the oldest known human remains on the continent {di Cesnola, 1964 #22452}.

Stefano Benazzi (AT), Katerina Douka (GB), Cinzia Fornai (AT), Catherine C. Bauer (DE), Ottmar Kullmer (DE), Jiří Svoboda (CZ), Ildikó Pap (HU), Francesco Mallegni (IT), Priscilla Bayle (FR), Michael Coquerelle (ES), Silvana Condemi (FR), Annamaria Ronchitelli (IT), Katerina Harvati (DE) and Gerhard W. Weber (AT) reanalyzed the deciduous molars from the Grotta del Cavallo (Southern Italy), associated with the Uluzzian and originally classified as Neanderthal. Their new chronometric data for the Uluzzian layers of Grotta del Cavallo obtained from associated shell beads and included within a Bayesian age model show that the teeth must date to ~45–43 K BP. The Cavallo human remains are therefore the oldest known European anatomically modern humans, confirming a rapid dispersal of modern humans across the continent before the Aurignacian and the disappearance of Neanderthals {Benazzi, 2011 #20361}.



“Error is modern while truth is ancient.” Ramón y Cajal {Ramón y Cajal, 1928 #5587}.


"Only when some important function... lends itself to ready observation or quantitative measurement are the conditions suitable for making progress..." Edwin B. Hart, et al., {Hart, 1928 #3064}.


“As I view my contribution to the writing of our time, it seems to me to consist of a double affirmative, saying first that an awareness and experience of Nature is necessary to Man if he is to have his humanity, and saying in the second place that that same awareness must have something of a religious quality, the Italian pieta, if you will.

Nature is a part of our humanity, and without some awareness and experience of that divine mystery man ceases to be man. When the Pleiades and the wind in the grass are no longer a part of the human spirit, a part of the very flesh and bone, man becomes, as it were, a kind of cosmic outlaw, having neither the completeness and the integrity of the animal nor the birthright of a true humanity.” Henry Beston {Beston, 1928 #10805}.


Adolf Otto Rheinhold Windaus (DE) was awarded the Nobel Prize in Chemistry for research into the constitution of the sterols and their connection with the vitamins.


Charles Jules Henri Nicolle (FR) was awarded the Nobel Prize in Physiology or Medicine for his work on typhus.


Kurt Heinrich Meyer (DE) and Herman Francis Mark (DE) proved the existence of polymeric chain molecules by examining the crystalline structure of polymers with x-rays {Mark, 1928a #12198;Mark, 1928b #12199;Mark, 1928c #12200}.


Otto Fritz Meyerhof (DE-US) and Karl Lohmann (DE) isolated argininephosphoric acid from arthropod (crustacean) muscle. It was found to be the invertebrate analogue of the vertebrate phosphagen (creatine phosphate) {Meyerhof, 1928a #8625;Meyerhof, 1928b #8626;Meyerhof, 1928c #8627}.

Karl Lohmann (DE) and Hermann Lehmann (DE-GB) demonstrated that phosphagen (creatine phosphate) reacts with ADP in minced crustacean muscle {Lohmann, 1935a #8628;Lehmann, 1936 #8629}. This provided, for the first time, a mechanism for utilization of phosphate energy. This became known as the Lohmann reaction.


Marjory Stephenson (GB) obtained the first cell-free preparation of non-NAD dependent lactic dehydrogenase, which oxidized lactate to pyruvate in the presence of methylene blue {Stephenson, 1928 #19715}.

Bengt Andersson (SE) provided evidence for identifying the coenzyme necessary for reducing pyruvate with NAD {Andersson, 1933 #19713;Andersson, 1934 #19714}.


Oliver Kamm (US), Thomas B. Aldrich (US), Irvine W. Grote (US), Louis W. Rowe (US), Edwin P. Bugbee (US) started the isolation and purification of vasopressin and oxytocin from the posterior lobe of the pituitary gland {Kamm, 1928 #5453}. Note: Arginine vasopressin (AVP) is synthesized in the supraoptic nuclei of the hypothalamus and transported within axons to the posterior lobe of the pituitary for storage. The gene encoding AVP is arginine vasopressin–neurophysin II (AVP–NPII), located on chromosome 20p13.


George William Marshall Findley (GB) demonstrated the association of skin cancer with exposure to ultraviolet light {Findlay, 1928 #23764}.


Leslie Julius Harris (GB) and Thomas Moore (GB) confirmed the toxic action of irradiated ergosterol in excess and concluded that "a weight of evidence pointed to the toxic action of excess of vitamin D {Harris, 1928 #23387}.


Donald Dexter van Slyke (US) and James A. Hawkins (US) presented a gasometric method for determination of reducing sugars, and its application to analysis of blood and urine {van Slyke, 1928b #10940}.


Robert Robison (GB) and Walter Thomas James Morgan (GB) isolated crystalline trehalose from trehalosemonophosphate {Robison, 1928 #18991}.


F. Gottwalt Fischer (DE) and Kurt Löwenberg (DE) determined the structure of phytol (the major esterifying alcohol of chlorophyll at position 7 of the macrocycle) {Fischer, 1928 #12254}.

F. Gottwalt Fischer (DE) and Kurt Löwenberg (DE) synthesized phytol from pseudoionone {Fischer, 1929 #12255}.


James Waddell (US), Conrad Arnold Elvehjem (US), Harry Steenbock (US), Edwin Bret Hart (US), Evelyn Van Donk (US), and Blanche M. Riising (US) discovered that inorganic iron salts are ineffective at increasing the hemoglobin levels of anemic rats, but that ashed residues from dried beef liver, dried lettuce, and yellow corn were very effective in curing anemia. From these results they inferred that the extracts contained some other inorganic substance that was necessary to produce hemoglobin. They later discovered that this necessary substance is copper {Waddell, 1928 #21464}.

James Waddell (US), Harry Steenbock (US), Conrad Arnold Elvehjem (US), Edwin Bret Hart (US), and Evelyn Van Donk (US) found that copper-containing liver extracts and a copper sulfate solution both served equally well in curing anemia in rats. This led them to the conclusion that, “the deficiency (in this anemia) is inorganic in nature and that this inorganic deficiency is copper only” {Waddell, 1929 #21465}.


Hubert Bradford Vickery (CA-US) and Charles S. Leavenworth (US) carried out an analysis of the basic amino acids in horse hemoglobin and concluded that the protein contains 7.64% histidine, 3.32% arginine, and 8.10% lysine. These results agreed with the assumption that hemoglobin has 33 molecules of histidine, 13 molecules of arginine, and 37 molecules of lysine {Vickery, 1928 #21450}.


Bertil Hanstöm (SE) discovered that the X-organ, located in the eyestalk of crustaceans, secretes neurohormones {Hanström, 1928 #15543;Hanström, 1931 #19447;Hanström, 1933 #19446}. Neurosecretory cells in the X-organ (part of the brain) produce a molt-inhibiting hormone that is stored in the sinus gland of the eyestalk, while a molting hormone is produced in the Y-organ. Interactions of these two hormones control the molting process.


Bernhard Zondek (DE-IL) and Selmar Aschheim (DE) isolated from the anterior pituitary gland a gonadotropic hormone they named prolan (it was luteinizing hormone-LH-and/or follicle stimulating hormone-FSH) {Zondek, 1928 #14593}. Prolan is a term no longer in use.


Fritz Albert Lipmann (DE-US), Vladimir Aleksandrovich Engelhardt (RU), and A.P. Barkash (RU) presented evidence for the metabolic pathway from glucose-6-phosphate to triose phosphate by way of ribose-5-phosphate. They called it the hexose monophosphate shunt {Lipmann, 1928 #8346;Engelhardt, 1938 #9269}.


Hans Sachs (DE) and Ernst Witebsky (DE-US) developed a precipitation reaction for serological diagnostics of syphilis {Sachs, 1928 #13576}.


Frederick Griffith (GB) discovered the transforming factor while working with Diplococcus pneumoniae (Streptococcus pneumoniae). He demonstrated that this factor, which he isolated from smooth colonies, is a chemical that is present in the extract of dead smooth colonies but missing from live rough colonies. When mixed with living rough colonies the transforming factor converted many of them to living smooth colonies {Griffith, 1928 #621}.

James Lionel Alloway (US), working in the laboratory of Oswald Theodore Avery (CA-US), broke open cells of the smooth form of Pneumococcus and collected the cell contents, which were passed through a filter that would remove all wall material. When this extract was added to a culture of growing rough cells some of them were transformed into smooth cells. When the transforming extract was treated with alcohol a thick syrupy precipitate formed {Alloway, 1932 #708}.

Oswald Theodore Avery (CA-US) would later show that the transforming factor is DNA. See, Sanfelice, 1893 and Avery, 1944.


Andrew C. Ivy (US) and Eric Oldberg (US) discovered and named the hormone cholecystokinin (CCK) which controls gall bladder function {Ivy, 1928 #3164}.

Viktor Mutt (EE) and J. Erik Jorpes (SE) determined the structure of cholecystokinin (CCK) {Mutt, 1968 #12233}.


Julian Walawski (PL) and Jerzy Kaulbersz (PL) discovered enterogastrons produced in the large intestine, which inhibit the stomach secretion {Walawski, 1928a #17345;Walawski, 1928b #17346}.


Albert Imre Szent-Györgyi (HU-US) using the adrenal cortex, cabbage, and orange as raw material isolated and crystallized a factor which he knew was in some way involved in the mechanism of biological oxidation. He named this acidic carbohydrate hexuronic acid {Szent-Györgyi, 1928 #2524}.

Charles Glen King (US) and William A. Waugh (US) succeeded in obtaining Szent-Györgyi’s hexuronic acid in crystalline form then equated it with vitamin C {King, 1932 #2975;Waugh, 1932 #2976}. Today it is known as ascorbic acid (ascorbic meaning, no scurvy) or vitamin C.

Szent-Györgyi (HU-US) and Walter Norman Haworth (GB) suggested that vitamin C be renamed ascorbic acid {Szent-Györgyi, 1933 #2971}.

Walter Norman Haworth (GB), Luszló Vargha (HU), Ernest G. Cox (GB), Edmund Langley Hirst (GB), and Reginald John William Reynolds (GB) would contribute to the determination of its chemical structure {Haworth, 1933 #2977;Vargha, 1932 #2978;Cox, 1932 #2979}.

Walter Norman Haworth (GB) and Edmund Langley Hirst (GB) synthesized ascorbic acid {Haworth, 1933 #2977}.


Cornelis Bernardus Kees van Niel (NL-US), working with propionic acid bacteria, provided the first quantitative picture of the products derived by these bacteria from lactate, glycerol, glucose, and starch. His taxonomic treatment determined the veracity of the genus Propionibacterium. During these studies he identified diacetyl as the compound responsible for the characteristic aroma of high quality butter {van Niel, 1928 #10778}.


Pierre Stricker (FR) and Fritz Grüeter (FR) induced mammary gland development and the secretion of milk by injecting anterior pituitary extract into castrated virgin rabbits {Stricker, 1928 #12241;Stricker, 1929 #19792}. This strongly suggested the presence of a lactation hormone (prolactin).

Herbert McLean Evans (US) and Miriam E. Simpson (US) found that extracts from the anterior pituitary cause hypertrophy of the mammary glands in virgin rats {Evans, 1929 #10429}.

Oscar Riddle (US) and Pela Fay Braucher (US) showed that extracts of the anterior lobe of the pituitary gland could cause the enlargement and functioning of the crop-glands in pigeons {Riddle, 1931 #12242}.

Oscar Riddle (US), Robert Wesley Bates (US), and Simon William Dykshorn (US) reported their discovery of a hormone produced in the anterior pituitary gland, which stimulates the crop-gland in pigeons and is lactogenic in guinea pigs; they named it prolactin {Riddle, 1932 #4845;Riddle, 1933 #3382}.

Abraham White (US), Hubert Ralph Catchpole (GB-US), and Cyril Norman Hugh Long (GB-US) crystallized the lactogenic hormone prolactin {White, 1937 #6709}.

Choh Hao Li (CN-US), Jonathan S. Dixon (US), Tung-bin Lo (TW), Knud D. Schmidt (DK), and Yuri A. Pankov (RU) determined the primary amino acid sequence of the sheep lactogenic hormone prolactin {Li, 1969b #15410;Li, 1970b #10176}.

Henry George Friesen (CA), Harvey J. Guyda (CA), and Jules Hardy (CA) discovered prolactin in humans {Friesen, 1965 #9823;Friesen, 1970 #9824}.

G. Michael Besser (US), Lynne Parke (GB), Christopher R.W. Edwards (GB), Isabel A. Forsyth (GB), and Alan S. McNeilly (US) found that they could successfully treat galactorrhoea by reducing plasma prolactin levels using brom-ergocryptine {Besser, 1972 #9351}.

Eva Nagy (HU-CA) and Istvan Berczi (HU-CA) reported that regulation of the immune response by the neuroendocrine hormone prolactin (PRL) occurs via its interaction with the prolactin receptor. Interference with this ligand-receptor interaction inhibits both in vitro and in vivo immune responses {Nagy, 1978 #14597}.

Charles V. Clevenger (US), Amy L. Sillman (US), Michael B. Prystowsky (US), Yi-Ping Rao (US), Donna J. Buckley (US), Mark D. Olson (US) and Arthur R. Buckley (US) showed that prolactin is able to physically enter the cell, travel straight to the cell’s DNA, and directly activate the process that turns on genes and triggers the growth of breast cancer cells. It does this by binding to a protein called cyclophilin B, or CYPB for short {Clevenger, 1990 #14599;Rao, 1995 #14600}.

Charles V. Clevenger (US), Tracey L. Plank (US), Susan E. Hankinson (US), Walter C. Willett (US), Dominique S. Michaud (US), JoAnn E. Manson (US), Graham A. Colditz (US), Christopher Langcope (US), Bernard Rosner (US), Frank E. Speizer (US), Marcela V. Maus (US), and Sean C. Reilly (US) presented evidence that the somatolactogenic hormone prolactin, a naturally occurring peptide hormone needed for milk production following pregnancy, stimulates the movement, or motility of breast cancer cells causing the cells to essentially pick up and move, and actually trigger invasive potential of these cells {Clevenger, 1997 #14596;Hankinson, 1999 #14595;Maus, 1999 #14594}.


Georg von Békésy (HU-US) elucidated all the physical events at every strategically important point in the transmission system of the ear by recording events in this fragile biological miniature system, micro dissection, advanced teletechniques for stimulation and recording, and high magnification stroboscopic microscopy. He elucidated the vibration patterns of the eardrum and the interplay of the ossicle movements, provided experimental and clinical data confirming Helmholtz’s assumption that the frequency of the sound waves determines the location along the basilar membrane at which stimulation occurs. He found that movements of the stirrup footplate evokes a wave complex in the basilar membrane, which travels from the stiffer basal part to the more flexible part in the apex of the cochlea. The crest of the largest wave first increases, thereafter quickly decreases. The position of the maximal amplitude was found to be dependent on the frequency of the stimulating sound waves in such a way that the highest crest of the travelling wave appears near the apex of the cochlea at low-frequency tones and near its base at high frequencies {von Békésy, 1928 #14150;von Békésy, 1930 #17668;von Békésy, 1960 #9178;von Békésy, 1967 #6599;von Helmholtz, 1863 #12831}.


Donald Dexter van Slyke (US) and Julius Sendroy, Jr. (US) reported studies of gas and electrolyte equilibria in blood in which they present line charts for graphic calculations by the Henderson-Hasselbalch equation, and for calculating plasma carbon dioxide content from whole blood content {van Slyke, 1928a #9326}. Van Slyke continued to improve this method through eleven more papers.


Thomas Hunt Morgan (US) presented the theory of the gene when he said, “The theory states that the characters of the individual are referable to paired elements (genes) in the germinal material that are held together in a definite number of linkage groups; it states that the members of each pair of genes separate when the germ-cells mature in accordance with Johann Gregor Mendel’s first law, and in consequence each germ-cell comes to contain one set only; it states that the members belonging to different linkage groups assort independently in accordance with Johann Gregor Mendel’s second law; it states that an orderly interchange—crossing over— also takes place, at times, between elements in corresponding linkage groups; and it states that the frequency of crossing over furnishes evidence of the linear order of the elements in each linkage group and of the relative position of the elements with respect to each other” {Morgan, 1928 #2473}.


Hermann Joseph Muller, Jr. (US) and Theophilus Shickel Painter (US), using Drosophila, carried out parallel investigations in which phenotypic variations brought about by x-rays were related to physical changes in the chromosomes, e.g., deletions and translocations {Muller, 1928 #12182;Painter, 1929 #10777}.


Johann Heinrich Emil Heitz (DE) coined the terms euchromatin and heterochromatin {Heitz, 1928 #12384}.


Karl Heinrich Bauer (DE) proposed the theory that cancers arise because of mutations {Bauer, 1928 #23771}. Ernest Edward Tyzzer (US) coined the phrase “somatic mutation” {Tyzzer, 1916 #23772}.


Eggert Hugo Heiberg Møller (US), John F. McIntosh (US), and Donald Dexter van Slyke (US) determined that a constant volume of blood is being cleared of urea in each minute’s time course. This came to be referred to as standard blood urea clearances {Møller, 1928 #3366}.


Adrian Stokes (GB), Johannes H. Bauer (US), N. Paul Hudson (US), Constant Mathis (FR), Andrew Watson Sellards (US), and Jean Laigret (FR), working in West Africa, proved that yellow fever is caused by a virus and transferable to monkeys {Mathis, 1928 #6977;Stokes, 1928a #6978;Stokes, 1928b #9120}. Note: Stokes died of a yellow fever infection.


Renjiro Kaneko (JP) and Yoshio Aoki (JP) determined that the etiological agent of Japanese B encephalitis was probably a virus {Kaneko, 1928 #6974}.

Michitomo Hayashi (JP) transmitted Japanese encephalitis one of the leading causes of acute encephalopathy— from an infected human to monkeys by way of an intracerebral injection to prove the viral etiology of the disease {Hayashi, 1934 #15528}.

Itsuma Takaki (JP), Tenji Taniguchi (JP), M. Hosokawa (JP), and S. Kuga (JP) isolated the viral etiological agent of Japanese B encephalitis {Takaki, 1926 #6975;Taniguchi, 1936 #6976}.


Stefan Nicolau (GB) and Ian Galloway (GB) discovered the virus of Borna disease and enzootic encephalo-myelitis of sheep and cattle {Nicolau, 1928 #25960}. Note: the only bornavirus


Jean-Louis Verge (FR) and N. Christoforoni (FR) were the first to associate a virus with feline panleukopenia {Verge, 1928 #25961}. Note: the first parvovirus


Thomas Milton Rivers (US) edited the first major virology book, Filterable Viruses. Harold L. Amoss, Jacques J. Bronfenbrenner, Alexis Carrel, and others, were contributors {Rivers, 1928 #25962}.


Arthur T. Henrici (US) reported that the average size of bacterial cells might vary considerably from one growth phase to another during a growth cycle {Henrici, 1928 #7116}.


Clifford Dobell (US) was the first to describe encystment of a parasitic amoebic form, Entamoeba histolytica {Dobell, 1928-1938 #20491}.


Kenneth L. Burdon (US) described Bacteroides melaninogenicus in the stools of patients suffering from chronic amebic dysentery, and in puerperal sepsis. This bacterium is often associated with the mouth, tonsils, infected abdominal wounds, and focal infections of the kidneys {Burdon, 1928 #2247}.


Henry E. Meleney (US) presented clear evidence of the development of six immunologically distinct strains of Borrelia recurrentis {Meleney, 1928 #106}.


Karl Johannes (Hans) Kniep (DE) and Arthur Henry Reginald Buller (CA) had previously presented the broad picture of the genetic control system and the developmental sequence from spore to spore in the basidiomycetes {Kniep, 1928 #10011;Buller, 1909 #10010}. Buller was also a poet. One of his most recognized works is this limerick, first published in the Dec. 19, 1923 issue of Punch:

There was a young lady named Bright

Whose speed was far faster than light;

She set out one day,

In a relative way

And returned on the previous night.


Émile Eugène Aldric Topsent (FR), spongiologist, described the Atlantic and Mediterranean collections of Prince Albert of Monaco in three monumental volumes and erected many genera and families. His work is the basis for the current classification system of the Porifera {Topsent, 1928 #6730}. Topsent is commemorated by Acheliderma topsenti Burton, 1932, Axinyssa topsenti Lendenfeld, 1897; Chondropsis topsenti Dendy, 1895; Corticium topsenti Pouliquen, 1972; Cryptotethya topsenti Thiele, 1900, Desmacella topsenti Burton, 1930; Desmanthus topsenti Hentschel, 1912; Dragmatyle topsenti Burton, 1954; Dysideopsis topsenti Hentschel, 1912; Echinodictyum topsenti De Laubenfels, 1936; Erylus topsenti Lendenfeld, 1903; Eurypon topsenti Pulitzer-Finali, 1983; Grayella topsenti Babic, 1922; Haddonella topsenti I. Sollas, 1903; Halichondria topsenti De Laubenfels, 1936; Hymenotrocha topsenti Burton, 1930; Hymerhabdia topsenti Lévi, 1952; Jaspis topsenti Thiele, 1900; Leucandra topsenti Breitfuss, 1929; Raspailia topsenti Dendy, 1924; Reniera topsenti Thiele, 1905; Rhabderemia topsenti Van Soest & Hooper, 1993; Rhabdoploca topsenti Hentschel, 1912; Spongosorites topsenti Dendy, 1905; Stelletta topsenti Thiele,1903; Suberella topsenti Burton, 1929; Tedania topsenti De Laubenfels, 1930; Topsentia Berg, 1899; and Tylaspis topsenti Lévi & Lévi, 1983.


Remington Kellogg (US) wrote The History of Whales—Their Adaptation to Life in the Water, still one of the best summaries of the subject {Kellogg, 1928 #10951}.


John Walton (GB) presented evidence of mosses and liverworts from the Carboniferous deposits in England {Walton, 1928 #5134}.

Thomas M. Harris (GB) presented excellent evidence of mosses in the Triassic of England {Harris, 1938 #5135;Harris, 1939 #5136}.


Royal N. Chapman (US) used flour beetles (Tribolium confusum Duval) to devise an experimental single-species model for studying the growth and regulation of closed, single-species systems {Chapman, 1928 #1779}.

Jerzy Neyman (US), Thomas Park (US), and Elizabeth L. Scott (US) later christened it the Tribolium model {Neyman, 1956 #1780}.


Herbert Friedmann (US) described social parasitism among birds and wrote, The Cowbirds: A Study in the Biology of Social Parasitism, a definitive treatment of brood parasitism {Friedmann, 1928 #10689;Friedmann, 1929 #10688}. This type of behavior is seen among the cuckoos, cowbirds, honey guides, and weaverbirds.


Clara M. Davis (US) pioneered experiments on self-selection of diet by healthy infants that led to new studies of eating behavior, including sensory-specific satiety {Davis, 1928 #23436}.


Archibald Philip Bard (US) discovered that the emotion we call rage depends on neurons in the caudal half of the hypothalamus {Bard, 1928 #6719;Bard, 1929a #17743;Bard, 1929b #17744;Bard, 1930 #17745}.


Edgar V. Allen (US), Jean Paul Pratt (US), Quitman U. Newell (US), and Leland J. Bland (US) used dogs to develop surgical methods of recovering embryos for application in humans {Allen, 1928 #20333}.


Hans Zinsser (US) and H. Yu (US) were among the first to suggest that diseases such as rheumatic fever and glomerulonephritis result from hypersensitivity to toxins produced by certain strains of streptococci {Zinsser, 1928a #6919;Zinsser, 1928b #6920}.


Georgios Nikolaou Papanikolaou; George Nicholas Papanicolaou (Greek-US) and Herbert Frederick Traut (US) developed the Cervical Smear Test (Pap smear for Papanicolaou) for the detection of uterine cancer. It is based on the cytological examination of cells exfoliated from the uterus {Papanicolaou, 1928 #13442;Papanicolaou, 1941 #11859;Papanicolaou, 1943 #13443;Papanicolaou, 1949 #23773}. Note: This test was perfected as the first clinical exam useful as an early detection method for cervical cancer.

Olive Gates (US) and Shields Warren (US) wrote important articles and a handbook on cervical cytopathology {Gates, 1945 #27345;Gates, 1947 #27346}.


Selmar Aschheim (DE) and Bernhard Zondek (DE-IL) reported their pregnancy test (Aschheim-Zondek Pregnancy Test) for humans, “Our test is carried out with morning urine…. The urine is injected subcutaneously into the infantile [female mice] …. Only the ovarian findings are of significance for the pregnancy reaction [enlargement with follicular maturation] …. We have examined 78 cases of pregnancy. In 76 cases the reaction was definitely positive…. In [the] 198 control cases the reaction was positive twice…. The reaction thus has a precision that one cannot hope to surpass with a biologic method” {Aschheim, 1928a #3386;Aschheim, 1930 #3388}.


Owen Harding Wangensteen (US) and George W. Waldron (US) did studies in intestinal obstruction, which led to the development of the Wangensteen suction technique to relieve increased internal viscus pressure in the stomach and intestine resulting from excess gastric and intestinal secretions in an atonic intestine {Wangensteen, 1928 #10752}.


Walter Edward Dandy (US) devised an operation to relieve Ménière’s disease (inflammation and congestion of the semicircular canals) {Dandy, 1928 #13177;Dandy, 1941 #13178}.


Harvey Williams Cushing (US) and Percival Bailey (US) provided the first extensive classification and description of angiomatous malformations and hemangioblastomas {Cushing, 1928b #13174}.


Percival Bailey (US) and Harvey Williams Cushing (US) were the first to describe the condition known as fugitive acromegaly, in which patients may exhibit physical stigmata of acromegaly without biochemical evidence of the disease {Bailey, 1928 #16886}.


Howard Christian Naffziger (US) devised an operation to relieve severe malignant exophthalmos using orbital decompression {Naffziger, 1928 #13422}.

Jörgen H. Vogt (NO) and Arne Torkildsen (NO) named this Naffziger’s operation {Vogt, 1947 #13423}.


Andrew Ellicott Douglass (US) rediscovered that annual growth of tree rings can be used to construct a window on the weather of the past. Trees add a layer of wood to their trunks every year - a wide ring during wet years and a thin one during dry years. By matching ring patterns in living trees to the patterns in old timbers, the record could be extended further back into history, i.e., dendrochronology {Douglass, 1928 #7461}. This method is most accurate from the present to 12,000 years ago. See, Leonardo da Vinci c.1497; Henri-Louis Duhamel du Monceau and Georges-Louis Leclerc de Buffon 1737.


Adolf Remane (DE), working in the Kiel Bight (Baltic Sea) in 1928, discovered the new phylum Gnathostomulida (jaw worms), a small (0.5 mm long) marine worm-like form with worldwide distribution {Remane, 1936 #19790}.

Peter Ax (DE) originally described Gnathostomulida as an order of the Platyhelminthes {Ax, 1956 #18459;Ax, 1965 #16812}.


John Burdon Sanderson Haldane (GB-IN), in 1928, wrote the essay, On Being the Right Size, which Jane Jacobs and others have since referred to as Haldane's principle. This is that sheer size very often defines what bodily equipment an animal must have: “Insects, being so small, do not have oxygen-carrying bloodstreams. What little oxygen their cells require can be absorbed by simple diffusion of air through their bodies. But being larger means an animal must take on complicated oxygen pumping and distributing systems to reach all the cells” {Haldane, 1985 #14745}.



“Everything is determined… by forces over which we have no control. It is determined for the insect as well as for the star. Human beings, vegetables, or cosmic dust—we all dance to a mysterious tune, intoned in the distance by an invisible piper.” Albert Einstein {Einstein, 1929 #1473;Clark, 1971 #1474}.


Arthur Harden (GB) and Hans Karl August Simon von Euler-Chelpin (DE-SE) were awarded the Nobel Prize in Chemistry for their investigations on the fermentation of sugar and fermentative enzymes.


Christiaan Eijkman (NL) for his discovery of the antineuritic vitamin (thiamine, vitamin B1) and Frederick Gowland Hopkins (GB) for his discovery of the growth-stimulating vitamins shared the Nobel Prize in physiology and medicine.


Dame Kathleen Yardley Lonsdale (GB) was the first to use x-ray diffraction to solve the structure of an organic molecule, she showed that hexamethylbenzene is planar and hexagonal - and gave its precise dimensions {Lonsdale, 1929 #14584}.


Sergei Yakovievich Sokolov (RU) suggested the concept of the pulse-echo ultrasonic metal flaw detector {Sokolov, 1929 #12237}. This instrument was the precursor of subsequent pulse-echo medical ultrasonic (ultrasound) devices operating in the uni-directional A-mode and used as early as 1949 by Lars Leksell (SE) and J.C. Turner (GB) for examining brain lesions {Leksell, 1949 #12239;Leksell, 1951 #7672}.

George Döring Ludwig (US), Francis W. Struthers (US), Richard H. Bolt (US), Theodor F. Hueter (US), and Henry Thomas Ballantine, Jr. (US) made significant progress toward understanding the usefulness and limitations of ultrasound for clinical examination {Ballantine, 1950a #20150;Ballantine, 1950b #20151;Ludwig, 1949 #20146;Ludwig, 1950a #4876;Ludwig, 1950b #20147;Ludwig, 1950c #20148}.

George Andrew Douglas Gordon (GB), J.C. Turner (GB), William Valentine Mayneord (GB), Stigg Jeppson (SE), Brita Lithander (SE), Marinus de Vlieger (NL), Kenji Tanaka (JP) and Toshio Wagai (JP) were among the first practitioners to use ultrasound in clinical settings.


Walter Norman Haworth (GB) determined that a number of stereo isomeric structures are possible for the pyranose ring of each sugar and showed that glucose in the six-member (pyranose) form is more stable than when it is in the five-member (furanose) form or in the Fischer form, i.e., Haworth structures for sugars {Haworth, 1929 #12202}.


Karl Lohmann (DE), Cyrus Hartwell Fiske (US), and Yellapragada SubbaRow (US), found a new organic phosphate related to muscle adenylic acid and named it adenylpyrophosphate {Fiske, 1929a #2565;Lohmann, 1929 #2564}. Note: Priority for the discovery of ATP was awarded to Lohmann.

Karl Lohmann (DE) determined its structure and recognized it to be adenosine-5’-triphosphate (ATP). Lohmann (DE) also characterized adenosine-5’-diphosphate (ADP) {Lohmann, 1935b #2560}.

Otto Fritz Meyerhof (DE-US) and Karl Lohmann (DE) stated that the role of adenylpyrophosphate (ATP), “… appears to consist in the fact that the esterification of phosphate, which precedes the cleavage of carbohydrate to lactic acid, occurs with the simultaneous cleavage of adenylpyrophosphate, which is resynthesized during the further cleavage [of carbohydrate]. In this manner, the adenylpyrophosphate cycle maintains the lactic acid formation. The synthesis of phosphagen (creatine phosphate) is therefore made possible … by the cleavage energy of the adenylpyrophosphate, while the energy of lactic acid formation (from phosphate esters) serves to resynthesize the cleaved pyrophosphate” {Meyerhof, 1931 #2566}.


John Howard Northrop (US) isolated swine pepsin in pure crystalline form by techniques, which were later used by him and other workers to crystallize trypsin, chymotrypsin, carboxypeptidase, and pepsinogen {Northrop, 1929 #19737}.


Marjorie Martland (GB) and Robert Robison (GB) observed that during the hydrolysis of fructosediphosphate (fructose-1,6-diphosphate) by preparations of bone phosphatase, a part of the sugars liberated, following the removal of the phosphoric acid groups, suffered an intramolecular change and gave rise to a mixture of aldolase and ketose sugars {Martland, 1929 #18992}.


Ernst Waldschmidt-Leitz (CZ) and Arnulf Purr (CZ-US) identified the enzyme carboxypolypeptidase (carboxypeptidase) in bovine pancreatic extracts. They found that it cleaves C-terminal amino acids from acetylated peptides {Waldschmidt-Leitz, 1929 #12170}.


Phoebus Aaron Theodor Levene (RU-US) and Efim Semenovich London (RU) isolated and described 2-deoxyribose (they called it thyminose because it came from thymus nucleic acid, i.e., DNA, as belonging to a group of pentoses lacking an oxygen atom, and hence named desoxypentoses then later named deoxypentoses {Levene, 1929a #14636;Levene, 1929b #12240}.


George Oswald Burr (US), Mildred M. Burr (US), and Elmer S. Miller (US) demonstrated the existence and necessity of the so-called essential fatty acids. At that time, essentiality meant promotion of growth and prevention of the dermatitis observed when a fat-free diet was fed to rats {Burr, 1929 #2744;Burr, 1930 #3072;Burr, 1932 #3073}.

Osmo Turpeinen (US) later proved that arachidonic acid is an essential fatty acid {Turpeinen, 1938 #3074}.


Adolf Friedrich Johann Butenandt (DE) isolated the hormone estrone (3-hydroxy 17-keto delta 1,3,5-estratriene) from the urine of pregnant women {Butenandt, 1929 #14605}.

Clement D. Veler (US), Sidney Allen Thayer (US), and Edward Adelbert Doisy (US) independently isolated estrone (3-hydroxy 17-keto delta 1,3,5-estratriene) from the urine of pregnant women. They called it Theelin {Veler, 1930 #12245}.

Sidney Allen Thayer (US), Louis Levin (US), and Edward Adelbert Doisy (US) characterized estrone (Theelin) {Thayer, 1931 #12246}.


Michel Macheboeuf (FR) isolated the first plasma lipoprotein to exhibit a constant composition {Macheboeuf, 1929 #5459}. This fraction was later characterized as an alpha-1-globulin that we now recognize as high-density lipoprotein (HDL).


Henry Hallett Dale (GB) and Harold Ward Dudley (GB) isolated histamine and acetylcholine from the spleens of ox and horse {Dale, 1929 #5282}.


Charles Robert Harington (GB) and Sydney Stewart Randall (GB) isolated 3,5, di-iodotyrosine from thyroglobulin of the thyroid gland {Harington, 1929 #3102}.


Karl Paul Gerhard Link (US), Herbert Raleigh Angell (GB-AU), Allan D. Dickson (US), and John Charles Walker (US) established for the first time a specific chemical difference between a resistant host (pigmented onion) and a non-resistant host (the white onion). They determined that the brown-pigmented onion produces the antifungal agent protocatechuic acid, which the white onion lacks {Link, 1929a #10390;Link, 1929b #10388;Link, 1929c #11184}.


Leo Loeb (CH-US) and Robert B. Bassett (US) isolated thyroid-stimulating hormone (TSH) from the anterior pituitary gland of cattle and demonstrated its effect on guinea pigs {Loeb, 1929 #7817}.

Max Aron (DE) independently isolated thyroid-stimulating hormone (TSH) from the anterior pituitary {Aron, 1929 #13063}.


Carl Richard Moore (US) Thomas F. Gallagher (US) and Fred C. Koch (US) obtained male sex hormone in relatively pure form. Moore and his colleagues introduced the term testosterone {Moore, 1929a #12243;Moore, 1929b #12244}.


George Washington Corner (US) and Willard Myron Allen (US) found, ” that alcoholic extracts of the corpus luteum, freed of phospholipids, contains a substance which when injected into castrated adult female rabbits induces a characteristic alteration of the endometrium identical with the progestational proliferation previously shown to be due to the presence of corpora lutea in the ovaries.” They called this substance progestin (progesterone) {Corner, 1929 #3380}.


Barnet Woolf (GB) was the first to site an example of an enzyme forming a complex with two substrates; later called ternary complexes {Woolf, 1929 #5549}. He proposed a theory of enzymatic action in which the binding of substrate or substrates leads to chemical transformation of the substrate at the specific binding site of the enzyme {Woolf, 1931 #5550}.


Carl Ferdinand Cori (CZ -US) and Gerty Theresa Cori, née Radnitz (CZ -US), concluded in 1929 from work begun in 1925 that, “Formation of liver glycogen from lactic acid is thus seen to establish an important connection between the metabolism of muscle and that of the liver. Muscle glycogen becomes available as blood sugar through the intervention of the liver, and blood sugar in turn is converted into muscle glycogen. There exists therefore a complete cycle of the glucose molecule in the body, which is illustrated by the diagram: blood glucose à muscle glycogen à blood lactic acid à liver glycogen à blood glucose.

Epinephrine (adrenaline) was found to accelerate this cycle in the direction of muscle glycogen to liver glycogen and to inhibit it in the direction of blood glucose to muscle glycogen; the result is an accumulation of sugar in the blood. Insulin, on the other hand, was found to accelerate the cycle in the direction of blood glucose to muscle glycogen, which leads to hypoglycemia and secondarily to a depletion of the glycogen stores of the liver … There is also a possibility that other hormones besides epinephrine [adrenaline] and insulin influence this cycle.” Earl W. Sutherland, Jr. (US), and Sidney P. Colowick (US) later helped refine the concept {Cori, 1925e #23145;Cori, 1927 #23146;Cori, 1929 #2593;Cori, 1931a #13159;Cori, 1941 #10709;Sutherland, 1941 #13160;Cori, 1945-1946 #13161;Cori, 1931b #23147}. Note: This circulation of carbohydrate material in the animal organism has since been called the Cori cycle.


Carl Peter Henrik Dam (DK) discovered vitamin K while working with chickens on synthetic diets. It seemed to be necessary for normal blood clotting so he named it vitamin K, for koagulation (the German spelling). Dam, Fritz Schönheyder (DK), and Erik Tage-Hansen (DK) discovered that the blood of chickens became depleted of prothrombin when they were placed on a vitamin K deficient diet {Dam, 1929 #1975;Dam, 1933 #3075;Dam, 1934a #3076;Dam, 1934b #3077;Dam, 1935a #23362;Dam, 1935b #3078;Dam, 1936 #3079}.

Sidney Allen Thayer (US), Donald W. MacCorquodale (US), Stephen B. Binkley (US), and Edward Adelbert Doisy (US) crystallized vitamin K {Thayer, 1938 #3085}.

Donald W. MacCorquodale (US), Lee C. Cheney (US), Stephen B. Binkley (US), Walter F. Holcomb (US), Ralph W. McKee (US), Sidney Allen Thayer (US), and Edward Adelbert Doisy (US) determined the constitution of vitamin K and synthesized it {Binkley, 1939 #23382;MacCorquodale, 1939a #14734;MacCorquodale, 1939b #12247;Thayer, 1939a #14732;Thayer, 1939b #14733;Thayer, 1940 #16186}.


Ernst Franz Moro (AT-DE) introduced the use of raw apples, as well as, a carrot soup in the treatment of diarrheic conditions in infants {Moro, 1929 #19089}. Note: This was based on a custom among German peasants.

Tobias L. Birnberg (US) reported the successful treatment of diarrhea, dysentery, colitis, and celiac disease in children by restricting their diet to raw apple {Birnberg, 1933 #2899}.


Michael Heidelberger (US) and Forrest E. Kendall (US) established the principle of quantitative immunochemistry when they perfected the quantitative precipitin reaction {Heidelberger, 1929 #7306}.


William Hay Taliaferro (US) wrote The Immunology of Parasitic Infections, a pioneering book in this branch of immunology {Taliaferro, 1929 #10836}.


Theodosius Grigorievich Dobzhansky (UA-US), Theophilus Shickel Painter (US), and Hermann Joseph Muller, Jr. (US) showed that while the linear sequence of genes is the same for genetic and cytological maps, physical distances and crossover map distances did not coincide {Dobzhansky, 1929a #5026;Dobzhansky, 1929b #5027;Painter, 1933b #5025;Muller, 1929 #5028}.


Frank Macfarlane Burnet (AU) and Margot McKie (AU) attributed the permanency of the lysogenic character in bacteria to the presence of an entity they called anlage which is capable of liberating bacteriophage. Anlage was conceived as a normal hereditary constituent of lysogenic bacteria and, unless activated, no liberation of bacteriophage resulted {Burnet, 1929 #5720}.


C. Eugene Woodruff (US) and Ernest William Goodpasture (US) supplied the first direct evidence of the relation of viral inclusion bodies and elementary bodies to virus {Goodpasture, 1929-30 #2322;Woodruff, 1929 #5725}.


Francis O. Holmes (US) arrived at the first practical and accurate method for the quantification of plant virus infectivity by counting the local lesions developing in the leaves of tobacco (Nicotiana species) {Holmes, 1929 #5723}.


Howard Bancroft Andervont (US) showed that the Herpes virus of man could be transmitted to mice by intracerebral inoculation, and thereby provided the means of studying this human virus in an animal {Andervont, 1929 #21851}.


Aage Nyfeldt (DK) reported that Listeria monocytogenes is capable of causing an infection in man (listeriosis) although it had been known to be infectious for domestic and feral animals since 1911. The organism derives its name from the striking monocytic blood reaction it causes in the infected host {Nyfeldt, 1929 #3159}.


Clifford Dobell (GB) and Ann Bishop (GB) described the life cycle of Entamoeba histolytica {Dobell, 1929 #6415}.


Warren Harmon Lewis (US) and Margaret Reed Lewis (US) were the first to develop time-lapse microscopic motion pictures to record observations on living cells in culture. Their films became important teaching resources in cytology and led them to develop mechanical theories of cell motion. In one of their first films they were able to see the early development of rabbit embryos, from the first ovum cleavage to the blastocyst stage.


Max Hartmann (DE) and Bjorn Føyn (NO) discovered alternation of isomorphic generations in the green algae {Hartmann, 1929 #5107;Føyn, 1929 #5108}.

E. Juller () discovered alternation of heteromorphic generations in the green algae {Juller, 1937 #5109}.


Alfred Henry Sturtevant (US) was the first to use genetic mosaics (Drosophila simulans) for developmental studies {Sturtevant, 1929 #3189}.


Eli Kennedy Marshall, Jr. (US) described the aglomerular kidney of the toadfish (Opsanus tau) {Marshall, 1929 #14488}.


Homer William Smith (US) became the first researcher to report that the gills are the major sites of nitrogen excretion in freshwater fish. His experimental subjects were the common carp (Cyprinus carpio) and goldfish (Carassius auratus) {Smith, 1929 #21194}.


Sybil Cooper (GB), Charles Scott Sherrington (GB), and Derek Ernest Denny-Brown (NZ-GB-US) observed and defined the distinctive properties of red and white muscles {Cooper, 1926 #12294;Denny-Brown, 1929a #13183}.


Derek Ernest Denny-Brown (NZ-GB-US) developed the technique of antidromic stimulation for the analysis of motor neuron responses {Denny-Brown, 1929b #18457}.

Edgar Douglas Adrian (GB-US) and Detlev Wulf Bronk (US) developed an electromyograph. Bronk’s invention of the coaxial needle electrode greatly enhanced electromyography {Adrian, 1929 #19067}.

Derek Ernest Denny-Brown (NZ-GB-US) introduced electromyography (EMG) as a clinical tool and initiated the procedure of muscle biopsy as a means of seeking a direct tissue diagnosis of neuromuscular diseases {Denny-Brown, 1949b #18456}.

Hans Piper (DE) authored the first extensive clinical EMG study {Piper, 1912 #18470}.


George Bernays Wislocki (US) and James Peter Hill (GB) determined that all the primates, except the lemurs, have discoidal or doubly discoidal placentas of hemochorial type. In Old World forms, the shape of the placenta in the Ceropithecidae (macaques, langurs, etc.) is usually doubly discoidal; the baboons, in which it is singly discoidal, are the exception. In the Hylobatidae, Pongidae, and Hominidae (anthropoid apes and man) it is always singly discoidal. A completely villous discoidal placenta is seen only in the gorilla, orangutan, and man {Wislocki, 1929 #10649;Hill, 1932 #10650}. At this point in history the placenta of the chimpanzee had not received a thorough microscopic study.


Moriz Oppenheim (AT) and Erich Urbach (AT-US) described a new metabolic skin condition: necrobiosis lipoidica diabeticorum {Oppenheim, 1929 #13349;Oppenheim, 1932 #13350;Urbach, 1932 #11848}.


Heinrich Pette (DE) proposed that inflammatory diseases of the nervous system be divided into two groups: (1) acute inflammatory diseases predominantly of the gray matter, and (2) acute inflammatory diseases predominantly of the white matter {Pette, 1929 #8079}.


Fuller Albright (US), Walter Bauer (US), Marian Ropes (US), and Joseph C. Aub (US) found that an increased phosphorus excretion is the primary effect of parathormone {Albright, 1929 #7813}.


Maxwell Myer Wintrobe (CA-US) invented what became known as the Wintrobe hematocrit {Wintrobe, 1929 #10779}.


Donald Macomber (US) and Morris B. Sanders (US) reported their results of analyzing the spermatozoa count to determine its value in the diagnosis, prognosis and treatment of sterility. They reported the normal sperm density to be 100 million sperm/mL. Their number was based on the sperm counts of 294 individuals without regard to fertility status. In addition, they reported that men with sperm densities less than 60 million/mL rarely can initiate a pregnancy {Macomber, 1929 #11536}.


Philip Duryeé McMaster (US) and Douglas R. Drury (US) revealed that the liver is the source of blood fibrinogen {McMaster, 1929 #10891;Drury, 1929 #10892}.


Edgar Otto Conrad von Gierke (DE) described a common member of a group of hereditary glycogen-storage diseases. This progressive disease is an inborn error of glycogen metabolism due to glucose-6-phosphatase (G6P) deficiency, involving chiefly the liver and kidneys {von Gierke, 1929 #13275}. The liver may become huge and contain as much as 15 percent of glycogen. It is sometimes called Gierke's disease.


Luis Morquio (UY) and James Frederick Brailsford (GB) simultaneously and independently described a disease leading to dwarfism, waddle gait and deafness, but not mental retardation. It was later called Morquio syndrome (also referred to as mucopolysaccharidosis IV, MPS IV, Morquio-Brailsford syndrome, or Morquio) {Morquio, 1929 #22630;Brailsford, 1929 #22631}.


Otto Hermann Krayer (DE-US) mastered the heart-lung-preparation (HLP) originally developed by Starling in England. He used the HLP to show that an oxidation product of Neosalvarsan is toxic to vascular beds in several organs {Krayer, 1929 #10789;Krayer, 1930a #10790}. This is one of the reasons that this anti-syphilitic arsenical drug ceased to be used.

Otto Hermann Krayer (DE-US) perfected the management of the heart-lung-preparation (HLP) to the point that he could make quantitative measurements of the activity of cardioactive drugs. He also developed a standard HLP procedure to study drug effects on the failing heart {Krayer, 1930b #10791;Krayer, 1931a #10792;Krayer, 1931b #10793;Krayer, 1931c #10794;Krayer, 1932a #10795;Krayer, 1932b #10796;Krayer, 1933 #10797}.

Wilhelm Siegmund Feldberg (DE-GB) and Otto Hermann Krayer (DE-US) used both intact dogs and cats, as well as, dog and cat HLPs to show that an “acetylcholine-like substance” is released into the coronary circulation of mammals upon electrical stimulus of the vagus nerve {Feldberg, 1933 #10799}.

Albert Wollenberger (DE) and Otto Hermann Krayer (DE-US) demonstrated a method for quantitatively determining the limits of cardiac sufficiency in response to specific measured changes in right arterial pressure. They developed a specific “competence index” to express the heart’s response numerically. This method allows a clear distinction to be made between drugs that primarily affect heart rate and those (like digitalis) that truly improve the work capacity of the impaired muscle {Wollenberger, 1948 #10798}.


Herbert Spencer Gasser (GB) and Joseph Erlanger (GB) found that within the A fiber group of nerves of the frog and the dog, cocaine blocks fibers of smaller diameter and slower conduction rate before larger fibers; but as they found that ‘blocking of some of the fast fibers occurs before the slow ones’, they were unable to formulate a satisfactory theory to account for all the facts they observed {Gasser, 1929 #23256}. Note: Type A fibers are thickest and fastest conducting, myelinated, with a diameter of 1.5-20 microns, and 4-120 m/sec conduction of impulse.


Philip Drinker (US) and Charles F. McKhann (US) invented a new negative pressure ventilator apparatus (the iron lung) for the administration of artificial respiration over prolonged periods of time {Drinker, 1929 #7908}. Note: This apparatus was invented primarily to aid the thousands of children with polio (infantile paralysis).


Walter Edward Dandy (US) introduced the practice of removing an intervertebral disk to alleviate lower back pain, sciatica, and other symptoms caused by a ruptured disk {Dandy, 1929 #12218}.


Harold Brunn (US) reported six lobectomies (removals of lung lobes) for bronchiectasis with only one death. In bronchiectasis one or more bronchi or bronchioles are chronically dilated and inflamed, with copious discharge of mucus mixed with pus. The secret of Brunn's success was the use of intermittent suction after surgery to keep the cavity free of secretions until the remaining lobes of the lung could expand to fill the space {Brunn, 1929 #14736}.


Jacques Forestier (FR) introduced gold therapy for rheumatoid arthritis {Forestier, 1929 #13252}.


Frederic Edward Clements (US), John Ernst Weaver (US), and Herbert C. Hanson (US) stated that one of the important processes directing plant succession is competition between similar plants leading ultimately to a climax community {Clements, 1929 #8294}.


Kurt van Neergaard (CH) identified the function of the pulmonary surfactant in increasing the compliance of the lungs by reducing surface tension {von Neergaard, 1929 #24044}.

Mary Ellen Avery (US) and Jere Mead (US) found that in lung extracts of immature infants and infants dying with hyaline membrane disease, surface tension is higher than expected. This deficiency of surface-active material may be significant in the pathogenesis of hyaline membrane disease (idiopathic respiratory distress syndrome of the newborn) {Avery, 1959 #24043}. Note: Pulmonary surfactant is a surface-active lipoprotein complex (phospholipoprotein) formed by type II alveolar cells.


Percival Bailey (US) and Paul Clancy Bucy (US) were the first to confirm the existence of tumors that were, in fact, composed of oligodendroglia and to establish this type of glioma as a definite entity {Bailey, 1929 #16887}.


Sergei Sergeevich Brukhonenko (RU) demonstrated successful total body perfusion after removal of animal hearts {Brukhonenko, 1929 #11273}.

John Heysham Gibbon, Jr. (US) was the first to attempt a bypass of the heart using an external circulation. Partially replacing the circulation between the heart and the lungs, he managed to keep a cat alive in this way for four hours. Two years later, under sterile conditions, 3 out of 13 cats survived for more than 250 days following similar procedures, with the remaining animals living for between 1 and 23 days {Gibbon, 1937 #10213;Gibbon, 1939 #10214}. See, J.J.C. LeGallois, 1813, M. von Frey, 1885.

Clarence Dennis (US), Dwight S. Spreng, Jr. (US), George E. Nelson (US), Karl E. Karlson (US), Russell M. Nelson (US), John V. Thomas (US), Walter Phillip Eder (US), and Richard L. Varco (US) reported the first case of open-heart surgery with machine-supported circulation and oxygenation. It was performed on April 1, 1951. This was apparently the first occasion for such an attempt to be made anywhere. The patient was a 4-year old child with a known inter-atrial septal defect who was previously operated upon with hope of repair by closed technique. When an attempt was made to take the patient off perfusion, the patient’s heart could not maintain the circulation, and the patient expired. Postmortem examination revealed that the lesion was not the simple secundum type of defect but rather an atrioventricular canal, a complicated set of anomalies {Dennis, 1951 #11341}.

Forest Dewey Dodrill (US), Edward Hill (US), and Robert A. Gerisch (US) performed the first clinically successful total left-sided heart bypass in a human on July 3, 1952. The machine was used to substitute for the left ventricle for 50 minutes while a surgical procedure was carried out to repair the mitral valve; the patient's own lungs were used to oxygenate the blood {Dodrill, 1952a #14520;Dodrill, 1952b #14521}.

John Heysham Gibbon, Jr. (US) successfully applied extracorporeal circulation in an 18-year-old female with an atrial septal defect. Unfortunately, he was unable to repeat this success in other humans {Gibbon, 1953 #11323}.

John Heysham Gibbon, Jr. (US), Arthur R.C. Dobell (US), and George B. Voigt (US) reported the closure of interventicular septal defects on dogs during open cardiotomy with the maintenance of the cardio-respiratory functions by a pump oxygenator {Gibbon, 1954 #11274}.


Karl S. Lashley (US) promulgated the theory of cortical specialization for sensory and motor functions. He challenged the ongoing concept of cortical localization. Lashley brought the controversy between localization and holistic emphasis of brain function into focus. He is remembered as a great psychologist who approached learning and memory by assessing the effects of brain damage in laboratory animals {Lashley, 1929 #17614;Lashley, 1930 #14606}.


Wolfgang Köhler (DE) and Kurt Koffka (DE) promoted a pattern theory of memory. Diffuse neural groupings mediated memory. They hypothesized that multiple memory traces were formed in the cortex. Visual memories involved successive and simultaneous stimuli in different parts of the visual field. They postulated that new records might be inscribed on top of old patterns these could affect one another leading to a newly organized group-unit {Koffka, 1935 #17754;Köhler, 1929 #17753}.


Friedrich von Lucanus (DE) published Zugvögel und Vogelzug (Migratory Birds and Bird Migration) {von Lucanus, 1929 #26643}.

Ernst Schüz (DE) and Hugo Weigold (DE) published the first atlas of bird migration {Schüz, 1931 #26642}.


Johanna Gabrielle Ottilie Edinger (DE-US) demonstrated that the evolution of the brain could be studied directly from fossil cranial casts {Edinger, 1929 #8972}. She later showed that the progression of brain structure does not proceed at a constant rate in a given family but varies over time; also, that the enlarged forebrain evolved several times independently among advanced groups of mammals and there was no single evolutionary scale {Edinger, 1948 #8973}.



“The most beautiful thing we can experience is the mysterious. It is the source of all art and science,” Albert Einstein {Einstein, 1930 #4529;Maslow, 1970 #7766}.


“The scientific attitude of mind involves a sweeping away of all other desires in the interest of the desire to know—it involves suppression of hopes and fears, loves and hates, and the whole subjective emotional life, until we become subdued to the material, without bias, without any wish except to see it as it is, and without any belief that what it is must be determined by some relation, positive or negative, to what we should like it to be or to what we can easily imagine it to be.” Bertrand Russell {Russell, 1930 #6366}.


“No myth of miraculous creation is so marvelous as the fact of man’s evolution.” Robert Briffault {Briffault, 1930 #15303}.


“The truly scientific mind is altogether unafraid of the new, and while having no mercy for ideas which have served their turn or shown their uselessness, it will not grudge to any unfamiliar conception its moment of full and friendly attention, hoping to expand rather than to minimize what small core of usefulness it may happen to contain.” Wilfred Batten Lewis Trotter {Trotter, 1930 #22827}.


“Any scientific problem must be attacked by research into detail; the natural scientist did not win his victories until he left meditation on the great riddles of the world and began a careful study of special problems; our knowledge - of more general associations and of far-reaching laws has grown out of the results of such research.” Hans Karl August Simon von Euler-Chelpin, Nobel Lecture 1930.


Hans Fischer (DE) was awarded the Nobel Prize in Chemistry for research into the constitution of hemin and chlorophyll and especially for the synthesis of hemin.


Karl Landsteiner (AT-US) was awarded the Nobel Prize in Physiology or Medicine for his discovery of human blood groups.


A.A. Lebedeff (RU) designed and built the first interference microscope {Lebedeff, 1930 #14609}.


Alessandro Vallebona (IT) constructed equipment and published the first clinical body-section imaging material ever in 1930{Vallebona, 1930 #27907;Vallebona, 1933 #27908}.

Bernhard Ziedses des Plantes (NL) devised the first ideal planigraphic method for clinical material in 1932 {Plantes, 1932 #27909}.

William Watson (GB) in 1937 {Watson, 1937-1939 #27911}, Jean Kieffer (US) in 1938 {Kieffer, 1938 #27910}, and Shinji Takahashi in 1947 {Takahashi, 1969 #27912} independently described methods for transverse axial tomography . Note: The limitation in sensitivity of the photographic medium was revealed in parallel to this development. See, Oldendorf, 1961


Sterling Brown Hendricks (US) and William H. Fry (US) presented what is arguably the most important elucidation of the nature and properties of soils. They conclusively proved the crystalline nature of colloidal clay with the prevalence of negative charges that will absorb and release cations. These findings led to an understanding of the chemistry necessary to maintain high potential in soil productivity, and in providing a valid chemical basis for the reclamation of the alkali soils of arid regions {Hendricks, 1930 #9960}.


William Thomas Astbury (GB), Henry J. Woods (GB), A. Street (GB), William Henry Bragg (GB) and using x-ray diffraction, demonstrated for the first time a measurable change in protein structure at the most intimate molecular level—interatomic shifts of a few ångströms. This change was found to be reversible. They called the two forms alpha-keratin and beta-keratin {Astbury, 1930 #19754;Astbury, 1931 #19755;Astbury, 1933 #19756}.


Hermann Bortels (DE) reported that nitrogen fixation by Azotobacter has a requirement for molybdenum {Bortels, 1930 #16811}.


Rudolph John Anderson (US) reported myoinositol as a lipid constituent in the phospholipids of mycobacteria {Anderson, 1930 #16131}.


Otto Heinrich Warburg (DE), Otto Fritz Meyerhof (DE-US), Einar Lundsgaard (DK) and Hermann Karl Felix Blaschko (DE-GB) demonstrated that ATP is generated from ADP in coupled enzymatic reactions during the anaerobic breakdown of glucose to lactic acid in muscle {Meyerhof, 1930a #774;Warburg, 1939b #121}.


Edward Adelbert Doisy (US), Sidney Allen Thayer (US) and Clement D. Veler (US) crystallized the ovarian hormone, estrogen, which induces estrus from the urine of pregnant women {Doisy, 1930 #12296}. Note: probably 17-beta estradiol.


Hans Karl August Simon von Euler-Chelpin (DE-SE), Karl Zeile (SE), and Harry Hellström (SE) showed that catalase contains a hemin residue {von Euler-Chelpin, 1930 #9135}.


John Tileston Edsall (US) and Alexander L. von Muralt (US) isolated myosin from muscle {Edsall, 1930 #11242;von Muralt, 1930a #11243;von Muralt, 1930b #11244}.


Wilbur Willis Swingle (US), Joseph J. Pfiffner (US), Frank A. Hartman (US), and Katherine A. Brownell (US) were the first to prepare extracts from the adrenal cortex, which successfully controlled the symptoms of adrenal insufficiency both in adrenalectomized animals and in patients who had Addison’s disease {Hartman, 1930 #9122;Swingle, 1930 #9123;Swingle, 1931 #16251}.

Leonard George Rowntree (CA-US), Carl H. Greene (US), Wilbur Willis Swingle (US), and Joseph J. Pfiffner (US) proved the medical efficacy of the Swingle-Pfiffner extract {Rowntree, 1930 #19953}.


C.L. Ruiz (AR), L.L. Silva (AR), and L. Libenson (AR) were the first to report the hypoglycemic effect of some sulphonamide-like compounds {Ruiz, 1930 #3421}.


William Smith Tillett (US), Walther Frederick (US), and Oswald Theodore Avery (US) discovered C-polysaccharide (the C-fraction), a non-capsular antigenic component of pneumococci {Tillett, 1930a #10408}.

William Smith Tillett (US) and Thomas Francis, Jr. (US) discovered that the C-fraction carbohydrate from pneumococci stimulates the production of a non-antibody globulin, which they dubbed C-reactive protein (CRP) {Tillett, 1930b #10409}.

Subsequently it was found that C-reactive protein appears in the blood of patients in response to many infections. During recovery from infection the C-reactive protein (CRP) diminishes in amount and within a few days disappears entirely. The serum of an animal immunized to CRP is used in a Precipitation Test to detect CRP in sera of persons suspected of having one of the diseases in which the protein appears, e.g. staphylococcal infection.

Colin Munro MacLeod (US) and Oswald Theodore Avery (CA-US) purified C-reactive protein (CRP) {Macleod, 1941 #25493}.

Maclyn McCarty (US) crystallized C-reactive protein (CRP) {McCarty, 1947 #25494}.


 Vladimir Aleksandrovich Engelhardt (RU) recognized that nucleated erythrocytes catalyze an aerobic process linked to phosphorylation, i.e., oxidative phosphorylation. This experiment initiated the history of oxidative phosphorylation {Engelhardt, 1930 #9274;Engelhardt, 1932 #105}.


Ejnar Lundsgaard (DK) discovered that frog muscle poisoned with iodoacetic acid—which inhibits glycolysis—can contract without the formation of lactic acid, but with the disappearance of creatine phosphate. Once the creatine phosphate was exhausted the muscles went into a rigor mortis like condition. He wrote, “… phosphagen (creatine phosphate) is the substance directly supplying the energy for contraction, while lactic acid formation in the normal muscle continually provides the energy for its resynthesis” {Lundsgaard, 1930 #2561}. Lundsgaard had discovered that the muscle machine can be driven by phosphate-bond energy.


Ragnar S. Nilsson (SE) isolated phosphoglyceric acid from natural sources. Based on the action of dried yeast on a mixture of glucose, hexosephosphate, and acetylaldehyde he suggested that glyceraldehyde-phosphate might play a role in the glycolytic breakdown of carbohydrate {Nilsson, 1930 #1718}.


Friedrich Breinl (CZ) and Felix Haurowitz (CZ-US) published their template theory of antibody formation {Breinl, 1930 #2156}. Although incorrect, this theory and others stimulated research.

Gustav Joseph Victor Nossal (AU), Gordon Leslie Ada (AU), Caroline M. Austin (AU), John Pye (AU), and Gail M. Williams (AU) demonstrated that antibody making cells do not contain any antigen around which to shape an antibody {Nossal, 1965a #4363;Nossal, 1965b #4364;Nossal, 1965c #4365;Nossal, 1967 #4366}.


H. Lyndhurst Duke (GB) and James Montague Wallace (GB) gave the first description of a complement receptor activity on erythrocytes {Duke, 1930 #8257}.

James Montague Wallace (GB) and Arthur Wormall (GB) proposed that complement is required for this adherence reaction {Wallace, 1931 #8258}.


Haldan Keffer Hartline (US) and Clarence H. Graham (US) used tiny electrodes to determine how single retinal nerve cells in horseshoe crabs and frogs receive information and transfer it to the brain {Hartline, 1930 #8204}.

Haldan Keffer Hartline (US) mapped the activity of the visual receptive field to reveal a system of many convergent pathways from many photoreceptors {Hartline, 1940 #10767}. This work laid the foundation for modern concepts of parallel processing by specialized channels.


Yandell Henderson (US), Howard W. Haggard (US), Pol N. Coryllos (GR-US) and George L. Birnbaum (US) found that in dogs with experimentally induced pneumonia the lungs may be cleared, and the pneumonia cured by placing the animals in an atmosphere of about 8% carbon dioxide for 12 to 24 hours. In support of the claim that these are real cures is the fact that pneumococci are inhibited in growth or even killed by a lowering of pH no greater than carbon dioxide may induce. A lowering of the pH by carbon dioxide contributes also to the autolysis and liquefaction of the exudate responsible for the consolidation of the lungs in pneumonia. Many cases of pneumonia have now been treated by inhalation of a carbon dioxide-oxygen mixture in a special tent introduced by Henderson and Haggard. This treatment is decidedly superior to treatment with oxygen alone {Henderson, 1930 #11118}.


Ernest Glen Wever (US) and Charles W. Bray (US) discovered the cochlear microphonic potential, i.e., the bioelectric signals generated in the inner ear in response to sound stimuli {Wever, 1930a #11074;Wever, 1930b #11075;Wever, 1930c #6597;Wever, 1930d #6598}.


Benjamin Freeman Kingsbury (US) in his studies of neuroembryonic development provided the first detailed discussion of the possible functions of the floor plate in neural development {Kingsbury, 1930 #6300}.


Ronald Aylmer Fisher (GB-AU) wrote The Genetical Theory of Natural Selection. This book contained his fundamental theorum of natural selection: “The rate of increase in fitness of any organism at any time is equal to its genetic variance in fitness at that time.” Natural selection is viewed as always tending to increase fitness, in the sense of reproductive fitness, and the course of evolution being determined by the momentary advantage of one allele over the other. This work formalized the relation of particulate genes to the evolutionary process. It overturned the old theory of heredity as blending, on which Charles Robert Darwin (GB) had based his views of the operation of natural selection {Fisher, 1930 #4833}. Note: Fisher borrowed from the works of Alfred James Lotka (GB-US) without acknowledgment. See, Lotka, 1920, 1925, and 1956


Lewis John Stadler (US) devised and perfected methods for determining rate of spontaneous mutation in maize, finding that different genes mutate at widely different rates {Stadler, 1930 #12299;Stadler, 1932 #12300;Stadler, 1951 #12298}.

Norman H. Giles, Jr. (US) found that in Neurospora crassa quantitative mutability of different loci varies in spontaneous mutation {Giles, 1951 #16197}.

Milislav L. Demerec (Yugoslavian -US), Zlata Hartman (US), Philip Emile Hartman (US), Takashi Yura (JP), Joseph S. Gots (US), Haruo Ozeki (JP), and Stuart W. Glover (GB) found that in bacteria, quantitative mutability of different loci varies in spontaneous mutation {Demerec, 1956a #16198}.


Herbert Friedmann (US) studied the honey-guides—family Indicaroridae—of Southern Rhodesia (Zimbabwe) and South Africa. He confirmed, first hand, that these birds do indeed lead a symbiont, often man, who opens the hive and removes some of the honey, the bird, then feeds on the wax comb {Friedmann, 1930 #10696;Friedmann, 1943 #10690;Friedmann, 1954 #10691;Friedmann, 1955 #10692;Friedmann, 1958 #10695}.

Herbert Friedmann (US) and Jerome Kern (US) determined that honey-guides possess a digestive enzyme and a microbial symbiont both of which attack the beeswax, thus making it available for further digestion {Friedmann, 1956a #10693;Friedmann, 1956b #10694}. Wax eating is called cerophagy.


Charles Benedict Davenport (US) published a short pedigree, which demonstrates linkage between hemophilia and red-green color-blindness in man {Davenport, 1930 #22133}.


Julia Marchal (GB) discovered the mousepox (Poxvirus muris) virus and called it infectious ectromelia {Marchal, 1930 #7016}. Marchal bodies are cell inclusion bodies observed in infectious ectromelia.


Wilbur A. Sawyer (US), Stuart F. Kitchen (US), Martin M. Frobisher, Jr. (US), and Wray Lloyd (US) determined the relationship of yellow fever of the Western hemisphere to that of Africa and leptospiral jaundice {Sawyer, 1930 #12317}.


Charles Cyril Okell (GB) and Adelaide V. Blake (GB) determined that the Shiga exotoxin is released following autolysis or disruption of the bacterial cells; suggesting that it is an endotoxin {Okell, 1930 #1826}.


Bert Cyril James Gabriel Knight (GB) and Paul Gordon Fildes (GB) initiated studies, which led to the understanding that some heterotrophic bacteria can grow on a simple medium containing inorganic ions, ammonium as nitrogen source and glucose as source of carbon and energy. Other species are unable to grow on such a medium unless it is supplemented by an amino acid or one of the B group of vitamins. Still others, particularly those found in chemically complex habitats, require several amino acids and several vitamins before they could grow. Fildes and his colleagues postulated that the nutritional requirements reflect the synthetic disabilities of the organisms concerned: that an organism that requires a specific amino acid for growth has, during its evolution, lost the ability to synthesize that amino acid which is nevertheless an essential part of its cellular material. This postulate meant that nutritional studies could be used to study stages in biosynthesis {Knight, 1930 #15789}.


Max Theiler (ZA-US) and Hugh H. Smith (US) grew the yellow fever virus in rhesus monkeys (Macacus rhesus), and then passed it to mice. In mice, it developed as encephalitis. They passed it from mouse to mouse, and then eventually back to monkeys. By this time, it had attenuated, producing a very mild attack in the monkeys, but producing a full immunity to the most virulent form of the virus. This attenuated strain could also be used to vaccinate man {Theiler, 1930a #2367;Theiler, 1930b #12186}.

Max Theiler (ZA-US), Hugh Hollingsworth Smith (US), Henrique A. Penna (BR), and Adhemar Paoliello (BR) carried out successful field trials of their yellow fever vaccine {Theiler, 1937a #8818;Smith, 1938 #8819}.


Gail Monroe Dack (US), William E. Cary (US), Oram C. Woolpert (US), and Hazel Wiggers (US) ingested filtered supernatant fractions of Staphylococcus aureus cultures to demonstrate that it can produce an extracellular enterotoxin, which causes gastroenteritis {Dack, 1930 #15790}.


Samuel Phillips Bedson (GB), George T. Western (GB), Samuel Levy Simpson (GB), and John O.W. Bland (GB) were outstanding early students of the Bedsonia —later called Chlamydiae. They discovered that Chlamydia psittaci/Chlamydophila psittaci is the etiological agent of parrot fever (psittacosis) {Bedson, 1930 #12333;Bedson, 1932 #13110}.

Walter Levinthal (DE), Ralph Dougall Lillie (US), and Alfred C. Coles (GB) independently discovered the etiological agent of psittacosis {Levinthal, 1930a #13364;Levinthal, 1930b #13365;Lillie, 1930 #13366;Coles, 1930 #13367}.


M. Ruiz Castañeda (MX) developed an excellent method for staining rickettsia {Castañeda, 1930 #19954}.

Attilio Macchiavello (CL) developed an excellent differential stain for observation of Rickettsia and Chlamydia {Zinsser, 1939 #22453}. Note: Macchiavello did not independently publish his staining procedure.


Bernard Schlesinger (GB) showed that hemolytic streoptococcal infection could cause acute rheumatism in children {Schlesinger, 1930 #13492}.


Homer William Smith (US) determined that for marine teleosts to maintain their blood salinity below that of the surrounding waters they rely upon a relatively impermeable skin, drink sea water to replace water lost through the gill membranes, and secret salt using a special transport mechanism located in the gills. Elasmobranch fishes solve this problem by making their tissues isosmotic or slightly hyperosmotic to seawater using a build-up of urea. He found that the elasmobranch kidney conserves urea by tubular reabsorption {Smith, 1930a #5832;Smith, 1932 #5833;Smith, 1936 #5835}.

Malcolm S. Gordon (US), Knut Schmidt-Nielsen (DK-US), and Hamilton M. Kelly (US) found that the crab-eating frog (Rana cancrivora) is the only known amphibian, which can tolerate seawater. It does so by raising its blood osmolarity using urea {Gordon, 1961 #5834}.

Homer William Smith (US) found that to prevent desiccation, lungfishes burrow deep into the mud and form a cocoon with a breathing channel to the surface. They may remain dormant for up to two years during which time course their body proteins are gradually utilized, and their tissue urea levels may reach 3 percent {Smith, 1930b #5839}.


Charles Haskell Danforth (US) established the basic conditions for understanding the mechanisms for production of sex plumage in birds. He found in Ring-necked pheasants the sex characteristics of the plumage are dependent upon simultaneous action of both genic and hormonal factors. A single gene difference leads to divergent reactions to hormone in two races, Campines and Brown leghorns {Danforth, 1930 #11195}.


Hayes E. Martin (US) and Edward B. Ellis (US) introduced the technique of fine needle biopsy {Martin, 1930 #11571}.


Ernest Laurence Kennaway (GB) and Izrael Hieger (GB) showed for the first time that single polycyclic aromatic hydrocarbons (PAHs), such as dibenz[a,h]anthracene, are tumorigenic in mouse skin {Kennaway, 1930a #14559;Kennaway, 1930b #23774}.

James W. Cook (GB), Colin L. Hewett (GB), and Izrael Hieger (GB) discovered that pure benzo[a]pyrene induces skin cancer in mice {Cook, 1933 #14560}.


Grover A. Kempf (US) and Frederick S. McKay (US) determined that high levels of water-borne fluoride cause the discoloration of tooth enamel now called fluorosis {Kempf, 1930 #15805}.

Margaret Cammack Smith (US), Edith M. Lantz (US), and Howard V. Smith (US) performed experiments, which strongly supported the idea that excessive fluoride ingestion causes the hypoplasia characteristic of mottled teeth {Smith, 1931 #3105;Smith, 1932 #23372}.

Henry Trendley Dean (US) recalled that mottled tooth enamel is unusually resistant to decay from reading studies by Frederick S. McKay and Green Vardiman Black on fluorosis in Colorado. Dean wondered whether adding fluoride to drinking water at physically and cosmetically safe levels would help fight tooth decay.

Henry Trendley Dean (US), Frederick S. McKay (US), and Elias Elvove (US) made a critical discovery. Namely, fluoride levels of up to 1.0 ppm in drinking water did not cause enamel fluorosis in most people and only mild enamel fluorosis in a small percentage of people {Dean, 1938 #15806}.

Henry Trendley Dean (US), Francis A. Arnold, Jr. (US), and Elias Elvove (US) performed a statistical survey, which confirmed that the incidence of dental caries is significantly lower in communities where the water contains fluorine above certain concentrations {Dean, 1942 #3107}.

In 1945, Grand Rapids, MI became the first city in the world to intentionally fluoridate its drinking water. During the 15-year project, researchers monitored the rate of tooth decay among Grand Rapids' almost 30,000 schoolchildren. After just 11 years, Francis A. Arnold, Jr. (US), Henry Trendley Dean (US), Philip Jay (US), and John W. Knutson (US) announced an amazing finding. The caries rate among Grand Rapids children born after fluoride was added to the water supply dropped more than 60 percent {Arnold, 1956 #15808;Dean, 1956 #15807}.


Gregor T. Popa (RO) and Una Fielding (GB) discovered the vascular link between the pituitary and the hypothalamus; the hypophyseal portal system {Popa, 1930 #26511}.


Bernardo Alberto Houssay (AR) and Alfredo Biasotti (AR) removed the anterior lobe of the pituitary gland and pancreas from animals. They showed that the anterior lobe of the pituitary gland affects the course of sugar metabolism in the human. The anterior lobe seemed to produce one hormone that had the opposite effect of insulin {Houssay, 1930 #13324;Houssay, 1931a #3404;Houssay, 1931b #13325;Houssay, 1936 #13326}. Their work explained spontaneous remission of diabetes mellitusvanishing diabetes – by a destructive lesion in, or surgical removal of, the anterior pituitary gland.

József Baló (AT-HU) was the first to describe this phenomenon (vanishing diabetes) in humans {Baló, 1924 #13327}.


Ludvig Hektoen (US) determined the infectious nature of acute endocarditis {Hektoen, 1930 #15881}.


Karl Theodore Fahr (DE) described intracerebral calcification of the small vessels of the deep cortex and lenticular and dentate nuclei {Fahr, 1930-1931 #22629}. This became known as Fahr’s disease.  


Frederick E. Becker (US) first recognized Colorado tick fever as a separate clinical entity and gave it its name. He gave the first clear description of the disease {Becker, 1930 #21725}. Colorado tick fever (CTF) (also called Mountain tick fever, American tick fever, and "American mountain tick fever") is a viral infection (Coltivirus) transmitted from the bite of an infected Rocky Mountain wood tick (Dermacentor andersoni). It should not be confused with the bacterial tick-borne infection, Rocky Mountain spotted fever. Colorado tick fever virus (CTFV) infects hemopoietic cells, particularly erythrocytes.


Donald Dexter van Slyke (US), Edgar Stillman (US), Eggert Hugo Heiberg Møller (US), William E. Ehrich (US), John F. McIntosh (US), Louis Leiter (US), Eaton M. MacKay (US), R. Roger Hannon (US), Norman S. Moore (US), and Christopher D. Johnson (US) related the changes occurring at different stages of renal deterioration to the quantitative changes taking place in kidney function {van Slyke, 1930 #10941}.


John Carew Eccles (AU) and Charles Scott Sherrington (GB) showed central inhibition of flexor reflexes {Eccles, 1930a #12306;Eccles, 1930b #12307;Eccles, 1931a #12308;Eccles, 1931b #12309;Eccles, 1931c #12310;Eccles, 1931d #12311;Eccles, 1931e #12312}.

John Carew Eccles (AU) wrote, The Physiology of Snapses which gives a comprehensive review of chemical and electrical synapses for both vertebrates and invertebrates {Eccles, 1964 #24234}.


Robert G. Green (US), Newell R. Ziegler (US), Benjamin B. Green (US), and Evelyn T. Dewey (US) discovered that epizootic fox encephalitis is caused by a virus, a.k.a., infectious canine hepatitis virus {Green, 1930 #25959}. Note: the first adenovirus


Arnold Rice Rich (US) separated jaundice into two types based on pathogenesis. The first, retention jaundice, results from the overproduction of the bile pigment in conditions that are associated with a decrease in excretory power of the liver, such as fever, anoxemia, and immaturity. The second type—regurgitation jaundice—is caused by reflux of bile from the liver canaliculi into the blood stream in the presence of duct obstruction or liver cell necrosis {Rich, 1930a #10102}.


Arnold Rice Rich (US) demonstrated that acquired resistance in the host is independent of the hypersensitive inflammatory reaction, and the latter, injurious to the host, may be eliminated by desensitization without impairment of immunity {Rich, 1930b #10106;Rich, 1941 #10104}.


James Wenceslas Papez (US) and Grant L. Rasmussen (US) determined that the superior olivary complex (SOC) exhibits a descending projection directed to the cochlea. Axons of SOC neurons build the so-called olivo-cochlear bundle (Bundle of Rasmussen) {Papez, 1930 #11634;Rasmussen, 1946 #11633}.


Alexander A. Maximow (RU-US) posthumously authored A Textbook of Histology, which was one of the most influential histology texts of the twentieth century {Maximow, 1930 #10669}. After Maximow’s death his book was completed and edited by William Bloom (US).


Anton Elschnig (AT) developed the method of corneal grafting introduced by Eugen von Hippel (DE) and produced good results on the human eye {Elschnig, 1930 #13223}.


Magnus Hirschfield (DE) was the first true sex change pioneer. He coined the word transsexualism and founded, in Berlin, the first sexology institute. It was here that the first complete male-to-female sex change operation was performed in 1930. The patient was a Danish painter, Einer Wegener-Andreas Sparre, who had his genital organs removed, ovaries transplanted into him, and attempts made to furnish him with an artificial vagina. He died because of the series of operations, but before his death his marriage was annulled by the Danish authorities who issued him a new birth certificate as a female, with the name Lili Elbe {Elbe, 1933 #14310}.

Harold Delf Gillies (NZ-GB) and Ralph Millard (GB), in 1945, performed the first woman-to-man sex change operation {Gillies, 1957 #14737}.


Chester Scott Keefer (US) described the pathologic changes in the beriberi heart as: (1) degeneration of peripheral nerves, including both the motor and sensory elements; (2) edema of the subcutaneous tissues and muscles, and effusion of fluid into the serous cavities and (3) dilatation of the heart, particularly of the right side, with fatty infiltration and a moderate degeneration of the myocardium.

These changes lead to peripheral neuritis, edema of the tissues and signs of cardiac insufficiency {Keefer, 1930 #23368}.


Louis Wolff (US), John Parkinson (GB), and Paul Dudley White (US) discovered what became known as the WPW (Wolff-Parkinson-White) Syndrome. This is a bundle-branch block with short P-R interval in healthy young people prone to paroxysmal tachycardia {Wolff, 1930 #19141}.

Agustin Castellanos, Jr. (CU), Eduardo Chapunoff (CU), Cesar Castillo (CU), Orlando Maytin (CU), and Louis Lemberg (CU) made major contributions to understanding the WPW-Syndrome {Castellanos, 1970 #19177}.


Owen S. Gibbs (CA) demonstrated an artificial heart consisting of two bellows within a round brass container {Gibbs, 1930 #18471;Gibbs, 1933 #18472}.

Tetsuzo Akutsu (JP) and Willem Johan Kolff (NL-US) reported the development of a totally artificial heart in an animal model. They implanted a totally artificial heart into a living dog that subsequently survived for 90 minutes {Akutsu, 1958 #14526}.

Lyle D. Joyce (US), Willem C. DeVries (US), W. Larry Hastings (US), Don B. Olsen (US), Robert K. Jarvik (US), and Willem Johan Kolff (NL-US) reported on the response of the human body to the first permanent implant of the Jarvik-7 Total Artificial Heart. In 1982, the heart was implanted into Barney Clark who lived 112 days following the implantation {Joyce, 1983 #4214}.


Sewall Green Wright (US) gave the principles of gene frequency fluctuations in small populations. According to what he called genetic drift—the Sewall Green Wright effect—, in small, non-randomly mating populations, gene frequencies are found to fluctuate purely by chance {Wright, 1929 #25214}.


Alfred Adler (AT) and Philip Mairet (GB) suggested that neurosis is a defect or failure in adjustment to the social environment and arises as a failure to defend the ego {Adler, 1930 #22633}.


The United States Congress passed the Plant Protection Act (PPA), which provided for the patenting of asexually reproducing varieties. In 1970, the Plant Variety Protection Act (PVPA) was extended to include breeder’s rights to sexually reproducing varieties.


Sydney Savory Buckman (GB), published dozens of papers on ammonites, named hundreds of them, and invented a new way of dating rocks by time zones called hemera, each with their characteristic ammonites. His hemeral scheme for the Jurassic Period contained 370 hemera and 47 ages, the latter roughly corresponding to Oppel's sediment Zones {Buckman, 1909-1930 #7460}.



“Concern for man himself and his fate must always be the chief interest of all technical endeavors … in order that the creations of the mind shall be a blessing and not a curse to mankind. Never forget this, in the midst of your diagrams and equations.” Albert Einstein {Einstein, 1931 #12}.


Otto Heinrich Warburg (DE) was awarded the Nobel Prize in Physiology or Medicine for his discovery of the nature and mode of action of the respiratory enzyme (Atmungsferment); a system of cytochromes and their oxidases that participate in the respiratory process; often, specifically, cytochrome oxidase.


William Joseph Elford (GB) and Christopher Howard Andrewes (GB) developed the first filters (graded collodion membranes) in which pore size could be precisely determined. They used these to determine that viruses range in size from large protein molecules to tiny bacteria {Elford, 1931 #8812;Elford, 1932 #5726}.


Linus Carl Pauling published his first essays on The Nature of the Chemical Bond, detailing the rules of covalent bonding {Pauling, 1931 #724}.


Sterling Brown Hendricks (US), William L. Hill (US), Kenneth D. Jacob (US), and Merrill E. Jefferson (US) showed the structural characteristics of apatite-like substances and composition of phosphate rock and bone as determined from microscopic and x-ray diffraction examinations {Hendricks, 1931 #9961}.


Paul Karrer (RU-CH) and Alois Helfenstein (CH) determined the chemical formula for squalene {Karrer, 1931b #12313}.


Hans Fischer (DE) and Richard Hess (DE) determined that the structure of bilirubin and strongly implied that it is derived from hemin {Fischer, 1931 #12810}.

Hans Fischer (DE) and Hans Plieninger (DE) accomplished the complete synthesis of bilirubin {Fischer, 1942 #18989}.


Ulf Svante Hansson von Euler-Chelpin (SE) and John H. Gaddum (GB) isolated Substance P as a tissue extract that caused intestinal contraction in vitro {von Euler-Chelpin, 1931 #22387}.

Michael M. Chang (US), Susan E. Leeman (US), and Hugh D. Niall (US) determined the eleven-amino-acid structure of the Substance P peptide {Chang, 1971 #22390}.

Norma P. Gerard (US), Levi Alexander Garraway (US), Roger L. Eddy, Jr. (US), Thomas B. Shows (US), Hideya Iijima (JP), Jean Luc Paquet (FR), and Craig Gerard (US) identified the endogenous receptor for substance P as neurokinin 1 receptor (NK1-receptor, NK1R) {Gerard, 1991 #22388}.

Jane Yip (US) and Loris A. Chahl (GB) found that Substance P and the NK1 receptor are widely distributed in the brain and are found in brain regions that are specific to regulating emotion (hypothalamus, amygdala, and the periaqueductal gray) {Yip, 2001 #22389}.


Otto Heinrich Warburg (DE) and Walter Christian (DE), and Alfred Griese (DE) found that adenine, nicotinamide, pentose, and phosphate were present in the ratio 1:1:2:3 in what was being called co-ferment or cozymase. They began referring to it as hydrogen-transporting co-ferment and stated, “The pyridine component of the co-ferment is its active group, because the catalytic action of the co-ferment depends on the alteration of the oxidation state of the pyridine part” {Warburg, 1931 #19061;Warburg, 1935a #2549;Warburg, 1935b #2550;Warburg, 1935c #2551;Warburg, 1936b #19260}. Note: Over the years their Wasserstoffübertragendes Co-ferment or hydrogen transport coenzyme has been called: codehydrogenase II (Co II); phosphocozymase; codehydrase II; codehydrogenase II; coenzyme II (Co II); triphosphopyridine nucleotide (TPN); and nicotinamide adenine dinucleotide phosphate (NADP). The location of the third phosphate was established after 1950 in the laboratory of Arthur J. Kornberg (US).


Adolf Friedrich Johann Butenandt (DE) and Kurt Tscherning (DE) isolated, crystallized, and identified androsterone (andro = male, ster = sterol, one = ketone) from male urine. They presented this discovery at a Hamburg chemical meeting on October 23, 1931. Using microanalysis techniques, they were able to learn enough about it to predict its structure and draw up the full constitutional formula {Butenandt, 1934a #14613}. Androsterone is now called testosterone.

Karoly Gyula David (NL), Elizabeth Dingemanse (NL), Janos Freud (NL) and Ernst Laqueur (NL) crystallized male hormone from testicles, coining the name testosterone for their newly identified hormone (testo = testes, ster via sterol, one = ketone) {David, 1935 #9150}.

Adolf Friedrich Johann Butenandt (DE), Günter Hanish (DE), Leopold Stefan Ruzicka (HR-CH), and Albert Wettstein (CH) partly synthesized the sex hormone testosterone from cholesterol {Butenandt, 1935a #12315;Butenandt, 1935b #9153;Ruzicka, 1935a #9149;Ruzicka, 1935b #9151;Ruzicka, 1935 #9152}. Note: This was the first synthesis of a sex hormone and the first proof of the relationship between cholesterol and sex hormones.

John McLean Morris (US), reported a series of 82 individuals (80 cases collated from the literature and two cases of his own) who had a female phenotype despite the presence of bilateral testes. He referred to this sndrome as "testicular feminization" {Morris, 1953 #13417}. Note: Since his initial description, studies of the endocrinology, pathophysiology, biochemistry, and molecular biology of the androgen insensitivity syndrome (AIS) have provided insights into the role of androgens in male sex differentiation, the mechanisms of androgen action, and aspects of the structure/function relationships of the androgen receptor. AIS is an archetypal example of a hormone resistance disorder. Androgens are secreted by the testes of these 46,XY individuals in normal or increased amounts; however, due to defective androgen receptor (AR) function, there is loss of target organ response to the hormone, and the effects of androgens are reduced or absent . comprehensive description of the clinical manifestations of Note: Most clinicians have abandoned the term ‘‘testicular feminization’’ and adopted the term ‘‘complete androgen insensitivity syndrome’’ (CAIS).

Arun K. Roy (US), Yan Lavrovsky (US), Chung S. Song (US), Shuo Chen (US), Myeong H. Jung (US), Nadarajan K. Velu (US), Baoyuan Bi (US), and Bandana Chatterjee (US) show that the androgen receptor (AR), also known as NR3C4 (nuclear receptor subfamily 3, group C, member 4), is a type of nuclear receptor that is activated by binding any of the androgenic hormones, including testosterone and dihydrotestosterone in the cytoplasm and then translocating into the nucleus {Roy, 1998 #27198}.


Joseph L. Svirbely (US), Frederick L. Smith (US) and Charles Glen King (US) isolated vitamin C and reported its anti-scorbutic activity just two weeks ahead of Albert Imre Szent-Györgyi (HU-US) {Svirbely, 1931 #4843;Smith, 1931 #4844}.


Gertrude Maud Robinson (GB) and Robert Robinson (GB) were among the first to isolate, separate, and identify the anthocyanin and anthoxanthin plant pigments {Robinson, 1931 #1624;Robinson, 1932 #1625;Robinson, 1934 #1626}.


Phoebus Aaron Theodor Levene (US) and Lawrence Bass (US) introduced the chemical components and the basic structure of DNA {Levene, 1931 #14666}.


Ernst Friedheim (CH-US) and Leonor Michaelis (DE-US) found that at acidic pH values, the oxidation-reduction of the natural pigment pyocyanine involves the step-wise transfer of one electron, with the intermediate formation of a free radical, which Michaelis termed a semiquinone {Friedheim, 1931 #19799}.

Bene Elema (NL) discovered that the green coloring matter from Bacillus chloraphis represents a natural two-stage oxidation-reduction system {Elema, 1933 #1713}.


James Gordon Horsfall (US) found that dusting tomato seed with copper sulfate monohydrate combated damping-off {Horsfall, 1931 #11089}.


Edgar Douglas Adrian (GB) and Frederik Buytendijk (NL) may have been the first to present direct evidence that the brainstem contains all the neural elements necessary to generate breathing. Their experimental subject was the goldfish. They found that the rhythmic activity of the respiratory center in the goldfish could occur in the entire absence of sensory impulses {Adrian, 1931b #21189}.


Harry Leonard Fevold (US), Frederick Lee Hisaw (US), and Samuel Leeson Leonard (US) discovered that the anterior lobe of the hypophysis (pituitary gland) produces follicle-stimulating hormone (FSH) and interstitial cell-stimulating hormone (ICSH). The interstitial cell-stimulating hormone (ICSH) is also known as luteinizing hormone (LH) {Fevold, 1931 #3381}.


Elsa R. Orent (US), Elmer Verner McCollum (US), Arthur R. Kemmerer (US), Conrad Arnold Elvehjem (US), and Edwin Bret Hart (US) demonstrated that mammals require manganese as a nutrient {Orent, 1931 #3043;Kemmerer, 1931 #3047}.


Cornelis Bernardus Kees van Niel (NL), after elucidating the nature of bacterial photosynthesis, proposed that bacterial and plant photosynthesis are special cases of a general process in which light energy is used to photodecompose a hydrogen donor, H2A, with the released hydrogen in turn reducing carbon dioxide by dark, enzymatic reactions. The hypothesis envisaged that in plant photosynthesis H2A is water, whereas in green sulfur bacteria, e.g., H2A is H2S, with the results that oxygen becomes the by-product of plant photosynthesis and elemental sulfur the by-product of bacterial photosynthesis {van Niel, 1931 #5502;van Niel, 1941 #5503;van Niel, 1949b #5504}.

Keita Shibata (JP), in 1931, independently proposed the photo dissociation of water during oxygenic photosynthesis and a metabolism of anoxygenic photosynthetic bacteria very similar to the scheme proposed by van Niel {Shibata, 1975 #13015}.


Heinz Ohle (DE) postulated that the anaerobic breakdown of glucose begins with its phosphorylation to glucose-6-phosphate, which is converted into fructose-1, 6-diphosphate (via fructose-6-phosphate); the hexose diphosphate was considered to undergo a series of oxidations and reductions leading to the formation of glyceraldehyde-3-phosphate and dihydroxyacetone phosphate {Ohle, 1931 #2546}.

Otto Fritz Meyerhof (DE-US) would establish the formation of phosphoglyceraldehyde in hexosediphosphate cleavage {Meyerhof, 1938a #9265}.


Elsa R. Orent (US) and Elmer Verner McCollum (US) performed experiments with rats, which demonstrated that magnesium is essential to that animal’s diet. Deficiency symptoms included, dilatation of cutaneous blood vessels, extreme hyperirritability, and tetanic seizures initiated by sound {Orent, 1931 #3043}.


Hsien Wu (CN-US) proposed that in addition to peptide covalent linkages between amino acids the native organization of proteins was due mainly to polar groups. He was the first to propose that protein denaturation was a purely conformational change, i.e., corresponded to protein unfolding and not to some chemical alteration of the protein {Wu, 1931 #14614}.

Alfred Ezra Mirsky (US) and Linus Carl Pauling (US) stated, “Our conception of a native protein molecule (showing specific properties) is the following: The molecule consists of one polypeptide chain which continues without interruption throughout the molecule (or in certain cases, of two or more such chains); this chain is folded into a uniquely defined configuration, in which it is held by hydrogen bonds between the peptide nitrogen and oxygen atoms and also between the free amino and carboxyl groups of the diamino and dicarboxyl amino acid residues…. The characteristic specific properties of native proteins we attribute to their uniquely defined configuration. The denatured protein molecule we consider to be characterized by the absence of a uniquely defined configuration {Mirsky, 1936 #2446}.

Walter J. Kauzmann (US) concluded that the most important energetic driving force behind the three-dimensional structure of proteins is a hydrophobic bond {Kauzmann, 1959 #16044}.


Albert Jan Kluyver (NL) in his treatise, The Chemical Activities of Microorganisms, recognized the potential significance of the ideas that Neuberg, Wieland, Warburg, Harden, and a few others had advanced to account for specific biochemical events. Soon it became evident to him that these concepts could be welded together into a very few general principles, applicable to all biochemical phenomena. The most basic of these generalizations is the extension of the ideas of Neuberg and of Wieland to their ultimate limits. Thus, any biochemical process, whether oxidation, fermentation, or synthetic reaction, was considered as a chain of step reactions, each one of which represented a simple mechanism in which hydrogen is transferred from one molecule, the H-donor, to another, the H-acceptor. The apparent exception to this principle was exhibited in the metabolism of complex molecules, composed of several simple entities, for example the polysaccharides (complexes of simple sugars), proteins (complexes of amino acids), and fats (complexes of fatty acids and glycerol). Such complexes would first be converted to their constituent units by hydrolytic cleavages, with the products subsequently undergoing the various hydrogen-transfer reactions. In this manner the existence of the many hydrolytic enzymes—glucosidases, proteinases, lipases, etc.—could be fitted into the general picture.

He advocated the use of microorganisms for comparative biochemical studies because of the ease of handling them under controlled and reproducible conditions, and because of the enormous biochemical versatility encountered within this group. It is often possible to select a specific microorganism as singularly appropriate for a given problem because it carries out a certain type of reaction to the exclusion of almost any other. But it is equally important to realize that one may find among these creatures the best examples of seemingly quite different biochemical properties with respect to the conversion of a particular substrate.

The recognition of this unity at the molecular level is Kluyver’s great contribution; it is also the starting point of comparative biochemistry {Kluyver, 1931 #103}.


Warren Harmon Lewis (US) characterized the process of pinocytosis {Lewis, 1931 #12248}.


Johann Heinrich Emil Heitz (DE) and Barbara McClintock (US) demonstrated that the nucleolus in somatic cells is regularly associated with a particular locus on one or more chromosomes. McClintock defined this region as the nucleolar organizer {Heitz, 1931 #6123;McClintock, 1934 #6124}.


Frank L. Howard (US) was the first to describe the life cycle of the slime mould Physarum polycephalum {Howard, 1931 #27927}.


Edgar Douglas Adrian (GB) reported that isolated ventral nerve cords of a water beetle Dytiscus marginalis produced rhythmic output {Adrian, 1931a #21146}.

Donald M. Wilson (GB) explored the control of rhythmic locomotion of flight mechanics in the desert locust Schistocerca gregaria. He provided evidence to reject the proprioceptive chain reflex model by rigorously demonstrating that the full motor pattern of locust flight could be generated by fully deafferented thoracic ganglia, which could not receive the inputs required by the proprioceptive chain model. He proposed that ‘the basic co-ordination of flight is an inherent function of the central nervous system but that peripheral feedback loops influence the frequency of operation and details of pattern' {Wilson, 1961 #21144}.

Keir G. Pearson (CA) and John F. Iles (CA) focused on the hind legs of the cockroach Periplaneta americana and, in particular, on a subset of the muscles that raise (levate) and lower (depress) the limbs during the swing and stance movements of a step cycle, respectively. They demonstrated for the first time that alternating patterns of levator and depressor motor activity like those seen in walking were generated centrally in the absence of leg sensory feedback. Put simply, the basic motor pattern was generated within the central nervous system {Pearson, 1970 #21165}.

Keir G. Pearson (CA) confirmed the work in the 1970 paper then went on to provide strong evidence that sensory feedback acted to modify the centrally generated pattern of activity {Pearson, 1972 #21166}.

Donald M. Wilson (GB) emphasized the corrective role of sensory feedback: ‘the importance of sensory feedback in behavior patterns appears not to lie in the cueing of sequences but rather in the correction of errors inherent in genetically determined motor programs’ {Wilson, 1972 #21145}.


John Charles Walker (US) developed and released peas resistant to Fusarium wilt and near-wilt disease {Walker, 1931 #11187}.


Karl Friedrich Meyer (CH-US), Clarence M. Haring (US), and Beatrice Howitt (US) isolated the virus of Western equine encephalomyelitis from the brains of infected horses during an outbreak in California {Meyer, 1931 #2379;Meyer, 1956 #12302}.


Wilbur A. Sawyer (US) and Wray Lloyd (US) developed a Serum Neutralization Test for the presence of yellow fever {Sawyer, 1931 #2368}.

Fred L. Soper (US), Elmer R. Rickard (US), and Peter J. Crawford (US) perfected the postmortem diagnosis of yellow fever by viscerotomy (histological examination of the liver) {Soper, 1934 #2369}. This greatly aided in field surveys of yellow fever.

Wilber A. Sawyer (US), Stuart F. Kitchen (US), and Wray Lloyd (US) developed a vaccine to yellow fever using the attenuated virus. The vaccine had two parts: a ten-percent suspension of mouse-brain tissue with yellow fever virus in fresh sterile human serum, and human immune serum from people recently recovered from yellow fever {Sawyer, 1932 #21850}.


Alice Miles Woodruff (US) and Ernest William Goodpasture (US) developed the first practical method for cultivating large quantities of a virus in the laboratory—by growing it on the exposed membrane of a chick embryo {Woodruff, 1931 #5724}.


Charles E. Clifton (US), Edwin William Schultz (US), and Louis P. Gebhardt (US) used ultrafiltration collodion membranes to estimate that poliomyelitis virus is less than 50 nanometers in diameter {Clifton, 1931 #15896}. Subsequent work by them placed its size at close to 25 nanometers.

Frank Macfarlane Burnet (AU) and Annie Jean Macnamara (AU) discovered that there is more than one strain of the poliomyelitis virus {Burnet, 1931 #22748}.

David Bodian (US), Isabel Merrick Morgan (US), and Howard A. Howe (US) identified 14 strains of poliovirus, which are divisible into three immunological types capable of causing poliomyelitis {Bodian, 1949 #26024}.

James D. Trask (US), Alfred J. Vignec (US), and John R. Paul (US) isolated poliovirus from human feces {Trask, 1938b #1617;Trask, 1938a #1618}.

John Franklin Enders (US), Thomas Huckle Weller (US), and Frederick Chapman Robbins (US) were the first to grow poliovirus to high titer in cell culture. They used human embryonic extraneural tissue {Enders, 1949 #5652;Weller, 1949 #9157;Enders, 1952 #9158}.

Jordi Casals (ES-US), Peter K. Olitsky (US), and Ralph O. Anslow (US) adapted type 2-poliomyelitis virus to suckling mice and showed that the brains of these animals contained antigen in sufficient concentration to fix complement with poliomyelitis antisera {Casals, 1951 #14864}.

Joseph Louis Melnick (US) and Nada Ledinko (US) reported immunity following oral administration of poliomyelitis virus to monkeys {Melnick, 1951 #21739}.

Hilary Koprowski (PL-US), George A. Jervis (US), and Thomas W. Norton (US) created the world's first polio vaccine, based on oral administration of attenuated poliovirus. In researching a potential polio vaccine, they had focused on live viruses that were attenuated (rendered non-virulent) rather than on killed viruses. They developed the polio vaccine by attenuating the virus in brain cells of the cotton rat (Sigmodon hispidus), a New World species susceptible to polio. They then conducted the first human trial of this attenuated oral poliovirus vaccine, first treating themselves, then at a New York State facility for intellectually disabled children and children with epilepsy {Koprowski, 1952 #21737}.

Jonas Edward Salk (US) prepared a vaccine of chemically inactivated poliovirus {Salk, 1953a #5653;Salk, 1953a #7855;Salk, 1955a #11516;Salk, 1955b #18502}. Why this vaccine was chosen for widespread use in lieu of either the Koprowski or Sabin vaccines is interesting.

Albert Bruce Sabin (PL-US) developed a polio vaccine containing live attenuated viruses from the three known strains of poliovirus. He tried the vaccine on himself first then on prison volunteers. The vaccine did not displace the Salk vaccine until 1960 when its use abroad on more than 100 million people made it apparent that it was superior {Sabin, 1955 #5654;Sabin, 1960 #11514}.

Herald Rea Cox (US), Victor J. Cabasso (US), Floyd S. Markham (US), Max J. Moses (US), Arden W. Moyer (US), Manuel Roca-Garcia (US), and James M. Ruegsegger (US) developed a live attenuated trivalent oral poliomyelitis vaccine for humans {Cox, 1959 #25825}.


Richard E. Shope (US) and Paul A. Lewis (US) found that two infectious agents are necessary to induce the most severe form of swine influenza, the influenza virus and a bacterium he called Haemophilus influenzae suis. The virus alone produced a mild form of the disease {Shope, 1931a #2358;Lewis, 1931 #10915;Shope, 1931b #10914}. He later speculated that survival of the virus during inter-epidemic periods involves virus persisting in a latent form in the parasitic pulmonary nematode (lung worm) of the pig. The virus is incorporated in the ova of the nematode, which is later eaten by an earthworm that serves as its intermediate host. After the lung worm reaches a certain stage of development the earthworm is eaten by the pig and influenza virus inside the nematode returns to the lungs of the same or another pig {Shope, 1935 #6991;Shope, 1936d #6992;Shope, 1936e #10918;Shope, 1939 #5732;Shope, 1943 #6989}.


Robert Daubney (GB), James Ralph Hudson (GB), and Percy Cyril Claude Garnham (GB) were the first to isolate the Rift Valley fever virus (RVFV), a type of bunyavirus (Phlebovirus). They found it in East African sheep {Daubney, 1931 #8280}.


Sara Elizabeth Branham (US), Clara Enola Taft (US), and Sadie A. Carlin (US) identified a new organism, Neisseria flavescens, as a rare cause of meningitis and septicemia in humans, but one requiring careful differentiation from meningococcus {Branham, 1931 #15810;Branham, 1932 #15811}. In 1970 Branham was honored posthumously by the name of a new genus, Branhamella.


Ernst Lowenstein (DE) and Kai Adolf Jensen (DK) developed a culture medium for the isolation and cultivation of mycobacteria and as bases for selective, differential, and enriched media for mycobacteria. Malachite green is used to inhibit non-mycobacterial organisms {Jensen, 1932 #17583;Lowenstein, 1931 #17582}.


James S. Anderson (GB), Frank C. Happold (GB), James Walter McLeod (GB), and J.G. Thomson (GB) discovered that severe cases of diphtheria are nearly always caused by a genetic variety of Corynebacterium diphtheriae referred to as gravis. The mild cases were usually associated with the variety called mitis. The two varieties were easily distinguishable from one another because of their characteristic colonies formed on solid media. They devised a chocolate agar tellurite medium for the growth of the diphtheria bacilus, and by its use were able to divide the Klebs-Loeffler bacillus into three types-gravis, mitis, and "intermediate" {Anderson, 1931 #104}.


Margaret Pittman (US) determined that pathogenicity in Haemophilus influenzae is influenced by variation in encapsulation types. She found that type b (called Hib, for Haemophilus influenzae type b) caused nearly all cases of Haemophilus influenzae meningitis. It would later be confirmed that Hib could also cause many other serious diseases, including infections of the blood, bone, and joints {Pittman, 1931 #8811}.


René Jules Dubos (US) and Oswald Theodore Avery (CA-US) discovered Bacillus brevis, an organism that breaks down the capsular polysaccharide of Type 3 Streptococcus pneumoniae and protects mice against pneumonia {Dubos, 1931 #12418}.


A.S. Michailoff (DE) was the first to report on a mutation in the honeybee, Apis mellifera Linn. It was the ivory eye color which he referred to as white {Michailoff, 1931 #1896}.


Harriet B. Creighton (US) and Barbara McClintock (US), working with corn (Zea mays), obtained cytological proof that the inheritance of novel gene combinations during crossing over is due to the physical exchange of specific chromosome segments {Creighton, 1931 #1086}.

Curt Stern (DE-US), using cytologically abnormal X-chromosomes of Drosophila, one with the X-Y translocation, and one with an X-IV translocation, demonstrated that the genetic phenomenon of crossing over is accompanied by a physical exchange between the chromosomes {Stern, 1931 #1085}. See, Janssens, 1909.


G.K. Chrustschoff (RU), A.H. Andres (RU), W.I. Iljina-Kakujewa (RU), E.A. Berlin (RU), and P. Zhivago (RU) were the first to use cultures of peripheral blood cells for the study of chromosomes {Chrustschoff, 1931 #1227;Andres, 1933 #1228;Chrustschoff, 1935 #1229}.

P. Zhivago (RU), B. Morosov (RU), and A. Ivanickaya (RU) discovered that a much better spread of chromosomes occurs if water is used instead of isotonic solutions during the preparation of a chromosome squash {Zhivago, 1934 #1208}.

Tao-Chiuh Hsu (US), Sajiro Makino (JP), Isao Nishimmura (JP), and Arthur Hughes (GB) rediscovered the use of water instead of isotonic solutions in 1952 {Hsu, 1952 #7665;Hughes, 1952 #20155;Makino, 1952 #1209}.


Franklin R. Miller (US) clearly demonstrated that plasma cells were not derived from lymphocytes {Miller, 1931 #25571}.


Leonard H. Snyder (US) and Albert Francis Blakeslee (US), based on experiments using phenyl thiocarbamide (PTC), concluded that tasting is inherited as a Mendelian recessive trait, with "taste" being dominant to "non-taste." In addition, Blakeslee showed that there is a threshold concentration below which "tasters" cannot detect PTC {Blakeslee, 1931 #20646;Snyder, 1931 #20647}.


Sewall Green Wright (US) presented the first unified picture of evolution in terms of Mendelism by illustrating the relations between selection pressure, mutation rates, inbreeding, and isolation {Wright, 1931 #5042}.


John Punnett Peters (US) and Donald Dexter van Slyke (US) wrote the two volume Quantitative Clinical Chemistry, for many years the world authority {Peters, 1931 #10943;Peters, 1932 #10944}.


Charles Edward Spearman (GB) and Gerhard Kärber (DE) developed a nonparametric procedure for computing an ED50 estimate (and fiducial limits) during drug research {Kärber, 1931 #9408;Spearman, 1908 #20180}. This became known as the "Spearman-Kärber estimator", which is frequently used in drug screening.


Leopold Heine (DE) performed work, which made the manufacture of modern contact glasses possible {Heine, 1931 #14256}.


Philip Duryeé McMaster (US) and Stephen S. Hudack (US) proved that lymphatics instead of being passive drainage canals are very active in the process of fluid exchange. Their walls respond rapidly to various influences such as sunlight, warmth, or a state of shock that does not break the skin {McMaster, 1931 #10893;Hudack, 1931 #10894;McMaster, 1932b #10896;Hudack, 1932a #10897;Hudack, 1932c #10898;Hudack, 1932b #10899;McMaster, 1932d #10900;Hudack, 1933 #10901}.


Maxwell Myer Wintrobe (CA-US) developed the methodology for direct calculation of the average erythrocyte size, mean corpuscular volume (MCV) in cubic microns, mean hemoglobin content (MCH) in picograms, and mean corpuscular hemoglobin concentration (MCHC) in percent—quantifications that are standard clinical procedure today {Wintrobe, 1931 #10781}.


Vincent Brian Wigglesworth (GB) was the first to report on Rhodnius prolixus (Hemiptera, Reduviidae) Malpighian tubule physiology {Wigglesworth, 1931a #21229;Wigglesworth, 1931b #21230;Wigglesworth, 1931c #21231}.


Myrtelle May Canavan (US) was the first to describe the disease, which would bear her name {Canavan, 1931 #22626}. Canavan disease is an autosomal recessive degenerative disorder that causes progressive damage to nerve cells in the brain and is one of the most common degenerative cerebral diseases of infancy. It is caused by a deficiency of the enzyme aminoacylase 2.


Ernest Amory Codman (US) described an epiphyseal chondromatous giant cell tumor of the proximal humerus, hence the term "Codman tumor" {Codman, 1931 #27343}.


Alwin Max Pappenheimer, Jr. (US) and Marianne Goettsch (US) showed that growing chicks maintained on a diet consisting of milk powder, casein, starch, yeast, cod liver oil, salts and filter paper develop ataxia, tremors, retraction or twisting of the head, clonic spasms of the legs, and stupor. These symptoms may appear suddenly, usually between the 18th and 25th day, and may end in death. If recovery takes place, the chicks may go on to normal development.

Definite lesions are found in the cerebellum of the affected chicks. These consist of edema, necrosis and hemorrhages. Hyaline thrombi are found in the capillaries in and about the degenerated areas {Pappenheimer, 1931 #23390}.


Samuel S. Shouse (US), Stafford L. Warren, II (US), and George Hoyt Whipple (US) provided descriptions of the anatomic and functional effects of radiation injury {Shouse, 1931 #10571}.


Harvey Williams Cushing (US) removed a pituitary tumor from a patient on April 15, 1931. This was the 2000th verified brain tumor removed by Cushing in his remarkable career during which he reduced the mortality rate in brain surgery from more than 90 percent to a little over 8 percent. Cushing operated for an additional fifteen months before retiring {Riedman, 1962 #7656}. Note: In a supreme irony he died in 1939 of a brain tumor.

Harvey Williams Cushing (US) reported on the surgical-mortality percentages pertaining to the two thousand operations he had performed for the removal of intracranial tumors {Cushing, 1931 #18229}.


Ernst Ferdinand Sauerbach (DE) performed the first successful surgical intervention in the cardiac system when he operated on an aneurysm of the heart {Sauerbruch, 1931 #27012}.


Christian Georg Schmorl (DE) established the modern basis for understanding the intervertebral disc, by providing very clear discussions of herniations as well as degenerations {Schmorl, 1931 #16188}.


James Rognvald Learmonth (IR) observed that if the hypogastric nerve is stimulated the result is contraction of the ipsilateral ureteral orifice; tightening of the trigone; contraction of the internal sphincter; and contraction of the musculature of the prostate, seminal vesicles and ejaculatory ducts. Sectioning the hypogastric nerve produces relaxation of the ureteral orifice, trigone and internal sphincter, with no appreciable effect on the dome or lateral walls of the bladder {Learmonth, 1931 #18440}.

Derek Ernest Denny-Brown (NZ-GB-US), E. Graeme Robertson (AU), and Alex E. Roche (GB) studied the events of voiding and the volumetric changes of the bladder in relation to sphincter activity. They found spontaneous waves of pressure in the bladder during filling. As these increased in amplitude, subjective sensations of fullness or urinary urgency developed in the subjects. The subjects, when asked, could voluntarily suppress the summation of these pressure waves {Denny-Brown, 1933a #12278;Denny-Brown, 1933b #18439;Denny-Brown, 1936-1937 #12279}.


Michael S. Burman (US) carried out the first extensive academic research in arthroscope usage in the United States. Arthroscopy being the direct visualization of joints and other cavities such as the spinal canal via lumbar puncture {Burman, 1931 #11744}.


Rudolf Nissen (DE) successfully removed an entire bronchiectatic (irreversible dilatation of part of the bronchial tree) lung {Nissen, 1931 #13427}.


Paul Dudley White (US) virtually created the specialty of cardiology in the United States when he published his book, Heart Disease {White, 1931 #19140}.


Another outbreak of polio swept the U.S. during the summer of 1931, killing more than 4,000 people, about 12 percent of the reported cases {Kohn, 1995 #4873}.


George Gaylord Simpson (US) united the two families of anthropoid apes, Pongidae (Simiidae), and men, Hominidae, into a Superfamily, which he called Hominoidea, and distinguished from the other two superfamilies of the sub-order Anthropoidea, which he called the Cercopithecoidea and the Ceboidea. The superfamily Hominoidea includes the hominoids (gibbons, great apes, and humans). The term hominid refers only to humans {Simpson, 1931 #12329}.


Marius Pièry (FR), Julien Roshem (FR), and Vilhelm Moller-Christensen (DK) gave accounts of Stone Age homonoid skeletons diagnosed as exhibiting signs of tubercular damage. Evidence of pulmonary tuberculosis remains but, in the nature of the case, it is meager {Pièry, 1931 #5;Moller-Christensen, 1966 #6}.


Erik Andersson Stensiö (SE) and Gunnar Säve-Söderbergh (SE), of the 1929-1930 Danish scientific expeditions, found ichthyostegid fossils in the upper Devonian sediments in eastern Greenland. They appear to be intermediate between lobe-finned rhipidistians (Osteolepis) and early amphibians. These are the oldest known fossils that can be classified as amphibians {Säve-Söderbergh, 1932 #14165;Stensiö, 1931 #14166}.


Arthur Tindell Hopwood (GB), in 1931, discovered the fossils of three hominids near Lake Victoria, he would name them Proconsul in 1933 {Hopwood, 1932 #17776;Hopwood, 1933 #17775}. This was the oldest known ape found up to this time.

Wilfred E. Le Gros Clark (GB) and Louis Seymour Bazett Leakey (GB-KE), reported on a hominid skull and jaw found by Mary Douglas Nicol Leakey (GB-KE), in 1948, in Miocene deposits on Rusinga Island in Lake Victoria, Western Kenya. It was an excellent sample of Proconsul africanus (sometimes called Dryopithicus africanus or "woodland ape"). The specimen is approximately 16 million years old. Proconsul is a candidate for the distant ancestor from which all modern species of apes and all hominids—human beings included—evolved.


Henry Bryant Bigelow (US), cnidariologist, founded Woods Hole Oceanographic Institution (WHOI) in 1931 and became its first director until 1940. He is commemorated by Haliscera bigelowi Kramp, 1947, Bathyplotes bigelowi Deichmann, 1940, Braarudosphaera bigelowii Gran et Braarud Deflandre, 1947, Casco bigelowi Blake, 1929, Lucaya bigelowi Chace, 1939, Euphysora bigelowi Maas, 1905, Neoturris bigelowi Kramp, 1959, Calycopsis bigelowi Vanhöffen, 1911, and Octophialucium bigelowi Kramp, 1955.



“If a patient is poor he is committed to a public hospital as psychotic; if he can afford the luxury of a private sanatorium, he is put there with the diagnosis of neurasthenia; if he is wealthy enough to be isolated in his own home under constant watch of nurses and physicians he is simply an indisposed eccentric.” Pierre Marie Félix Janet {Janet, 1932 #13340}.


“If mankind is to profit freely from the small and sporadic crop of the heroically gifted it produces, it will have to cultivate the delicate art of handling ideas. Psychology is now able to tell us with reasonable assurance that the most influential obstacle to freedom of thought and to new ideas is fear; and fear which can with inimitable art disguise itself as caution, or sanity, or reasoned skepticism, or on occasion even as courage.” Wilfred Batten Lewis Trotter {Trotter, 1932 #22821}.


“It was not noisy prejudice that caused the work of Mendel to lie dead for thirty years, but the sheer inability of contemporary opinion to distinguish between a new idea and nonsense.” Wilfred Batten Lewis Trotter {Trotter, 1932 #22821}.


“The ordinary patient goes to his doctor because he is in pain or some other discomfort and wants to be comfortable again; he is not in pursuit of the ideal of health in any direct sense. The doctor on the other hand wants to discover the pathological condition and control it if he can. The two are thus to some degree at cross purposes from the first, and unless the affair is brought to an early and happy conclusion this diversion of aims is likely to become more and more serious as the case goes on.” Wilfred Batten Lewis Trotter {Trotter, 1941 #22826}.


Charles Scott Sherrington (GB) and Edgar Douglas Adrian (GB) were awarded the Nobel Prize in Physiology or Medicine for their discoveries regarding the functions of neurons.


Max Knoll (DE) and Ernst August Friedrich Ruska (DE) originated the concept of a transmission electron microscope (TEM) {Knoll, 1932 #14669}.

Ernst August Friedrich Ruska (DE) and Max Knoll (DE) built the first transmission electron microscope (TEM) at the Technische Hochschule of Berlin-Charlottenburg.

Albert F. Prebus (CA) and James Hillier (CA-US) made the TEM a practical instrument {Prebus, 1939 #14705}. See, M.M. Freundlich. 1963. Origin of the electron microscope. Science 142 (3589): 185-188.


Harold Clayton Urey (US), Ferdinand G. Brickwedde (US), and George M. Murphy (US) were the first to prove that hydrogen gas contains a few atoms in which the nuclear proton is accompanied by a neutron. This so called heavy hydrogen was given the name deuterium {Urey, 1932a #10994;Urey, 1932b #10995}.


Martin Gerhard Behrens (DE) used differential centrifugation to separate chloroplasts, nuclei, and cytoplasm {Behrens, 1932 #16810;Behrens, 1939 #16809}.


S. Furukawa (JP) isolated four terpenes in his quest to determine the active constituents of the Ginkgo extracts used in folk medicine {Furukawa, 1932 #3583;Furukawa, 1933 #3584;Furukawa, 1934 #3585}.

Koji Nakanishi (JP) determined the structures of the ginkgolides {Nakanishi, 1967 #3586}.

Elias James Corey (US), Myung-Chol Kang (US), Manoj C. Desai (US), Arun K. Ghosh (US), and Ioannis N. Houpis (US) carried out the complete synthesis of ginkgolide B {Corey, 1988 #3582}.


Wendell Meredith Stanley (US), Gerald H. Coleman (US), Clay M. Greer (US), Jacob Sacks (US), and Roger Adams (US) elucidated and synthesized chaulmoogric and hydnocarpic acids, the active ingredients of chaulmoogra oil, a folk remedy for leprosy {Stanley, 1932 #6915}.


Werner Schulemann (DE), Fritz Schönhöfer (DE), and August Wingler (DE) announced the synthesis of plasmoquine (pamaquine, plasmochin) for the treatment of malaria {Schulemann, 1932 #19058}.


Leonor Michaelis (DE-US) presented the first example of a reversible two-stage oxidation-reduction system {Michaelis, 1932 #1712}.


Heinrich Otto Wieland (DE) and Elisabeth Dane (DE) described the correct structure for cholesterol {Wieland, 1932 #19072}.


Sigmund Otto Rosenheim (GB) and Harold King (GB) worked out the correct structure for the rings found in bile acids and sterols (including cholesterol) {Rosenheim, 1932a #1714;Rosenheim, 1932b #1716;Rosenheim, 1932c #1715;Rosenheim, 1933 #1717}.


Cecil James Watson (US) crystallized stercobilin from human feces and proved that it is different from urobilinogen and mesobilirubinogen. In the process he discovered mesobiliviolin in nature for the first time {Watson, 1932 #10605}.


Manayath Damodaran (GB), Gunnar Jaaback (GB), and Albert Charles Chibnal (GB) isolated glutamic acid from the hydrolysate of a protein {Damodaran, 1932 #6010}.


Joseph K. Tillmans (DE), Paul Hirsch (DE), and Johannes Jackisch (DE) were the first to introduce a quantitative chemical test for the presence of vitamin C. If was based on the relationship between the vitamin C content of a food and its reducing power {Tillmans, 1932 #2974}.


Harry D. Kruse (US), Elsa R. Orent (US), and Elmer Verner McCollum (US) reported that when young rats are restricted to a ration containing only 1.8 parts per million of magnesium but adequate amounts of other dietary substances, they develop a characteristic and striking symptomatology that presages an early death. The animals pass successively through stages of vasodilatation, hyperirritability of the nervous system, cardiac arrhythmia, and fatal tonic-clonic convulsions. The conclusion was that magnesium is an essential element for normal functioning, growth and life {Kruse, 1932 #23385}.


Robert Emerson (US) and William A. Arnold (US) articulated the concept of a photosynthetic unit (photosystem), which includes the two components, antenna and reaction center {Emerson, 1932a #10518;Emerson, 1932b #10519}.


Robert F. Parker (US) and Ralph S. Muckenfuss (US) adapted the Complement Fixation Test for the detection of smallpox antigen in lesion fluid. This represents the first immunological test for viral antigens {Parker, 1932 #8718}.


Max Schlesinger (DE-GB) used the adsorption capacity of the bacterial cell for bacteriophages and the sedimentation velocity of bacteriophages to determine that the bacteriophage particle has a maximum linear dimension of about 0.1 micrometer and a mass of about 4 x 10-16 g. His studies of the kinetics of bacteriophage attachment imply that Brownian movement brings bacteriophage particles into random collisions with the bacterial surface. He purified a weighable amount of T-even bacteriophage by differential centrifugation and graded filtration; then by direct chemical analysis found that they consisted mainly of protein and DNA, in roughly equal proportions {Schlesinger, 1932 #19070;Schlesinger, 1934 #16822;Schlesinger, 1936 #5729}.


Max Bergmann (DE-US) and Leonidas Zervas (DE-US) developed a method for synthesis of polypeptides, which was superior to the Fischer synthesis because it allowed the inclusion of complex amino acids in the polymer {Bergmann, 1932 #2447}.


Otto Heinrich Warburg (DE) and Walter Christian (DE) isolated, from yeast, a yellow-red protein which they called oxygen-transporting-ferment and concluded that, “The yellow ferment is therefore not only an oxygen-transporting ferment but also a ferment of oxygen-less respiration. … It is probable that in life, the yellow ferment does not transfer molecular, but bound oxygen. Probably, in life, it is not an oxygen-transporting ferment but an oxidation-reduction ferment.” They went on to show that the pigment is a small molecule, which is released when the protein portion of the complex is denatured. The pigment was found to belong to the class of substances that Richard Johann Kuhn (AT-DE) had named flavins {Warburg, 1932 #2526;Warburg, 1933 #2527}. Note: The 1932 article includes the isolation of NADP.

Kurt Guenter Stern (GB-US) and Ensor Roslyn Holiday (GB) determined the chemical structure of the flavins {Stern, 1934 #2528}.

Richard Johann Kuhn (AT-DE), Karl Reinemund (DE), Friedrich Weigand (DE), Rudolf Ströbele (DE), Paul Karrer (RU-CH), B. Becker (CH), F. Benz (CH), P. Frei (CH), H. Salomon (CH), Kurt Schöpp (CH), and T.H. Quibell (CH) synthesized many flavins including lactoflavin, which was designated vitamin B2 or riboflavin {Kuhn, 1935a #2529;Kuhn, 1935b #3004;Karrer, 1935 #3003;Karrer, 1936 #3002}. Riboflavin alone was found to be inactive.

Axel Hugo Theodor Theorell (SE) showed that the coenzyme associated with Warburg’s yellow enzyme had a structure like riboflavin (vitamin B2), to which a phosphate group was attached {Theorell, 1935 #2544}.

Richard Johann Kuhn (AT-DE), Hermann Rudy (DE), and Friedrich Weygand (DE) found that the active form had a phosphate group at the five-prime position (vitamin B2 is riboflavin phosphate or flavin mononucleotide) and behaves as a prosthetic group carrying out its catalytic function while complexed with protein as a flavoprotein {Kuhn, 1936 #2530}.

Otto Heinrich Warburg (DE) discovered flavoproteins based on simple observations on lactobacilli that lack the red cytochromes. On exposure to air the intact cells become yellow. He isolated dehydrogenases, flavoproteins, and identified their coenzymes {Warburg, 1938a #119}.

Otto Heinrich Warburg (DE) and Walter Christian (DE) working with a preparation of D-amino oxidase isolated a flavin derivative, which would later be shown to be flavine-adenine-dinucleotide (FAD) {Warburg, 1938b #2532}.

Erwin Haas (US) showed that this FAD is also the prosthetic group of a flavoprotein isolated from yeast {Haas, 1938 #2531}.

Michael Doudoroff (RU-US) discovered that riboflavin (vitamin B2) is directly involved in bacterial luminescence {Doudoroff, 1938 #10678}.

Samuel M. Hunter Christie (GB), George Wallace Kenner (GB), and Alexander Robertus Todd (GB) carried out the total synthesis of flavin-adenine-dinucleotide (FAD) {Christie, 1952 #24379}.


Joachim Hämmerling (DE) and Charlotte Hämmerling (DE) performed a series of classic experiments using the alga Acetabularia spp. as his material. They concluded that there must be intermediate stages between the genetic information of the cell nucleus and expression of the genetic information in the cytoplasm. Furthermore, these intermediate stages must be represented by chemically defined substances (m-RNA), which were called morphogenetic substances, and even in darkness the nucleus continues to supply the cytoplasm with morphogenetic substances. In the 1934 reference Hämmerling found that chloroplasts divide in anucleate cells {Hämmerling, 1932 #8644;Hämmerling, 1934 #8645;Hämmerling, 1935 #8646;Hämmerling, 1943 #8647;Hämmerling, 1955 #8648;Hämmerling, 1959 #8650;Hämmerling, 1963 #8649}. Note: This work is remarkable as a harbinger of molecular biology.


Hans Adolf Krebs (DE-GB) and Kurt Henseleit (DE) worked out the way in which the body eliminates waste nitrogen, usually from deamination reactions, by breakdown and regeneration of a portion of the amino acid arginine. This urea cycle (or ornithine cycle), as it is called, ultimately leads to the excretion of waste nitrogen as urea. In their own words, “The primary reaction of urea synthesis in the liver is the addition of one molecule of ammonia and one molecule of carbonic acid to the d-amino group of ornithine, with the elimination of one molecule of water and the formation of a d-ureido acid, citrulline … The second reaction of urea synthesis is the combination of one molecule of citrulline with an additional molecule of ammonia, with the loss of a second molecule of water and the formation of a guanidino acid, arginine … The third reaction is the hydrolytic cleavage of arginine to ornithine and urea” {Krebs, 1932a #2600;Krebs, 1932b #21271;Krebs, 1932c #25553}.This was the first cyclic reaction path to be elucidated and aroused an enthusiastic response from the scientific community.


Rudolph Albert Peters (GB), Nicolai Gavrilescu (HU), Arnold Peter Meiklejohn (GB), and Reginald Passmore (GB) were the first to demonstrate in vitro that a vitamin (thiamine) has a specific enzymatic aiding action {Gavrilescu, 1932 #1651;Peters, 1936a #1650}.


V. Albrecht Bethe (DE) introduced the concept of ectohormones, now known as pheromones {Bethe, 1932 #8690}. See Fabre, 1879 and Karlson, 1959. One member of a species to affect the physiology or behavior of another member of the same species excretes these chemicals.

Hermann Otto Laurenz Fischer (DE) and Erich Baer (DE) synthesized DL-glyceraldehyde-3-phosphate to be tested as an intermediate in alcoholic fermentation {Fischer, 1932 #2547}.

Carl Vincent Smythe (DE) and Waltraut Gerischer (DE), in 1933, showed that yeast could ferment DL-glyceraldehyde-3-phosphate, consuming only one of the isomers {Smythe, 1933 #15882}. Later work showed that it was the D isomer being metabolized.


Edwin Martin Case (GB) discovered that muscle extract produces pyruvic acid from hexosediphosphate or glycogen {Case, 1932 #1728}.


Marcus Morton Rhoades (US) demonstrated in maize that crossing over among chromosomes occurs at the four-strand stage {Rhoades, 1932 #11095}.


Franz Schrader (DE-US) concluded that spindles in living cells do indeed have a fibrous structure, albeit an invisible one. In bipolar spindles he described these fibrous arrays as consisting of: 1) those running from a spindle pole to each chromosome or chromatid, namely, chromosomal fibers or half-spindle components; 2) continuous fiber systems from pole to pole; and, in some organisms 3) interzonal fibers of quite different nature connecting the ends of separating chromatids at anaphase {Schrader, 1932 #10704;Schrader, 1934 #10705}.


William Henry Chandler (US), Dennis Robert Hoagland (US), and Paul Llewellyn Hibbard (US) discovered that zinc deficiency causes several physiological disorders in plants, including little leaf in peaches, mottle leaf in citrus, and rosette in apples {Chandler, 1932 #10757}.

Howard S. Reed (US) and Jean Dufrenoy (US) described the signs and symptoms of zinc deficiency in plants {Reed, 1935 #3091}.


Richard Edwin Shope (US) showed that mad itch, a violent, distressing, and fatal disease of cattle in the American Midwest is caused by a virus transmissible to rabbits, and endemic among pigs. He identified mad itch as being identical to pseudorabies in Europe {Shope, 1932b #10919}.


Richard Edwin Shope (US) discovered the viral etiology of rabbit fibroma disease found in wild cotton tail rabbits (Sylvilagus) in the United States {Shope, 1932a #7017;Shope, 1936a #7018;Shope, 1936b #10916;Shope, 1936c #10917;Shope, 1937 #7019}. This is often called Shope fibroma.

Francis Peyton Rous (US), and Joseph W. Beard (US) successfully induced a tumor in mammals using a virus. The virus causes a papilloma in rabbits. Today it is called the Shope papilloma virus {Rous, 1935a #12336;Rous, 1935b #12337}.


Chester Hamlin Werkman (US) and George Francis Gillen (US) proposed the bacterial generic term Citrobacter for the citrate-positive, coli-aerogenes intermediates, with Citrobacter freundii Braak as the type species {Werkman, 1932 #2326}.


Henry Pinkerton (US) and George M. Hass (US) grew Rickettsia in tissue culture {Pinkerton, 1932 #6924}.


Carl Clarence Lindegren (US) worked out much of the basic genetics of the fungus, Neurospora {Lindegren, 1932 #986}.


Margaret Newton (CA-US) and Thorvaldur Johnson (CA) developed techniques to allow them to hybridize the rust fungus while it grew on barberry. They discovered that pathogenic characters segregate in a Mendelian fashion while some other characters appear to be inherited in a maternal pattern {Johnson, 1946a #1682;Johnson, 1946b #17184;Newton, 1932 #17844;Newton, 1940 #17845}.


Frans Verdoorn (NL) edited an excellent manual on the bryophytes (liverworts and mosses) which, among other things, discussed their classification, ecology, distribution, morphology, physiology, cytology, and genetics {Verdoorn, 1932 #5129}.


Gottfried Samuel Fraenkel (DE-US) determined the direction of air flow in the tracheal system of the locust {Fraenkel, 1932a #26684;Fraenkel, 1932b #26685}.


Helmuth Weese (DE) and Walther Scharpff (DE) reported on their first experiences with hexobarbital (the first short-acting barbiturate) for intravenous induction of general anesthesia. It is also called evipan and evipal {Weese, 1932 #14739}.


William F. Hamilton (US), Jack Walker Moore (US), James M. Kinsman (US), and Roy G. Spurling (US) performed experiments, which exposed hemodynamics under physiological and pathological conditions {Hamilton, 1932 #9415}.


Eliot R. Clark (US) and Eleanor Linton Clark (US) placed glass-windowed chambers in the rabbit ear and made exquisite drawings of the branching patterns of the blood vessels that entered the wound. Their work established the field of vascular biology {Clark, 1932 #16663}.


Philip Duryeé McMaster (US), Stephen S. Hudack (US), and Francis Peyton Rous (US) found that there is a relation of hydrostatic pressure to the gradient of capillary permeability {McMaster, 1932a #10895}.


James Augustine Shannon (US), Norman Jolliffee (US), Homer W. Smith (US), and Saul Fisher (US), in their studies of the kidney, developed a way to precisely measure the rate of formation of glomerular filtrate (i.e., the volume per unit time of plasma ultrafiltrate that, under the hydrostatic pressure in the arterial circulation, enters the kidney tubules and flows toward the urinary bladder) in a variety of species—amphibians, reptiles, birds, and mammals, including Homo sapiens. They went on to illuminate passive reabsorption from tubular lumen back across the tubular epithelium to the peritubular blood, simple diffusion along a chemical concentration gradient of urea created by the reabsorption of water, active reabsorption, and active secretion {Jolliffee, 1932a #11120;Shannon, 1932 #11121;Shannon, 1934 #11122;Shannon, 1935a #11123;Shannon, 1935b #11124;Shannon, 1935c #11125;Shannon, 1935d #11126;Shannon, 1936 #11127;Shannon, 1938a #11128;Shannon, 1938b #11129;Shannon, 1938c #11130;Shannon, 1939 #11131}. See, Eli Kennerly Marshall, Jr., 1923, and Joseph Treloar Wearn, 1924.

James Augustine Shannon (US), Saul J. Farber (US), and Leonard Troast (US) determined the transport maximum (Tm) for glucose in the normal dog kidney tubule. The value was found to be stable over months and not influenced by blood levels of glucose or insulin. They proposed that it be used as a marker of reabsorption capacity {Shannon, 1941 #16250}.


Allen Dudley Keller (US) and William Kendrick Hare (US) gave the heat regulating function of the brain a more precise location in the hypothalamus {Keller, 1932 #3350}.


James Gray (GB) produced three papers, which represent the cornerstone of our understanding of aquatic locomotion by animals {Gray, 1933a #15766;Gray, 1933b #15767;Gray, 1933c #15768}.


Marius von Senden (DE) reported that if congenital cataracts in children were removed then replaced by glass lenses the return to vision was long and arduous even though the retina seemed perfectly normal. Return to normal vision was exceptional {von Senden, 1932 #6145;von Senden, 1960 #6146}.


Fritz Schiff (US) and Hikaru Sasaki (US) established that the "secretor" antigen was inherited in the dominant mode. It was clear that secretor children had to have at least one secretor parent, and that nonsecretor parents did not ever have secretor children; the "secretor" trait was obviously dominant {Schiff, 1932 #24756}.


 Walther Kikuth (DE) introduced atebrin (mepacrine) for the treatment of malaria {Kikuth, 1932 #16836}.


John R. Paul (US) and Walls W. Bunell (US) found that heterophile antibodies, demonstrable in the form of sheep cell agglutinins, were recorded in rather high concentrations in the active stages of 4 cases of infectious mononucleosis. Apart from cases of serum disease, and one notable exception, the authors failed to note this finding in a large series of cases representing a variety of clinical conditions, including cases of Vincent's angina, lymphatic leukemia and other blood dyscrasias. These results pointed to an obvious clinical test for the diagnosis of infectious mononucleosis, the Paul-Bunell test {Paul, 1932 #23353}. Note: A heterophile antibody applies to antibodies having the capacity to react with certain antigens, which are quite different from, and phylogenetically unrelated to, the one instrumental in producing the antibody response. See, Forssman, 1911.


Clark W. Heath (US), Maurice B. Strauss (US), and William Bosworth Castle (US) formally established the efficacy of iron treatment. They proved that the active substance was iron when patients with hypochromic anemia were administered parenteral iron and showed a proportionate rise in hemoglobin {Heath, 1932 #20318}.


Jan Friedrich Tönnies (DE) developed the multichannel ink-writing EEG machine {Tönnies, 1932 #19076;Tönnies, 1933 #19077}.


Alfred Bielschowsky (DE-US) wrote Die Lähmungen der Augenmuskeln, a standard work on motility disturbances of the muscles of the eye {Bielschowsky, 1932 #13119}.


Peer Johannes Waardenburg (NL) first presented a clinical case characterized by dystopia of song and heterochromy of iris associated with deafness {Waardenburg, 1932 #24519}. Note: now named Waardenburg syndrome


Bayard Taylor Horton (US), Thomas B. Magath (US), and George Elgie Brown (US) reported inflammation of the temporal and other cranial arteries (temperal arteritis) {Horton, 1932 #22615;Horton, 1934 #22616}.


Edgar Alphonso Hines, Jr. (US) and George Elgie Brown (US) introduced a test designed to detect latent states of hypertension {Hines, 1932 #13135}. It became known as the Hines-Brown Test.


Albert Solomon Hyman (US) is credited with originating the concept of the artificial heart pacemaker. He stimulated the heart in animal experiments by inserting a transthoracic needle into the right ventricle, the other end of which was attached to a mechano-electrical pacemaker. He later employed the same technique in a human patient {Hyman, 1932 #11330}.

John Alexander Hopps (CA) devised a cardiac pacemaker. Hopps was trained as an electrical engineer at the University of Manitoba and joined the National Research Council in 1941, where he conducted research on hypothermia. While experimenting with radio frequency heating to restore body temperature, Hopps made an unexpected discovery: if a heart stopped beating due to cooling, it could be started again by artificial stimulation using mechanical or electric means.

Wilfred Gordon Bigelow (CA), John C. Callaghan (CA), and John Alexander Hopps (CA) developed and tested the first artificial cardiac pacemaker for human use {Bigelow, 1950a #10364;Callaghan, 1951 #19232}. This device was far too large to be implanted inside of the human body. It was an external pacemaker.

Paul Maurice Zoll (US) used the application of electric stimuli from an artificial external cardiac pacemaker by way of subcutaneous needle electrodes to produce effective ventricular beats in 2 patients with ventricular standstill after complete heart block {Zoll, 1952 #7914}. Some consider this the invention of the cardiac pacemaker. Dr. Zoll's later studies showed that externally applied, alternating current counter shocks are similarly effective in stopping ventricular fibrillation, and also in correcting several other serious, potentially lethal, abnormalities of rhythm. He developed the theory and technique of continuous cardiac monitoring of heart rhythm and was the first to apply this method clinically. See, James R. Jude, 1961.

Earl E. Bakken (US), in 1957, at the suggestion of Clarence Walton Lillehei (US) developed the world's first transistorized, battery-powered, external, wearable cardiac pacemaker. Very shortly thereafter Clarence Walton Lillehei (US), Vincent L. Gott (US), Paul Chesley Hodges (US), and David M. Long (US) installed this pacemaker on a patient to treat complete atrioventricular dissociation {Lillehei, 1960 #19078}. This gave patients mobility and eliminated concerns about a power failure. Bakken later formed the Medtronics Corporation, which developed the first fully implantable, self-contained, transistorized, wearable, battery operated pacemaker in 1960.

Rune Elmqvist (SE) developed the first fully implantable pacemaker. Ake Senning (SE), on 10/8/1958, implanted this devise into Arne H.W. Larsson (SE) who suffered from up to twenty fainting attacks per day {Elmqvist, 1960 #19233}. Mr. Larsson died on 12/38/2001.

William Chardack (US), Andrew Gage (US), and Wilson Greatbatch (US), devised a workable transistorized, implantable, pacemaker using primary cells as a power source. It was known as the Chardack-Greatbatch implantable pacemaker. It was the first to be implanted in a patient with AV block {Greatbatch, 1960 #16837}.

Yves Bouvrain (FR) and Fred I. Zacouto (FR) described a combination of devices they called a “resuscitation device”. This consisted of a heart monitor, a defibrillator, and a pacemaker {Bouvrain, 1961 #19246}.

Orestes Fiandra (UY) and Roberto Rubio (UY), in February 1960, inserted a totally implantable pacemaker into a 34-year-old patient with AV block. Its battery life was approximately 12-18 months {Fiandra, 1988 #19079}.

David A. Nathan (US), Sol Center (US), Chang-You Wu (US), and Walter Keller (US) reported the first implantable atrial-synchronous ventricular pacemaker (VAT), which sensed atrial activity and paced the ventricle accordingly {Nathan, 1963 #19234}.

Heinz-Joachim Sykosch (DE), Sven Effert (DE), Karl Georg Pulver (DE), and Fred I. Zacouto (FR) introduced the ventricular demand pacemaker (VVI) {Sykosch, 1963 #19235}.

Philippe Coumel (FR), Christian Cabrol (FR), Alexandre Fabiato (US), René Gourgon (FR), and Robert D. Slama (US) used programmed atrial and ventricular stimulation to unravel the mechanism and diagnosis of permanent junctional reciprocating tachycardia in a drug-refractory patient. They showed that tachycardia was sustained by a reentrant or reciprocal rhythm, then postulated that the process involved dual AV nodal pathways with a slow conducting retrograde pathway. They installed a permanent bipolar pacemaker in such a way as to relieve the tachycardia {Coumel, 1967 #19255}.

Barouh V. Berkovits (US), Agustin Castellanos, Jr. (US), Louis Lemberg (US), George Callard (US), and James R. Jude (US) introduced the bifocal pacemaker {Berkovits, 1971 #19236}.

Mieczyslaw Mirowski (PL-IL-US), Morton M. Mower (US), Alois A. Langer (US), and Marlin Stephen Heilman (US) designed the first automatic implantable defibrillator {Mirowski, 1973 #11347;Langer, 1976 #11348}.

Mieczyslaw Mirowski (PL-IL-US), Philip R. Reid (US), Morton M. Mower (US), Levi Watkins, Jr. (US), Vincent L. Gott (US), James F. Schauble (US), Alois A. Langer (US), Marlin Stephen Heilman (US), Steven A. Kolenik (US), Robert E. Fischell (US), and Myron L. Weisfeldt (US) implanted the first automatic implantable cardioverter/defibrillator into a human. This was done to terminate a malignant ventricular arrhythmia {Mirowski, 1980 #11349}.

Hermann D. Funke (DE) introduced the first DDD pacemaker, a device that senses intrinsic activity in the atrium as well as the ventricle, and can likewise pace both chambers {Funke, 1978 #19237}.

Benjamin Befeler (US), Barouh V. Berkovits (US), Juan M. Aranda, Jr. (US), Ruey J. Sung (US), Federico Moleiro (VE), and Agustin Castellanos, Jr. (US) performed a biventricular stimulation in humans by using a QRS triggered pacemaker connected by way of the great or middle cardiac veins {Befeler, 1979 #19238}.

Alexander Wirtzfeld (DE), Thomas Bock (DE), Anthony Francis Rickards (GB), Fawaz Akhras (GB), and David W. Barron (GB) were among the first to introduce a pacemaker capable of responding to a changing biological variable {Rickards, 1979 #19244;Wirtzfeld, 1978 #19243}.

Eric F.D. Wever (NL), Richard N.W. Hauer (NL), Frans J.L. Capelle (NL), Jan G.P. Tijssen (NL), Harry J.G.M. Crijns (NL), Ale Algra (NL), Ans C.P. Wiesfeld (NL), Patricia F.A. Bakker (NL) and Etienne O. Robles de Medina (NL) demonstrated the clinical benefit of biventricular pacing in patients who have survived heart failure {Wever, 1995 #19239}.

Werner Jung (DE) and Berndt Lüderitz (DE) successfully installed an implantable atrial defibrillator in a 64-year old female patient with symptomatic, drug-refractory atrial fibrillation {Jung, 1996 #19250}.

Werner Jung (DE) and Berndt Lüderitz (DE) successfully installed an implantable atrioventricular defibrillator in a 61-year-old female patient. This device automatically detects atrial and ventricular signals and delivers electrical therapy in the appropriate chamber to terminate the arrhythmia {Jung, 1997 #19251}.


Paul Chevalier (FR) and Jean Jacques Robert Bernard (FR) gave the first description of the use of high dosage radiotherapy in the treatment of Hodgkin's disease {Chevalier, 1932 #18745}.


Walter Edward Dandy (US) performed a partial resection of the sensory root of the trigeminal ganglion as treatment of trigeminal neuralgia {Dandy, 1932 #22611}.


Antoine Marcellin Bernard Lacassagne (FR) demonstrated that addition of the synthetic exogenous hormone folliculin (estrone benzoate) can induce cancer {Lacassagne, 1932 #23775}.

Edward Charles Dodds (GB) discovered diethylstilbestrol (DES), a powerful synthetic hormone used to treat prostate conditions, to fatten cattle, to treat women at risk for miscarriage and as a morning-after contraceptive {Dodds, 1938 #10076;Dodds, 1953 #19152}.

Wilhelmina F. Dunning (US) and Maynie R. Curtis (US) found that diethylstilbestrol (DES), a synthetic exogenous hormone, can induce cancer in rats {Dunning, 1952 #23776}.

Arthur L. Herbst (US), Howard Ulfelder (US), and David C. Poskanzer (US) conducted a retrospective investigation which revealed a linkage between adenocarcinoma of the vagina among young women and mothers who received diethylstilbestrol (DES) therapy during the first trimester of pregnancy {Herbst, 1971 #23807}.

Arthur L. Herbst (US), Robert J. Kurman (US), and Robert E. Scully (US) showed that the prescribed form of diethylstilbestrol (DES) was proved carcinogenic in adult women as well as in fetuses, when unusual types of endometrial cancer, reminiscent of the adenocarcinomas of the vagina of DES daughters, developed in young women treated with DES for five years or longer {Herbst, 1972 #25188}.

J.G. Stolk (NL), G. Peter Vooijs (NL), E.J. Aartsen (NL), and A. Peter M. Heintz (NL) reported on the teratogenic effect of diethylstilbestrol (DES) during pregnancy and the extent of the DES problem in the Netherlands {Stolk, 1982 #12463}.


 John Burdon Sanderson Haldane (GB) points out the importance of knowledge about the age/stage of gene expression - gametophytes and gametes to zygotes, embryos and immature and mature organisms in evolutionary studies {Haldane, 1932 #20454}.


Warder Clyde Allee (US) and Edith S. Bowen (US) introduced what would later be called the Allee effect. A phenomenon in biology characterized by a correlation between population size or density and the mean individual fitness (often measured as per capita population growth rate) of a population or species {Allee, 1932 #26877;Odum, 1953 #26878}. Note: This led them to conclude that aggregation can improve the survival rate of individuals, and that cooperation may be crucial in the overall evolution of social structure. The term Allee principle was introduced in the 1950s



“If all the arts aspire to the condition of music, all the sciences aspire to the condition of mathematics.” George Santayana {Santayana, 1933 #15306}.


“It is common sense to take a method and try it. If it fails, admit it frankly, and try another. But by all means, try something.” Franklin D. Roosevelt, Fireside Chat, March 12, 1933.


“The fundamental activity of medical science is to determine the ultimate causation of disease.” Wilfred Batten Lewis Trotter {Trotter, 1933 #22825}.


"The facts as revealed have just that degree of unexpectedness—if I may use the phrase—which was to be expected in a biochemical phenomenon. I often find myself compelled to assert that, though biochemical events are, of course, limited by chemical possibilities, they are not safely to be predicted by chemical probabilities, even when these are strong." Hans Adolf Krebs (DE){Hopkins, 1933 #25554}.


Thomas Hunt Morgan (US) was awarded the Nobel Prize in Physiology or Medicine for his discoveries concerning the role played by the chromosome in heredity.


Niels Henrik David Bohr (DK) introduced the idea of complementarity to biology. All properties of physical entities exist only in pairs, which Bohr described as complementary or conjugate pairs (which are also Fourier transform pairs). Physical reality is determined and defined by manifestations of properties, which are limited by trade-offs between these complementary pairs {Bohr, 1933 #14668}.


John Desmond Bernal (GB) and Ralph Fowler (GB) created their model of the structure of liquid water {Bernal, 1933 #19766}.

John Desmond Bernal (GB) and Helen Dick Megaw (IE) introduced the notion that hydrogen atoms in bulk liquid water can jump in concerted fashion between the two oxygen atoms that they link together {Bernal, 1935 #19767}.


A salt of dinitrophenol was used first in France in 1933 for control of annual forbs in cereals {Westgate, 1940 #24370}.


Roger J. Williams (US), Carl Morris Lyman (US), George H. Goodyear (US), John H. Truesdail (US), and Duncan Holaday (US) discovered and partially synthesized pantothenic acid (vitamin B5) {Williams, 1933 #3125}.

Roger John Williams (US), Carl M. Lyman (US), George H. Goodyear (US), John H. Truesdail (US), Duncan Holaday (US), Donald Herbert Saunders (US), Harry H. Weinstock, Jr. (US), Ewald Rohrmann (US), Hershel K. Mitchell (US), Curtis B. Meyer (US), and Charles H. McBurney (US) separated and named pantothenic acid (Greek pantos or pan meaning ubiquitous), vitamin B5, from other components of extracts of natural foods, secured a concentrated preparation of the acid, validated it as a vitamin using a yeast strain, found its approximate molecular weight and ionization constant, and further characterized its chemical properties. They isolated it as a pure calcium salt of the acid {Williams, 1933 #3125;Williams, 1934 #3126;Williams, 1935 #3127;Williams, 1939a #23383;Williams, 1939b #3128}.


Reginald William Herbert (GB), Edmund Langley Hirst (GB), Edmund George Vincent Percival (GB), Reginald John William Reynolds (GB), and Fred Smith (GB) determined the structure of ascorbic acid (vitamin C) {Herbert, 1933 #1727}.

Tadeus Reichstein (PL-CH), Andreas Grüssner (CH), Rupert Oppenauer (CH), R.G. Ault (GB), D.K. Baird (GB), H.C. Carrington (GB), Walter Norman Haworth (GB), Reginald William Herbert (GB), Edmund Langley Hirst (GB), Edmund George Vincent Percival (GB), Fred Smith (GB) and Maurice Stacey (GB) synthesized ascorbic acid (vitamin C) {Reichstein, 1933a #1725;Reichstein, 1933b #1726;Reichstein, 1933c #1724;Ault, 1933 #12212}.


André Félix Boivin (FR) Ion Mesrobeanu (RO), and Lydia Mesrobeanu (RO) developed a method for isolating endotoxin {Boivin, 1933 #8547}.

André Félix Boivin (FR) discovered that the somatic antigen (endotoxin) is present in all Enterobacteriaceae regardless of their pathogenicity {Boivin, 1946 #162}.


Rudolf Schoenheimer (DE-US) and Fritz Breusch (DE-US) concluded that cholesterol is continually destroyed as well as formed in the animal body {Schoenheimer, 1933 #17272}.


Karl Lohmann (DE) discovered glucose-6-phosphate isomerase (phosphoglucose isomerase), one of the phosphoglucose enzymes, which were subsequently found to catalyze the breakdown of sugar phosphates according to the glycolytic scheme. Typically, it catalyzes the reciprocal conversion of D-glucose 6-phosphate to D-fructose 6-phosphate {Lohmann, 1933 #20834}.


Morna MacLeod (GB) and Robert Robison (GB) isolated fructose-1-phosphate from the partial hydrolysis of fructose diphosphate by bone phosphatase {MacLeod, 1933 #1719}.


Jakov Borissovich Goldman (RU) developed a method for staining lipids in histologic slides and peripheral blood films using Sudan III {Goldman, 1933 #13287}.


William Smith Tillett (US) and Raymond L. Garner (US) found that certain strains of hemolytic streptococci produce a substance capable of inciting the rapid fibrinolysis of human plasma clots. They named the lytic agent fibrinolysin {Garner, 1934a #19083;Garner, 1934b #19084;Tillett, 1933 #7079;Tillett, 1934 #10397}. This represents the beginning of investigations of streptokinase.

Haskell Milstone (US) suggested that a plasma factor, which he called plasma lysing factor, is responsible for streptococcal mediated fibrinolysis {Milstone, 1941 #11316}.

L. Royal Christensen (US) and Colin Munro MacLeod (US) were able to describe the entire mechanism of streptococcal fibrinolysis. They showed that human plasma contains the precursor of an enzyme system, which they called plasminogen, and that the streptococcal fibrinolysin, which they named streptokinase, is an activator which can convert plasminogen to the proteolytic and fibrinolytic enzyme plasmin {Christensen, 1945a #7083;Christensen, 1945b #10395;Christensen, 1954 #7084}.

Stuart D. Elliott (US) was the first to extract streptokinase and its proenzyme (zymogene) in the crystallized form {Elliott, 1950 #10396}.

William Smith Tillett (US), Alan J. Johnson (US), and W. Ross McCarty (US) achieved an intravascular lytic state in man following streptokinase administration {Tillett, 1955 #10405}.

Sol Sherry (US) started using streptokinase in patients with acute myocardial infarction and changed the focus of treatment from palliation to “cure” {Sherry, 1981 #19613}.

Streptokinase (SK) has been used in medicine as fibrinolytic enzyme system - plasminogenous activator. SK is used for myocardial infarctions, vein thrombosis, lung artery emboli, occlusions of extremity arteries, retinal blood-vessel thrombosis and priapism.


Maurice W. Goldblatt (GB) and Ulf Svante Hansson von Euler-Chelpin (SE) independently discovered that extracts of human semen, monkey, sheep and goat seminal vesicular glands respectively cause contraction of smooth muscle in vitro and sharp decreases in the blood pressure in experimental animals {Goldblatt, 1933 #11250;Goldblatt, 1935 #11251;von Euler-Chelpin, 1935 #11252;von Euler-Chelpin, 1934 #13713;von Euler-Chelpin, 1983 #3857}. Ulf Svante Hansson von Euler-Chelpin called these substances prostaglandins because they were mistakenly believed to be made in the prostate gland.


Lárus Einarson (DK) suggested that the chromatic material of the Nissl Granules or Bodies in neurons is formed around the nucleolus and then diffuses out into the cytoplasm {Einarson, 1933 #5596}.


Paul Runar Collander (FI) and Hugo Bärlund (FI) made quantitative measurements of cell membrane permeability to non-electrolytes of varying molecular size and lipid solubility. Their results contributed enormously to our understanding of membrane structure {Collander, 1933 #19081}.


George Wald (US) found vitamin A (retinol) in the retina {Wald, 1933 #9213;Wald, 1935 #2953}.

George Wald (US) discovered retinene (retinal), an intermediate in the bleaching of rhodopsin, on the way to vitamin A (retinol). He concluded that rhodopsin in the retina, under the influence of light, engages in a cycle of reactions with retinene and vitamin A {Wald, 1934 #9215;Wald, 1935 #9216}.

George Wald (US) and Anna-Betty Clark (US) presented evidence that visual purple is a conjugated protein in which vitamin A (retinol) is a prosthetic group {Wald, 1935 #2953;Wald, 1936 #2954}.

George Wald (US) discovered that visual pigment from freshwater fish differs from that of mammals, birds, amphibia, and marine fish. He named it porphyropsin {Wald, 1937a #9224;Wald, 1939 #9225}.

George Wald (US), Paul K. Brown (US), and Patricia H. Smith (US) determined that all known visual pigments are built upon a common plan. Retinal is bound as chromatophore to a type of protein, called an opsin, found in the outer segments of vertebrate rods and cones and the analogous rhabdomeres of invertebrates. There are four major pigments known in vertebrate vision: 1) retinal 1 + rod opsin = rhodopsin, 2) retinal 1 + cone opsin = iodopsin, 3) retinal 2 + rod opsin = porphyropsin, and 4) retinal 2 + cone opsin = cyanopsin {Wald, 1937b #9226;Wald, 1953 #9228;Wald, 1955 #9227}.

Sadie Brenner (US), Lydia J. Roberts (US), John Elliott Dowling (US), and George Wald (US) demonstrated that night blindness is the earliest manifestation of vitamin A (retinol) deficiency {Brenner, 1943 #1972;Dowling, 1958 #1973}.

S. Ball (GB), Trevor Walworth Goodwin (GB), and Richard Alan Morton (GB) showed that retinine 1 is vitamin A aldehyde {Ball, 1948 #9218}. Retinene was later renamed retinal and vitamin A renamed retinol.

George Wald (US) and Ruth Hubbard (US) demonstrated that the visible pigment (rhodopsin) from the cones of the eye contains a protein (opsin) in combination with retinene (retinal). Retinene is very similar in structure to vitamin A (retinol) and is formed from vitamin A in the body. When light strikes rhodopsin, the protein and the retinene separate; they recombine in the dark {Hubbard, 1951 #19101;Wald, 1950 #19100}.

Ruth Hubbard (US) and George Wald (US) identified the initial molecular event in vision by showing that when 11-cis retinal absorbs a photon, it is converted to the all-trans form {Hubbard, 1952 #14956}.

Paul K. Brown (US) and George Wald (US) demonstrated that two different proteins, two opsins, are needed to form the red- and green-sensitive pigments suggesting that two genes are involved in red- and green-blindness {Brown, 1963 #9234}.

William B. Marks (US), William H. Dobelle (US), Edward F. MacNichol (US), Paul K. Brown (US), and George Wald (US) found that primate retinas possess, in addition to rod cells with their rhodopsin, three kinds of cone cells, blue-, green- and red sensitive, each containing predominantly or exclusively one of three color pigments with maximum absorption at 435, 540, and 565 micrometers respectively {Marks, 1964 #9231;Brown, 1964 #9232}.

George Wald (US) predicted that light activated rhodopsin might trigger a cascade of reactions much like the blood clot cascade {Wald, 1965 #10436}.

George Wald (US) discovered the primary event in vision to be when light triggers visual excitation by isomerizing the 11-cis retinal chromophore of visual pigments to the all-trans form {Wald, 1968a #14812;Wald, 1968b #6310;Wald, 1969 #14813}.


Moses Kunitz (RU-US) and John Howard Northrop (US) were the first to crystallize the enzyme chymotrypsin and its precursor chymotrypsinogen. They isolated and crystallized a new protein from the pancreas. It separated as elongated prisms and had no proteolytic action. However, when acted upon by a trace of active trypsin it was converted into a second protein, crystallizing in plates, which had a proteolytic activity about a third as great as crystalline trypsin-1. The new enzyme had less hydrolytic action on gelatin than trypsin-1 but had a powerful action on coagulating milk. They called the inactive form chymotrypsinogen and the trypsin activated form chymotrypsin {Kunitz, 1933 #1953;Kunitz, 1935 #5545}. They also isolated and crystallized trypsinogen, a trypsin inhibitor, and an inhibitor-trypsin compound {Kunitz, 1936 #5546}.


Richard Johann Kuhn (AT-DE), Paul György (HU-DE-GB-US), and Theodor Wagner-Jauregg (DE) discovered riboflavin (vitamin B2 or vitamin G) {Kuhn, 1933a #14740;Kuhn, 1933b #18141}.


Franklin E. Allison (US), Sam R. Hoover (US), and Dean Burk (US) isolated a vitamin, which is an indispensable nutrient for Rhizobium spp. They named the vitamin coenzyme R (biotin) {Allison, 1933 #3114}.

Fritz Kögl (NL) and Benno Tönnis (NL) isolated and crystallized a vitamin from the boiled yolks of duck eggs. They named it biotin {Kögl, 1936 #3112}.

Paul György (HU-DE-GB-US) and Thomas William Birch (US) isolated a vitamin in pure form, which would cure raw egg white injury (harm done when raw egg whites are the sole protein source in a diet). They named it vitamin H (biotin) {György, 1939 #11366;Birch, 1939 #3113}.

Paul György (HU-DE-GB-US), Donald B. Melville (US), Dean Burk (US) and Vincent du Vigneaud (US) proved that vitamin H, biotin, and coenzyme R are one and the same substance {György, 1940a #3111}.


Carl Alexander Neuberg (DE) and Maria Kobel (DE) demonstrated that phosphoglyceric acid is converted to pyruvic acid and phosphoric acid by Lactobacillus delbruckii {Neuberg, 1933 #11204}.


Dietrich Hans Franz Alexander Bodenstein (DE-US), working with caterpillars of the butterfly Vanessa urticae, concluded that the time course of molting is not determined by autonomous changes in the hypodermis, but rather that factors situated elsewhere within the caterpillar, most likely blood borne hormones, determine the time course of molting {Bodenstein, 1933 #10782}.


Evelyn Mary Anderson (CA) and James Bertram Collip (CA) were the first to successfully isolate a tropic substance. It was thyrotropic hormone (TSH) from the anterior pituitary gland {Anderson, 1933 #6708}.

James Bertram Collip (CA), Evelyn Mary Anderson (CA), and David Landsborough Thomson (GB-CA) prepared and tested extracts from the anterior pituitary gland and found that they contained a potent adrenotropic hormone (ACTH) {Collip, 1933 #3405}.

James Bertram Collip (CA), János Hugo Bruno “Hans” Selye (AT-HU-CA), and David Landsborough Thomson (GB-CA) purified a highly potent extract of growth hormone (somatotropic hormone/STH) from the anterior pituitary lobe {Collip, 1933 #6870}.


Otto Fritz Meyerhof (DE-US) and Wilhelm Kiessling (DE) isolated alpha-glycerophosphoric acid from a reaction mixture of animal muscle and either glycogen or hexose-phosphate {Meyerhof, 1933a #1721;Meyerhof, 1933b #1723;Meyerhof, 1933c #2421;Meyerhof, 1933d #12883}.

Otto Fritz Meyerhof (DE-US) and Wilhelm Kiessling (DE) published a detailed study of the intermediary phases of fermentation in yeast juice {Meyerhof, 1933c #2421}.

Otto Fritz Meyerhof (DE-US) worked out the chemical scheme of the cellular breakdown of sugar into alcohol {Meyerhof, 1933d #12883}.

Gustav Georg Embden (DE), Hans-Joachim Deuticke (DE), and Gert Kraft (DE) proposed an anaerobic glycolytic scheme within muscle cells which begins with the conversion of hexosediphosphate into triosephosphate which, by the oxidation reduction process, yields alpha-phosphoglycerol plus 3-phosphoglyceric acid; phosphoglyceric acid breaks down to pyruvic and phosphoric acids. In muscle extract, pyruvic acid is reduced to lactic acid at the expense of the phosphoglycerol, which is oxidized to triosephosphate. In yeast juice, pyruvic acid is converted by carboxylase into carbon dioxide plus acetylaldehyde. The latter was then believed to take part with glucose and inorganic phosphate in a rapid reaction in which hexosediphosphate plays the part of a catalyst and phosphorylation is coupled with an oxidation-reduction; the primary esterification product is oxidized to phosphoglyceric acid while the acetylaldehyde is reduced to alcohol {Embden, 1933 #1720}. This evidence suggested that it is pyruvic rather than lactic acid, which represents the true end product of anaerobic glycolysis.

Jakub (Jacob) Karol Parnas (PL), Pawel Ostern (PL), and Thaddeus Robert Rudolph Mann (PL-GB) resolved that, “… the resynthesis of phosphocreatine and adenosine triphosphate (ATP) is not linked to glycolysis as a whole, but to definite partial processes: and this leads further to the conclusion that this resynthesis does not involve a relationship that might be termed energetic coupling, but more probably involves a transfer of phosphate residues from molecule to molecule.” In this article they describe for the first time the presence of pyruvate kinase (phosphoenol transphorylase) and the fact that it catalyzes the magnesium- and potassium-dependent transphorylation between phosphoenolpyruvate and ADP {Parnas, 1934 #2567}.

Otto Fritz Meyerhof (DE-US) offered proof that, in isolated but otherwise intact frog muscle, the lactic acid formed is reconverted to carbohydrate in the presence of oxygen. He also prepared a KCl extract of muscle, which could carry out all the steps of glycolysis with added glycogen and hexose-diphosphate in the presence of hexokinase derived from yeast. In this system glucose was also glycolysed and this was the foundation of the Embden-Meyerhof-Parnas theory of glycolysis (glyco, sugar; glykis, sweet). The specific sequence of reactions from glucose to pyruvate is often called the Embden-Meyerhof-Parnas pathway {Meyerhof, 1935a #2548;Meyerhof, 1935b #9264;Meyerhof, 1935c #19614;Meyerhof, 1935d #19615;Meyerhof, 1935e #20835}. Meyerhof in the 1935c article gave the first description of phosphoglyceric acid mutase (phosphoglycerate mutase).

Hermann Lehmann (DE-GB) offered evidence that phosphoenolpyruvate plus ADP can be converted to pyruvate plus ATP and the reverse {Lehmann, 1935 #24391}.

Otto Fritz Meyerhof (DE) and Wilhelm Kiessling (DE) discovered triose-phosphate isomerase in muscle {Meyerhof, 1935f #24392}.

Pawl Ostern (PL), J.A. Guthke (PL), and J. Terszakowee (PL), in muscle brei, found 6-phosphofructokinase, which catalyzes the transfer of phosphate from ATP to D-fructose 6-phosphate {Ostern, 1936 #24393}.


Ernst Wolfgang Caspari (DE-US), working with the flour moth Ephestia kuhniella Zeller, provided an example of how a gene can control hormone action during development {Caspari, 1933 #7232;Caspari, 1971 #7233}.


John Belling (GB-US) states that “crossing over” exists apparently in all those flowering plants, which have been sufficiently investigated with regard to it. Billing’s modification of Janssens' hypothesis explains crossing over, and explains gene rearrangements, such as reversed crossing over, reciprocal translocation, inversion, deletion, and deficiency. The chromonemas were proved to not be split at leptotene in certain plants. Living (and fixed) chromonemas of resting “final” nuclei showed no split, in the plants examined. The secondary split was first seen at mid-pachytene. Both direct and oblique chiasmas were seen at pachytene in Lilium. After the chromomeres have split, the old longitudinal fibers are either alone visible; or are seen to be thicker than the new ones. In Lilium the opening-out at diplotene seems to be only at the primary split. In plants such as Datura, with no chiasmas at diaphase, it is probable that the diplotene opening-out alternates at the chiasmas. Since chiasmas arise at pachytene in certain liliaceous plants, they cannot arise from alternate opening-out at diplotene. Chiasmas seem to be due to overlaps, not twists. Overlaps may be sometimes mistaken for twists, under the microscope. There are 8 main kinds of double chiasmas, equally numerous by chance. Double chiasmas give, by chance, one non-crossover chromosome, two single-crossover chromosomes, and one double-crossover chromosome. If crossovers arise from chiasmas, then the distal recombinations from the end to the fusal chromomere should be 50 percent. If crossovers arise from chiasmas, then the chart crossovers divided by 50 should give the average number of chiasmas. The ascertained numbers of crossover X chromosomes of Drosophila melanogaster appear to lack about 7 percent of single crossovers, and about 2 percent of double crossovers, if they arose from chiasmas (neglecting triple crossovers). Flies with heterozygous attached X’s in Drosophila, should (on the Billing’s theory) give distal recessive homozygotes in a percentage equal to half the chart length minus one and a half times the percentage of double crossovers. This would be 17.5. In Belling's theory, the percentages of complementary and identical non-crossovers, of crossovers plus non-crossovers, and of complementary crossovers, in attached X’s, have been calculated from the chromosome chart. It is possible to explain reversed crossing over, heterologous interchange, terminal translocation, inversion, deletion, and deficiency, by the overlapping of two chromonemas when their chromomeres are dividing. The result is equivalent to the formation of a chiasma between synapsed homologues; but is less regular, so that genes may be lost at the junctions {Belling, 1933 #22114}.


Bryan H.C. Matthews (GB) showed that there exist in the mammal two distinct nerve-muscle systems in the skeletal musculature. The large motor nerve fibers set up the familiar motor unit twitch responses and the small motor nerve fibers increase the sensory discharges from muscle spindles, which consist of a number of so-called intrafusal muscle fibers {Matthews, 1933 #9413}.

Ichiji Tasaki (JP) and K. Mizutani (JP) found that extrafusal muscle fibers in amphibia are innervated by two distinct motor systems. Motor neurons with large axons cause the familiar large fast twitches with single stimuli, known as the twitch system, while motor neurons with small axons require repetitive stimulation to cause slow and relatively weak contractions, known as the tonic system {Tasaki, 1944 #21249}.

Bernard Katz (GB), in his studies of the frog, observed that stimulation of large, low-threshold, motor axons not only caused extrafusal contraction but also a short burst of afferent impulses. The afferent firing persisted when extrafusal contraction was blocked by critical dosage with the muscle relaxant curare, thus showing that the large motor axons branched to innervate intrafusal muscle fibres. When muscle shortening was allowed, the tendency of extrafusal contraction to silence the spindle was offset by the intrafusal contraction. Katz discussed the significance of this clearly, with the proposal that when an extended muscle is contracted actively in life, the inevitable simultaneous intrafusal contraction would ensure that afferent activity continues, which would support the contraction against loading by means of the stretch reflex {Katz, 1949 #21247}.


Richard Benedikt Goldschmidt (DE-US) interbred various geographical races of the gypsy moth, Lymantria dispar. The moths that resulted from this cross showed that characters distinguishing local varieties are transmitted to the offspring and, therefore, can be explained in terms of Mendelian laws as being determined by genes. This was the first genetic explanation of geographic variety {Goldschmidt, 1933 #12345}.


Rudolf Kaufmann (DE) studied the Upper Cambrian alum shales in Sweden, there he found that the trilobite genus Olenus occurred in an unbroken sequence of sediments covering a considerable period of geological time. He was thereby in a position to track the phylogenetic evolution of Olenus, that is, the rise and fall of species within the genus and the changes in their morphology. He coined the idea of species modification (Artabwandlung), which is the tendency of clade elements in the same environment to show the same morphological trends {Kaufmann, 1933 #26936}.


Teikichi Fukushi (JP) was the first to provide experimental evidence of plant virus multiplication in insects. He was also the first to demonstrate transmission of a plant virus through the eggs of the vector (transovarial passage) {Fukushi, 1933 #5727;Fukushi, 1935 #8549;Fukushi, 1940 #8550}.


Richard E. Shope (US) and E. Weston Hurst (US) discovered the cottontail rabbit papillomavirus (CRPV) {Shope, 1933 #8249}.


Ralph S. Muckenfuss (US), Charles Armstrong (US), and Howard A. McCordock (US) proved that the etiological agent of St. Louis encephalitis was a virus. They successfully infected monkeys with the virus of St. Louis encephalitis by intracerebral inoculation with human brain tissue from a patient who had succumbed to the disease {Muckenfuss, 1933 #6979}.


William W. Dimock (US), Phillip R. Edwards (US), Elvis R. Doll, Jr. (US), and John H. Kintner (US) discovered equid herpes virus 1 (EHV-1; equine abortion virus) and equid herpesvirus 4 (EHV-4; equine rhinopneumonitis virus) {Dimock, 1933 #25953;Doll, 1954 #25954}.


Fred L. Soper (US), Henrique de Azevado Penna (BR), Eleyson Cardoso (BR), Jose Serafim (BR), Martin Frobisher, Jr. (BR), J. Pinheiro (BR), Raymond C. Shannon (US), Loring Whitman (US), Mario Franca (BR), John C. Burgher (US), Jorge Boshell-Manrique (BR), and Manuel Roca-Garcia (BR) discovered the jungle cycle of yellow fever virus involving Haemagogus species in the Americas and as Aedes species in Africa {Burgher, 1944 #25955;Shannon, 1938 #25956;Soper, 1933 #25957}.


Frits Mari Muller (NL) reported on the anaerobic use of organic sources of reducing power by the purple sulfur photosynthetic bacteria. Growth was accompanied by the production or utilization of CO2 depending on the ‘redox level’ of the organic substrate {Muller, 1933 #14743}.


Ernest Witebsky (DE-US) and Werner Henle (DE) discovered that bacteria indistinguishable morphologically, and in culture, can be subdivided by immunological tests {Witebsky, 1933 #107}.


Arthur T. Hendrici (US) noted the tendency of aquatic bacteria to colonize submerged surfaces {Hendrici, 1933 #108}.


Robert E. Foster (US) and Carlton Earl Burnside (US) described a new disease found within broods of the honeybee, Apis mellifera Linn., which they named parafoulbrood {Foster, 1933 #7029;Burnside, 1933 #25081}. In 1935, they named and identified the etiological agent Bacillus para-alvei.


S. Tanaka (JP) discovered that black spot of Japanese pear is due to a host-specific toxin {Tanaka, 1933 #23122}. Tanaka reported a heat-labile, host-selective factor produced by Alternaria kikuchiana that when sprayed on plants produced lesions typical of black spot.

Frances Meehan (US) and Hickman C. Murphy (US) discovered that Victoria blight of oat is also the result of a host-specific toxin. They reported a metabolic by-product of Helminthosporium victoria, which proved toxic to a susceptible oat cultivar. This heat-stable toxin was found in hyphae and in the growth substrate; symptoms in oat were caused by the toxin {Meehan, 1947 #23123}. These two diseases became models for further study of phytotoxins.


Alexander Ivanovitch Petrunkevitch (RU-US) wrote, An Inquiry Into the Natural Classification of Spiders, Based on a Study of Their Internal Anatomy, which is the first comparative anatomy for any group of arthropods that can be used in classification {Petrunkevitch, 1933 #10745}.


Johannes Friedrich Karl Holtfreter (DE-US)) performed experiments where gastrula tissue from young embryos was transplanted into ectoderm of older embryos. He found that the reacting tissue developed according to its surroundings. The conclusion is that the whole is controlling the events in its parts {Holtfreter, 1933a #1110;Holtfreter, 1933b #1111;Holtfreter, 1936 #1112}.

Johannes Friedrich Karl Holtfreter (DE-US) performed exogastrulation studies wherein the dorsal mesoderm failed to contact the overlying ectoderm. In these instances, the ectoderm did not acquire a neural character, again suggesting that the inducing signal appeared to be transmitted vertically from the mesoderm to the ectoderm {Holtfreter, 1933d #19971}.

Conrad Hal Waddington (GB), Noel Joseph Terence Montgomery Needham (GB), Dorothy Moyle Needham (GB), Wiktor W. Nowinski (US), and Max Rudolf Lemberg (DE) showed that the ether extracts of adult newts could act as an organizer. Since this activity could turn presumptive epidermis into non-specific neural tissue. Waddington referred to this substance as evocator {Waddington, 1935a #19973;Waddington, 1935b #19974}.

Conrad Hal Waddington (GB), Noel Joseph Terence Montgomery Needham (GB), and Jean Louis Auguste Brachet (BE) hypothesized that the evocator substance was produced throughout the embryo, but it was just released or activated in one particular region {Waddington, 1936 #19975}.

Jean Louis Auguste Brachet (BE), Taina Kuusi (FI), and Simone Gothie (BE) suggested that movements of microsomes/ribosomes (which contain ribonucleic acid) from the archenteron roof to the overlying ectoderm are involved in neural induction {Brachet, 1952 #14868}.

Lauri Saxén (FI) demonstrated that neural induction could occur through a 150-micron thick, 0.8-micron pore size filter, strongly suggesting that the inducer was diffusible {Saxén, 1961 #19972}.

Scott F. Gilbert (US) reported that follistatin, chordin, noggin, Xenopus nodal-related-3 promoter, and cerberus all have organizer function {Gilbert, 1997 #19976}.


Aaron Bodansky (RU-US), Lois F. Hallman (US), and Kissel Bonoff (US) described the factors, which influence accuracy during the determination of serum phosphatase {Bodansky, 1933 #9414}. Elevated levels of these enzymes can be a useful index of abnormal conditions in certain tissues.


Rudolpho Margaria (IT), Harold T. Edwards (US), and David Bruce Dill (US) defined the components of the oxygen debt and described what become known as the anaerobic threshold, i.e., the rate of appearance of lactic acid in the blood and its influence on ventilation during exercise {Margaria, 1933 #7373}.


Karl Landsteiner (AT-US) and James van der Scheer (US), discovered that antibodies can be formed to and subsequently bind with exquisite specificity to completely synthetic compounds {Landsteiner, 1933 #2154;Landsteiner, 1936 #1681}.


Wilbert R. Todd (US), Conrad Arnold Elvehjem (US), and Edwin Bret Hart (US) presented data demonstrating that zinc was essential in the nutrition of the rat. They found that the rate of growth of rats on a synthetic diet low in zinc was accelerated by the addition of salts of this metal. There was also an interference with the development of a normal fur coat of the animals on the zinc-low diet {Todd, 1933 #23391}.

Thore Edvard Brandt (SE) described a zinc deficiency syndrome in infants, characterized by acral dermatitis, alopecia, diarrhea, steatorrhea, and anal pustular eruptions on the face and around body orifices {Brandt, 1936 #13127}. The syndrome is caused by the absence of the ligand essential for zing absorption, which is present in human but not cow milk. A similar disease picture may be seen in patients receiving artificial nutrition with low zinc content. Untreated, the disease is usually lethal.

David Keilin (PL-GB), Thaddeus Robert Rudolph Mann (PL-GB), Carl G. Holmberg (SE), Edwin L. Hove (NZ), Conrad Arnold Elvehjem (US), and Edwin Bret Hart (US) demonstrated that zinc is necessary in animal nutrition because it is a cofactor for certain enzymes {Keilin, 1939b #3093;Keilin, 1940 #23430;Holmberg, 1939 #3094;Hove, 1940 #3095}.


Beatrice J. Geiger (US), Harry Steenbock (US), and Helen T. Parsons (US) described lathyrism in the rat {Geiger, 1933 #23373}.

Lathyrism or neurolathyrism is a neurological disease of humans and other animals, caused by eating certain legumes (e.g. the grass pea) of the genus Lathyrus. The consumption of large quantities of Lathyrus grain containing high concentrations of the glutamate analogue neurotoxin β-oxalyl-L-α,β-diaminopropionic acid (ODAP, also known as β-N-oxalyl-amino-L-alanine, or BOAA) causes paralysis, characterized by lack of strength in or inability to move the lower limbs, and may involve pyramidal tracts producing signs of upper motor neuron damage. The toxin may also cause aortic aneurysm.

Ingestion of legumes containing the toxin results mostly from ignorance of their toxicity but can also be because poor people lack resources to detoxify and may be desperate for food {Schilling, 1954 #23374}.


Lionel Sharples Penrose (GB) was the first to show the significance of the mother’s age in Down’s syndrome. He determined that birth order, parity, and length of interval between pregnancies are not significant etiological factors {Penrose, 1933 #11255}.


Lemuel W. Diggs (US), Chester Frederick Ahmann (US), and Juanita Bibb (US) demonstrated that there is a distinct difference between people with sickle-cell anemia and those who carry a slight sickling "trait" but have none of the other symptoms. They documented the diminished sickling of red blood cells from erythrocytes in newborns and defined the ratio of patients with sickle cell anemia to carriers of sickle cell trait {Diggs, 1933 #21626}.


Edward Alfred Cockayne (GB) wrote the first book to be exclusively concerned with the genodermatoses and it contained numerous pedigrees, which had been culled from the literature. Cockayne’s stated purpose in writing the book was to draw the attention of dermatologists and geneticists to this potentially fruitful field of research {Cockayne, 1933 #22883}.


John Burdon Sanderson Haldane (GB-IN) suggested that the immune response of mice allowing them to reject tumors which arose in a different strain would be directed against normal cellular antigens unique to that strain rather than against tumor-specific antigens unique to the tumor {Haldane, 1933 #19087}.


Louis Barkhouse Flexner (US) proved that cerebro-spinal fluid is produced by a secretory process in the choroid plexuses {Flexner, 1933 #11113;Flexner, 1934 #11114}.


Cicely D. Williams (GB) described a nutritional disease among the children of the Gold Coast of Africa, which the natives called Kwashiorkor, meaning the red or brown boy. The symptoms of the disease are: (1) edema of the hands and feet, (2) profound wasting, (3) dry, scaly skin with patches devoid of pigmentation, (4) hair which is dry, sparse, and often a dull reddish, muddy color, (5) diarrhea and irritability, (6) fatty degeneration of the liver at necropsy. The disease was common to children between one and four years of age who typically had been weaned following a long period of breast-feeding. The diet of these children was mostly cereal, maize, with no milk {Williams, 1933 #1648;Williams, 1935 #1649}.

John Fleming Brock (ZA) and Marcel Autret (IT) found kwashiorkor in the whole tropical belt of Africa between Zanzibar and Dakar; it was also found in the Union of South Africa and in Egypt. The incidence of the disease in these areas varies greatly, which can be explained by diet. Where consumption of animal protein, such as meat, fish, and milk, occurs in reasonable quantities it is protective. There was some evidence that plant protein was protective {Brock, 1952 #23376}.


Joseph Godwin Greenfield (GB) described a fatal familial condition characterized by progressive loss of motor power with seizures, blindness, nystagmus, and mental deterioration in children {Greenfield, 1933a #22617;Greenfield, 1933b #22618}. Note: This collection of symptoms later became known as Greenfield’s disease.


James Robertson Dawson, Jr. (US) described subacute encephalitis {Dawson, 1933 #22619}.


John George Paxton Cleland (CA-US) gave an accurate description of the nerve pathways of uterine pain and clarified the sources of failure and success of regional obstetric block. He successfully applied paravertebral and low caudal blocks in obstetrics {Cleland, 1933 #22620}.


Joseph Clarence Hinsey (US) and Joseph E. Markee (US) proposed, “pathways from the hypothalamus must activate the posterior lobe of the hypophysis which in turn may exert an influence on the anterior lobe by hormonal transmission" {Hinsey, 1933 #6703}.

Joseph E. Markee (US), Charles Henry Sawyer (US), W. Henry Hollingshead (US), Geoffrey Wingfield Harris (GB), Barry Albert Cross (GB), John D. Green (GB), Bernard Dufy (FR), Luce Dufy-Barbe (FR), and Dominique Poulain (FR) later proved this experimentally {Cross, 1959 #6706;Dufy, 1974 #6707;Harris, 1948 #6705;Markee, 1946 #6704}.


Aldo Starker Leopold (US) and Allan Brooks (US) produced Game Management. This book established Leopold as the founding father of Wildlife Ecology. It includes discussions of game population dynamics, food chains, and habitat restoration, all in Leopold’s unmistakably lyrical tone {Leopold, 1933 #24203}.


Norman McOmish Dott (GB) performed the first planned intracranial operation for an arterial aneurysm. The target was the internal carotid artery {Dott, 1933 #13187}.


Geoffrey Douglas Hale Carpenter (GB), Edmund Brisco Ford (GB), Lincoln P. Brower (US), and John A. Endler (US) provided evidence from field observations and experiments that birds were often the agents of selection in insects, thus powerful agents in mimicry {Carpenter, 1933 #26689;Brower, 1988 #26690;Endler, 1986 #26691}. Note: the 1933 book is considered a masterpiece of evolutionary biology.


Theodore Christian Schneiria (US) discovered that Army ants operate on a 36-day cycle consisting of a 16-day nomadic pattern followed by a 20-day stationary phase. In 1934 he reported that ants follow a particular pattern when moving into new territory and that raids by these insects peak once during the morning and again in the afternoon. Sudden changes in weather also were found to give rise to sudden bursts of activity. In 1944 he showed that their raids were caused by the level of excitability of the ant colony and not by a scarcity of prey {Schneirla, 1933 #26165;Aronson, 1972 #26166}.


Andrei Vasilevich Martynov (RU) discovered the oldest undoubted fossils of Coleoptera (beetles) in Upper Permian deposits in North Russia {Martynov, 1933 #3142}.


Jesse D. Figgins (US) found large, heavy fluted stone points near Clovis, New Mexico. Mammoth bones in a deposit beneath a layer containing Folsom points and bison skeletons accompanied them. The robust points, now named Clovis, were recognized as even older than the Folsom points. Characteristic of both points is a flute, a flake struck off the base along the length of the point, presumably to facilitate hafting {Figgins, 1933 #12530}.

In 1964, C. Vance Haynes, Jr. (US) used radiocarbon dating to place the Clovis points at about 9.5-9 K BCE, and none before 10 K BCE {Haynes, 1964 #12531}.


Rene Neville (FR), in 1933, was the first to excavate fossil remains of Homo sapiens sapiens. The source was a cave site near Nazareth, Israel, on the southwest flank of Mount Qafzeh {Neuville, 1934 #17140}. Note: Subsequently more human fossil remains have been discovered at this site, all dated c. 100 K BP



"No single feature of man's past equals in importance his attempt to understand the forces of Nature and himself. It is a safe prediction that the historian of the future will be concerned increasingly with the chronicle of the intellectual acquisitions of man, for this deeper story includes not merely improvement in material comforts but mental enlargement which transcends every other feature of human evolution." Herbert McLean Evans {Evans, 1934 #13226}.


"Long ago I learned from my father to put old people to bed only for as short a time as was absolutely necessary, for they were like a foundered horse, if they got down it was difficult for them to get up, and their strength ebbed away very rapidly while in bed." Charles Horace Mayo {Mayo, 1934 #20015}.


Harold Clayton Urey (US) was awarded the Nobel Prize in Chemistry for his proof that hydrogen gas contains a few atoms in which the nuclear proton is accompanied by a neutron. This so called “heavy hydrogen” was given the name deuterium.


George Hoyt Whipple (US), George Richards Minot (US) and William Parry Murphy (US) were awarded the Nobel Prize in Physiology or Medicine for their discoveries concerning liver therapy in cases of anemia.


Irène Joliot-Curie (FR), Hans von Halban (AT), Peter Preiswerk (CH) and Frédéric Joliot (FR) discovered that radioactivity ca be induced artificially using alpha ray bombardment from radium. This methodology rapidly made available such valuable isotopes as 32P and 35S {Joliot-Curie, 1934 #14749;Joliot, 1936 #14748;Joliot-Curie, 1935 #14750}.

Marcus Laurence Oliphant (AU), Paul Harteck (GB), and Ernest Rutherford (NZ-GB) bombarded deuterium with the nuclei of deuterium atoms and produced tritium (hydrogen-3), the only radioactive form of hydrogen known. Tritium is much used in biochemical research {Oliphant, 1934 #14751}.


The first organic fungicide dithiocaramae was introduced. It proved valuable in foliar sprays for the control of a range of pathogenic fungi such as the scabs and rots of fruit and potato blight. ref


John Desmond Bernal (GB), and Dorothy Mary Crowfoot (GB) were the first to take an x-ray diffraction photograph of a crystalline protein. It was of the enzyme pepsin {Bernal, 1934 #723}.


Dorothy Mary Crowfoot (GB), in 1934, used x-rays crystallography to illuminate the structure of the protein, insulin {Crowfoot, 1935 #14670}.


Alfred Clarence Redfield (US) discovered that the atomic ratios between the chemical components of marine plankton, specifically nitrogen, phosphorus, and carbon are identical with their relative proportions in the open ocean. For every atom of phosphorus there are fifteen atoms of nitrogen and 105 atoms of carbon (carbonate and bicarbonate carbon are not considered organic) {Redfield, 1934 #10585}.


Karl Meyer (US) and John W. Palmer (US) isolated hyaluronic acid in pure form from vitreous humor, and determined its correct composition {Meyer, 1934 #15959}.


Robert Robinson (GB) hypothesized that cholesterol is formed by the cyclization of squalene, a polyisoprenoid hydrocarbon {Robinson, 1934 #9186}. See, E.A. Rudolph, 1925.


Alexander R. Kiesel (RU) and Andrei Nikolaevitch Belozersky (RU) were the first to isolate thymine and then DNA (thymonucleic acid) from higher plants {Belozersky, 1936 #19357;Kiesel, 1934 #19358}.

Andrei Nikolaevitch Belozersky (RU) and I.I. Dubrovskaya (RU) isolated DNA in the pure state for the first time {Belozersky, 1936 #19357}.


Leopold Stefan Ruzicka (HR-CH), Moses Wolf Goldberg (EE), Jules Meyer (), Heinrich Brüngger (CH), and E. Eichenberger () synthesized the sex hormone androsterone from epidihydrocholesterol. This was not only the first synthesis of a sex hormone, but also the first complete structural elucidation of such a compound, and the first exact proof of the relationship between a sex hormone and a sterol {Ruzicka, 1934 #9148}.


Hans Andersag (DE) discovered chloroquine, long the drug of choice in treating malaria {Andersag, 1934 #14752}.


Hans Lineweaver (US) and Dean Burk (US) determined that if the reciprocal of the velocity of an enzyme catalyzed reaction is plotted against the reciprocal of the substrate concentration a straight line is obtained, and that this cuts the two axes at the reciprocal of V (velocity) and the reciprocal of Km respectively. This is commonly referred to as a Lineweaver-Burk plot {Lineweaver, 1934 #5532}. Curiously enough this type of plot was first published in Kurt Guenter Stern’s German translation of John Burdon Sanderson Haldane’s book Enzymes (1932) where it was credited to Barnet Woolf (GB). Woolf likely failed to publish due to illness.


John R. Marrack (GB) proposed the lattice theory of antigen-antibody coupling with its fundamental requirement that antibody must have at least two antigen-combining sites {Marrack, 1934 #7282}.


Adolf Friedrich Johann Butenandt (DE) and Ulrich Westphal (DE), Willard Myron Allen (US), Oskar Wintersteiner (US), Max Hartmann (CH), Albert Wettstein (CH), Karl Heinrich Slotta (DE), Heinrich Ruschig (DE), and Erich Fels (DE) isolated and characterized progesterone in pure form {Allen, 1934 #15522;Allen, 1935 #19712;Butenandt, 1934b #14753;Hartmann, 1934 #15520;Slotta, 1934 #15521}.

Russell Earl Marker (US) and John Krueger (US), in 1939, developed a method for synthesizing progesterone in large quantities from fats and oils of plants. They developed the process, initially for Parke Davis and Co., for degrading sapogenins to C21 steroids {Marker, 1940 #19955}. Marker applied this process in 1941, to convert diosgenin from the wild Mexican yam into progesterone.


Vincent Brian Wigglesworth (GB) proved by experiment that during larval stages the corpus allatum (corpora allata) secretes a metamorphosis hormone, commonly called the juvenile hormone or neotenin, or youth substance. Under the influence of this hormone the larval characters are retained. When the larva is fully-grown, the corpus allatum (corpora allata) no longer secretes the hormone, the adult characters are developed, and metamorphosis occurs. His experimental animal was the large South American blood-sucking hemipteran Rhodnius prolixus.

In the 1940a paper he also argues that patterns derive from the diverse behavior of single cells — “the tiny fragment of cuticle laid down by a single cell may possess morphological characters controlled by the activity of that cell alone {Wigglesworth, 1934 #3783;Wigglesworth, 1940a #21139;Wigglesworth, 1940b #10642;Wigglesworth, 1971 #7240}. This physiological behavior has now been shown in almost all groups of insects.

Vincent Brian Wigglesworth (GB) found that the metamorphosis hormone is not genus specific and that egg formation in Rhodnius is dependent on the secretory function of the adult corpus allatum (corpora allata) {Wigglesworth, 1936 #10643}.

Jean-Jacques Bounhiol (FR) discovered that removal of the corpus allatum (corpora allata) results in precocious metamorphosis of the immature larval insect {Bounhiol, 1938 #4900}. By removing the corpus allatum (corpora allata) a conservative factor or status quo hormone is removed. Its function is to stabilize the larval tissues as larval tissues and the imaginal discs as imaginal discs. This status quo hormone became known as the juvenile hormone.

Vincent Brian Wigglesworth (GB) used implantation experiments to show that protocerebral neurosecretory cells are the source of the insect hormone that initiates the molting cycle {Wigglesworth, 1939 #16335}. This was the first experimental demonstration of an endocrine role for neural cells in any animal.


Edward Calvin Kendall (US), Harold L. Mason (US), Bernard F. McKenzie (US), Charles S. Myers (US), and Giles A. Koelsche (US) isolated and crystallized the adrenal cortical hormone, cortisone {Kendall, 1934 #3406}.

Harold L. Mason (US), Charles S. Myers (US), and Edward Calvin Kendall (US) were the first to separate cortisone as a new compound {Mason, 1936 #9124}.

Tadeus Reichstein (PL-CH), Oscar Paul Wintersteiner (US), Joseph J. Pfiffner (US), Harold L. Mason (US), Willard M. Hoehn (US), and Edward Calvin Kendall (US) determined the chemical structure of cortisone {Reichstein, 1936 #9127;Wintersteiner, 1936 #9126;Mason, 1938 #9128}.

Note: During the period 1934-1936, the Kendall group, the Reichstein group, and the Pfiffner-Swingle-Wintersteiner group were all purifying steroid hormones from the adrenal cortex. They were designating them with letters of the alphabet which created confusion because A in one group was not necessarily A in another group. The Kendall group isolated and crystallized five would be hormones. As the structure of these compounds was worked out, it became possible to name them. In the Kendall series Compound A is 11-dehydrocorticosterone, Compound B is corticosterone, Compound E is 17-hydroxy-11-dehydrocorticosterone (cortisone), and Compound F is 17-hydroxy-corticosterone (cortisol or hydrocortisone). In 1949, Edward Calvin Kendall (US), and Phillip Showalter Hench (US) gave the name cortisone to17-hydroxy-11-dehydrocorticosterone {Kendall, 1971 #11777}.

Marguerite Steiger (CH) and Tadeus Reichstein (PL-CH) synthesized desoxycorticosterone {Steiger, 1937 #14754}.

H. Reich () and Tadeus Reichstein (PL-CH) synthesized 11-dehydrocorticosterone {Reich, 1943 #14755}.

Lewis Hastings Sarett (US), working in the laboratory of Merck & Co., developed a method for synthesizing cortisone in quantities sufficient for clinical testing {Sarett, 1948 #9129}.

János Hugo Bruno “Hans” Selye (AT-HU-CA) coined the group names glucocorticoids for the 11-oxy steroid hormones of the adrenal cortex and mineralocorticoids for the hormone(s) of the adrenal cortex affecting the metabolism of electrolytes and water {Selye, 1943a #23547;Selye, 1941 #23548;Selye, 1942 #23549}.

János Hugo Bruno “Hans” Selye (AT-HU-CA), Christiane Dosne (CA), Lucy Bassett (CA), Joan Whitaker (CA), Eleanor L. Clark (CA), Octavia Sylvester (CA), Charles E. Hall (CA), and Charles Philippe Leblond (CA) explored the anti-inflammatory action of glucocorticoids and pro-inflammatory action of mineralocorticoids {Clark, 1943 #23554;Selye, 1949 #23555;Selye, 1940a #23556;Selye, 1940b #23557;Selye, 1944 #23558}.


Robert Russell Bensley (US) and Normand Louis Hoerr (US) used centrifugation to fractionate cell contents and isolate mitochondria {Bensley, 1934 #14756}.


Otto Fritz Meyerhof (DE-US) and Karl Lohmann (DE) isolated the enzyme from muscle extracts that cleaves fructose-1, 6-diphosphate into two triose phosphates. They initially called it zymohexase but later Meyerhof, Lohmann, and Philipp Schuster (DE) changed it to fructose-biphosphate aldolase. They also recount their discovery of triose-phosphate isomerase {Meyerhof, 1934 #9262;Meyerhof, 1935d #19615;Meyerhof, 1936a #14480;Meyerhof, 1936b #23187}.

Paul K. Stumpf (US) purified and characterized plant aldolase {Stumpf, 1950 #18929}.


Karl Lohmann (DE) and Otto Fritz Meyerhof (DE-US) showed that the formation of pyruvic acid involves the enzyme catalyzed migration of the phosphoryl group from the 3-position to the 2-position of glyceric acid, followed by the dehydration of 2-phosphoglyceric acid to 2-phosphoenolpyruvic acid (PEPA) by the enzyme enolase (phosphopyruvate hydratase). Enolase was observed to be strongly inhibited by fluoride, thus explaining the effect of fluoride on alcoholic fermentation and glycolysis {Lohmann, 1934b #15537}.

Otto Heinrich Warburg (DE) and Walter Christian (DE) isolated and crystallized the enzyme enolase (phosphopyruvate hydratase) {Warburg, 1941 #9400}.


Wilhelm Kiessling (DE) found that a ketotriose identical to synthetic dihydroxyacetone phosphate is a reaction product when hexosediphosphate is split into two trioses {Kiessling, 1934 #9263}.


Harland Goff Wood (US) and Chester Hamlin Werkman (US) were the first to isolate and identify pyruvic acid as an intermediate in the propionic acid fermentation {Wood, 1934 #11203}.


Karl Lohmann (DE) used dialyzed muscle extracts and found that the hydrolysis of creatine phosphate to creatine and phosphate is promoted by the addition of ATP, which is cleaved to adenylic acid (adenosine monophosphate) and two equivalents of inorganic phosphate. He concluded that the ATP acts as a coenzyme in the hydrolysis of creatine phosphate. He identified ATP as the coferment (coenzyme) of lactic acid formation in muscle and determined that it required the presence of the magnesium ion {Lohmann, 1934a #2562}.


Gottfried Samuel Fraenkel (DE-US) discovered the blood-borne factor we now know to be the insect molting hormone, ecdysone (ecdysterone) {Fraenkel, 1934 #26679;Fraenkel, 1935 #26680}.


Peo C. Koller (US) and Cyril Dean Darlington (GB) showed that sex-chromosomes of the Norway rat (Rattus norvegicus) each consist of a pairing segment and a differential non-pairing segment. The pairing segment includes the spindle attachment and chiasmata may be formed on one or both sides of it so that the first division is either reductional or equational for the differential segments. The differential segments will have complete sex-linkage, the pairing segments will have partial sex-linkage diminishing in proportion to the crossing-over distance from the differential segments. The shape, movements, and staining capacity of the sex-chromosomes in the rat and elsewhere agree in suggesting that they have a lower surface charge than the autosomes, and this is held to be responsible for the special mechanism of X-chromosome segregation in organisms lacking theY {Koller, 1934 #24501}.

Peo C. Koller (GB) confirmed the presence of an X-Y pairing segment in human meiosis and speculated about the possibility of partial sex linkage —proof of which had to wait for another 40 years {Koller, 1938 #24500}.


Isaac McKinney Lewis (US) measured the frequency of lactose-negative to lactose-positive mutations in Escherichia coli mutabile and found it to be on the order of one mutation/one hundred thousand cells/cell generation {Lewis, 1934 #18930}.


Phineas W. Whiting (US), working with Habrobracon, was the first to study what would later be called conditional lethal mutants {Whiting, 1934 #8606}.

Norman Harold Horowitz (US) and Urs Leupold (CH) were the first to isolate temperature-sensitive mutants of bacteria. Such mutants are only able to grow at temperatures lower than that at which the wild-type can grow {Horowitz, 1951 #8605}.

Ernst Hadorn (CH) coined the term conditional lethal mutant to describe mutants of Drosophila which were either lethal, or allowed relatively normal development to occur, depending upon the growth conditions imposed by the experimenter {Hadorn, 1951 #8607}.


Henry Hallett Dale (GB) and Wilhelm Siegmund Feldberg (DE-GB) reasoned that the substance released by stimulated nerves, which provokes the contraction of eserinized (eserine blocks acetylcholinesterase) leech muscle, must be acetylcholine {Dale, 1934 #5284}.


Ragnar Arthur Granit (FI-SE) and Per-Olof Therman (SE) established that details of the visual image are elaborated by the interplay of excitation and inhibition in the nervous center of the retina, i.e., that light can both excite and inhibit and that the two opposite processes are antagonistic. They did this by recording the mass discharge in the optic nerve together with the electroretinograph {Granit, 1934 #9236;Granit, 1935 #9237}.


Claud D. Johnson (US) and Ernest William Goodpasture (US) filtered the mumps agent through Berkefeld V and N filters then with the filtrate produced the disease in monkeys. This was proof that mumps (epidemic parotitis) is caused by a virus {Johnson, 1934 #2361}.

Karl Habel (US) succeeded in growing the mumps virus in the chick embryo. He also demonstrated a good correlation between the skin reactions to virus grown in the chick allantoic sac and in the monkey parotid gland. Because of the greater availability of egg-grown virus, the wider use of this reaction then became possible {Habel, 1945 #11645}.

Jeanette H. Levens (US) and John Franklin Enders (US) developed a hemagglutination test to titer the mumps virus {Levens, 1945 #11917}. They also showed that influenza A virus could be assayed by hemagglutination.

Thomas Huckle Weller (US), and John Franklin Enders (US) grew the mumps virus in cell culture consisting of fragments of chick amniotic membrane nourished with a balanced salt solution and ox serum ultrafiltrate. They succeeded, where others had failed, by incorporating the recently available penicillin into their cultures {Weller, 1948 #9156}.

Gertrude Henle (DE) and Friedrich Deinhardt (DE) propagated and performed primary isolation of mumps virus in monkey kidney tissue culture {Henle, 1955 #14915}.

Robert E. Weibel (US), Victor M. Villarejos (US), Gloria Hernandez (ES), Joseph Stokes, Jr. (US), Eugene B. Buynak (US), and Maurice Ralph Hilleman (US) developed a combined live measles-mumps virus vaccine {Weibel, 1973 #15286}.

William J. McAleer (US), Eugene B. Buynak (US), Robert E. Weibel (US), Victor M. Villarejos (US), Edgar M. Scattergood (US), Hermann E. Wasmuth (US), Arlene A. McLean (US), and Maurice Ralph Hilleman (US) developed measles, mumps and rubella virus vaccines prepared from virus produced by the unit process {McAleer, 1975 #15287}.


Albert Bruce Sabin (US) and Arthur M. Wright (US) isolated the Herpes simian virus (later named B virus) from the brain of a lab worker who died after being bitten by a Macacus rhesus monkey {Sabin, 1934 #22612}.

Joseph Louis Melnick (US) and Dushyant D. Banker (US) isolated B virus (herpes group) from the central nervous system of a rhesus monkey {Melnick, 1954 #22613}.


Charles Armstrong (US), and Ralph Dougall Lillie (US) appear to be among the first to have described an infection caused by the virus of lymphocytic choriomeningitis (acute aseptic meningitis, idiopathic meningitis). It appeared as a contaminant in monkeys inoculated with the virus of St. Louis Encephalitis {Armstrong, 1934 #12392}.

T.F. McNair Scott (US) and Thomas Milton Rivers (US) isolated and identified the lymphocytic choriomeningitis virus from human cases {Rivers, 1935 #2364;Rivers, 1936 #16187;Scott, 1936 #2363}.


Ivan Claude Jagger (US) and Norman Chandler (US) were the first to describe the Big Vein Disease of lettuce and report that it is soil-borne and infectious {Jagger, 1934 #2041}.


Carl Clarence Lindegren (US) found that in Neurospora all asci in a single perithecium arise from a single pair of nuclei associated at the initiation of the perithecium {Lindegren, 1934 #1688}.


Wendell H. Tisdale (US) and Ira Williams (US) working for the Du Pont Chemical Company discovered the dialkyldithiocarbamates as fungicides {Tisdale, 1934 #15524}. This group includes thiram, ferbam, maneb, zineb and mancozeb which are all surface acting agents.


Ladislaus Laszlo Marton (US), Stuart Mudd (US), and David Lackman (US) were among the first to publish electron photomicrographs of biological specimens in the United States {Marton, 1934a #8814;Marton, 1934b #14758;Mudd, 1941 #14757}. In 1937 Marton published the first electron micrograph of bacteria.


Arthur Felix (PL-GB) and Margaret R. Pitt (GB) discovered the heat labile Vi somatic antigen of the Salmonella {Felix, 1934 #110}.


Alice Catherine Evans (US), using the Clark phage which she renamed B563, was the first person to utilize a phage to classify bacterial strains, thus founding the analytical field of phage typing {Evans, 1934 #8815}.

Roy T. Fisk (US) developed the method of typing staphylococci by using bacteriophages {Fisk, 1942 #15913}.


Per Fredrik Thorkelsson Scholander (SE-NO-US) revised the lichen family, the Umbilicariaceae {Scholander, 1934 #10812}.


Sanford B. Hooker (US) and Edna M. Follensby (US) discovered that the erythrogenic toxin of scarlet fever is two closely related toxins, which they called erythrogenic toxins A and B {Hooker, 1934 #111}.


Malcolm H. Merrill (US), C. William Lacaillade, Jr. (US), and Carl Ten Broeck (US) showed that the virus of Eastern equine encephalitis is vectored by mosquitoes {Merrill, 1934 #25952}.


Florence Barbara Seibert (US) prepared purified protein derivative (PPD) from tuberculin. This enabled the first reliable tuberculin test {Seibert, 1934 #12324}.

Florence Barbara Seibert (US) and John T. Glenn (US) prepared a large batch of PPD that has served as the standard reference material (PPD-S) in the United States {Seibert, 1941 #12322}.


Louis Alphonse Julianelle (US) and Charlotte W. Wieghard (US) introduced the first classification of the staphylococci based on differences in antigenic structure. They recognized two serological types of staphylococci based of specific carbohydrates obtained by chemical fractionation of the organisms. Type A strains were found to be pathogenic and capable of fermenting mannitol, while type B was comprised of strains which failed to ferment mannitol and had little, if any, pathogenicity {Julianelle, 1934 #14628}.


Chester W. Emmons (US) modernized the taxonomic schemes of Sabouraud and others and established the current classification of the dermatophytes based on spore morphology and accessory organs {Emmons, 1934 #12373}.


Eric T.B. Francis (GB) authored The Anatomy of the Salamander (Salamander musculosa), an example of outstanding descriptive zoology {Francis, 1934 #5163}.


William T. Heron (US), William M. Hales (US), and Dwight Joyce Ingle (US) reported that repetitive contraction of skeletal muscle requires the activity of a substance(s), which can be extracted from the adrenal cortex {Heron, 1934 #10715}. This knowledge was developed into a bioassy for adrenal hormones that facilitated the purification of cortisone.


Balduin Lucké (US) suggested a viral etiology for a renal adenocarcinoma he observed in the Northern Leopard Frog, Rana pipiens. This is the first postulation of a virus infection of an amphibian {Lucké, 1934 #8651;Lucké, 1938 #16830}.


Wallace Osgood Fenn (US), Doris M. Cobb (US), Albert H. Hegnauer (US), Burton Sanford Marsh (US), Jeanne F. Manery (US), Walter R. Bloor (US), Thomas R. Noonan (US), Lorraine F. Haege (US), Lorin J. Mullins (US), and Robert B. Dean (US) made the first determinations of potassium, sodium, magnesium and calcium in nerve. They showed that intracellular potassium is mobile, and that muscle potassium shifts in response to various environmental factors. They found that during muscle contraction potassium is lost from muscle in exchange for sodium, and that the process is reversed in recovery. This was the first time that sodium was shown to penetrate muscle. This work laid the foundation for the Hodgkin-Huxley hypotheses concerning initiation and propagation of nerve and muscle impulses and the magnitude and polarity of electrical potential differences across cell membranes. Fenn said, “The explanation for a loss of potassium from a muscle during activity is a matter of fundamental theoretical importance. In terms of the theory which I have been using as a guide, it is interpreted as an increase in the permeability of the muscle membrane of sufficient extent to permit sodium which enters then displaces one molecule of potassium” {Fenn, 1936b #10874}. They showed that potassium escapes from muscle during contraction in situ and that a large part of this potassium appears in the liver. Potassium uptake was linked with carbohydrate metabolism, particularly with glycogen deposition, and tends to follow the Cori cycle. Radioactive potassium was ingested as part of the first study of the kinetics of potassium metabolism and the demonstration that it is taken up by erythrocytes. Nearly all muscle potassium was found to be exchangeable, supporting the notion that it is maintained by an active energetic process {Fenn, 1934a #10868;Fenn, 1934b #10869;Hegnauer, 1934 #10870;Fenn, 1934c #10871;Fenn, 1935 #10872;Fenn, 1936a #10873;Fenn, 1936b #10874;Fenn, 1936c #10875;Fenn, 1937a #10876;Fenn, 1937b #10877;Fenn, 1938a #10878;Fenn, 1938b #10879;Fenn, 1938c #10880;Fenn, 1939a #10881;Fenn, 1939b #10882;Fenn, 1940 #10883;Noonan, 1941a #10884;Noonan, 1941b #10885;Mullins, 1941 #10886;Dean, 1941 #10887;Fenn, 1942 #10888}.


Philip Duryeé McMaster (US), Stephen S. Hudack (US), and John G. Kidd (US) demonstrated that lymph nodes, draining skin sites injected with bacteria or viruses, formed antibodies against these agents in very high concentrations {McMaster, 1934 #10902;Hudack, 1934 #10903;McMaster, 1935 #10904;McMaster, 1936 #10905;McMaster, 1937 #10906}.


Frederick W. Madison (US) and Theodore L. Squier (US) defined the etiology of primary granulocytopenia (agranulocytic angina). They suggested that the primary granulocytopenia following the use of such drugs amidopyrine with a barbiturate might be the result of an allergic or anaphylactoid drug reaction {Madison, 1934 #19314}.


John Silas Lundy (US) brought to a climax a long series of trials by many workers when he used the intravenous introduction of Pentothal (thiopental sodium, a barbiturate) to put a patient peacefully to sleep on June 18, 1934. Pentothal rapidly became the standard induction agent, being much more pleasant than inhaling the pungent ether. It was not until the 1990s that propofol, a more rapidly metabolized agent with fewer side effects, finally replaced Pentothal. Lundy (US) is best known for introducing intravenous anesthesia into clinical practice {Lundy, 1935 #14783}.


Ralph Milton Waters (US) and Erwin R. Schmidt (US) published their paper on the physiologic and pharmacologic effects of cyclopropane on the human body {Waters, 1934 #12304}.


J. Roswell Gallagher (US) described an outbreak of bronchopneumonia in a group of 16 boys living at a preparatory school (a closed community). He especially stressed that these children did not have pneumococcal pneumonia but something different--something "atypical" {Gallagher, 1934 #11554}.

Robert A. Reimann (US) described a group of eight patients with chest infection but atypical clinical presentations and no chest pain. He coined the term "atypical pneumonia" because the manifestations in these patients differed greatly from those in patients presenting with acute pneumococcal pneumonia. Reimann's initial impression was that this illness was caused by a filterable agent, most likely a virus, but to this date the cause of his patients' illnesses remains unknown {Reimann, 1938 #11553}.


Clive M. McCay (US), Mary F. Crowell (US), and Leonard Amby Maynard (US) studied the effects of food restriction on the life span of rats. They concluded that if an animal ate what it should and little more its life span was increased {McCay, 1934 #12404;McCay, 1935 #12405}.


Ernst Klenk (DE) identified sphingomyelin as the stored phospholipid in cells from patients with Niemann-Pick disease {Klenk, 1934 #13685}.


Ivar Asbjørn Følling (NO) described phenylketonuria (PKU) and called attention to its association with serious mental deficiency. He developed a test for demonstrating phenylpyruvic acid found in the urine of a person with PKU {Følling, 1934 #13250}.

George A. Jarvis (US), Richard J. Block (US), Diana Bolling (US), Edna Kanze (US), Merrill Webb (US), Ernest Borek (US), Arthur Brecher (US), and Heinrich Waelsch (US) demonstrated that the biochemical defect lies in the failure of the affected individuals to oxidize phenylalanine to tyrosine in the normal fashion {Jervis, 1947 #23400;Jervis, 1950 #23401;Borek, 1950 #23402;Jervis, 1952 #23403;Jervis, 1953 #23404;Jervis, 1938 #23405;Jervis, 1940 #23406;Block, 1940 #23407}.

Marvin D. Armstrong (US) and Frank H. Tyler (US) performed experiments, the results of which, suggested that patients suffering from phenylketonuria would benefit from a restricted dietary intake of phenylalanine {Armstrong, 1955 #23399}.


Fuller Albright (US), Esther Bloomberg (US), Benjamin Castleman (US), Edward D. Churchill (US), Walter Bauer (US), and Joseph C. Aub (US) gave the initial clinical description of hyperparathyroidism {Albright, 1934a #10922;Albright, 1934b #10923}.


William Jason Mixter (US) and Joseph S. Barr (US) discovered herniated intervertebral disks as a pathological condition {Mixter, 1934 #9579}.


Lillian Lauricella (US), on April 17, 1934, gave birth to twin daughters, conceived by artificial insemination using donor sperm (AID) {, 1934 #18932}.


Walter Edward Dandy (US) outlined his theory of vascular compression as a cause of trigeminal neuralgia and pointed to the main problem with that theory; namely, that vascular contact occasionally occurs without the production of pain and may be absent when neuralgia is present {Dandy, 1934 #12213}.

Peter J. Hamlyn (GB) and Thomas T. King (GB) confirmed that vascular compression of the fifth cranial nerve is an anatomical abnormality specific to trigeminal neuralgia {Hamlyn, 1992 #12214}.


Alexandra Adler (AT-US) suggested that the thalamic arcuate nucleus of the brain is associated with taste {Adler, 1934 #17706}.

Harry Dickson Patton (US), Theodore Cedric Ruch (US), and A. Earl Walker (US) subjected monkeys to lesions of the thalamic arcuate nucleus and confirmed Adler’s suggestion {Patton, 1944 #17707}.

Harry Dickson Patton (US) found evidence that the most important part of the taste area laid buried in the operculum, just below the facial regions {Patton, 1950 #17708}.


Michael Ellis DeBakey (US), four years after entering medical school, designed the first roller pump, which remains the basis for all cardiac by-pass surgery performed today {DeBakey, 1934 #11290}.


Ralph A. Colp (US) was the first to describe a granulomatous inflammation of the terminal ileum and cecum called ileo-colitis {Colp, 1934 #15938}.


Arthur Evans (GB) was the first to successfully operate on the thoracic region of the esophagus for cancer. He reported a 23-year cure of cancer of the cervical esophagus by radical excision of the cervical esophagus and larynx {Evans, 1934 #19696}.


Philippe l'Héritier (FR) and Georges Teissier (FR) devised the population cage method for the experimental study of natural selection. They showed, among other things, that unfavorable mutations (such as bar) could be maintained in a population of Drosophila in a stable balanced polymorphism despite the deleterious effects of the mutation on the flies that carried it {l'Héritier, 1934 #14616;l'Héritier, 1937a #14617;l'Héritier, 1937b #14618}.


Nikolai Wladimirovich Timoféeff-Ressovsky (RU) experimentally measured the viability of strains of Drosophila funebris of different geographical origin {Timoféeff-Ressovsky, 1934 #14744}.


Francis Bertody Sumner (US) experimentally showed the selective value of protective coloration in fishes {Sumner, 1934 #15535}.


William Beebe (US) and Otis Barton (US) developed the deep diving bathysphere. This round steel submersible was used on August 5, 1934 for a drop to 3,028 feet into the ocean off Nonsuch Island {Beebe, 1934 #14619;Crane, 1934 #14620}. Beebe is commemorated by Protopelagonemertes beebei Coe, 1936, Metapenaeopsis beebei Burkenroad, and Aeginura beebei Bigelow, 1940.

Auguste Piccard (CH-BE) developed a bathyscaphe, i.e., a more maneuverable submarine balloon, but looking somewhat like a conventional submarine. In 1954 his second bathyscaph the Trieste descended 10,330 feet into the Mediterranean and in 1960 his third vessel Trieste II was lowered to 35,800 feet (10,900 m) when it touched bottom in the Marianas Trench.


George Edward Lewis (US) found the first Ramapithecus (Ramapithecus brevirostris), the earliest known hominid fossil, in deposits in the Siwalik Hills of Northern India {Lewis, 1934 #14164}. It was alive during the Miocene epoch, c. 24 M.



“The air of caricature never fails to show itself in the products of reason applied relentlessly and without correction. The observation of clinical facts would seem to be a pursuit of the physician as harmless as it is indispensable. [But] it seemed irresistibly rational to certain minds that diseases should be as fully classifiable as are beetles and butterflies. This doctrine ... bore perhaps its richest fruit in the hands of Boissier de Sauvauges. In his Nosologia Methodica published in 1768 ... this Linnaeus of the bedside grouped diseases into ten classes, 295 genera, and 2400 species.” Wilfred Batten Lewis Trotter {Trotter, 1935 #22824}.


“No one of the laity can realize how much time, pains, mistakes, imagination, mental distress, and money have gone into the making of an experienced surgeon.” Robert Tuttle Morris {Morris, 1935 #23284}.


"No one can be a one-hundred-per-cent doctor until he has himself had some serious illness or surgical operation." Robert Tuttle Morris {Morris, 1935 #23284}.


Hans Spemann (DE) was awarded the Nobel Prize in Physiology or Medicine for his discovery of the organizer effect in embryonic development.


Max Knoll (DE) demonstrated the feasibility of the scanning electron microscope; three years later a prototype was built by Manfred von Ardenne (DE) {Knoll, 1935 #14630;von Ardenne, 1938 #14631}.


Alexis Carrel (FR-US) and Charles A. Lindbergh (US) described an all-glass system for the perfusion of whole organs {Carrel, 1935 #15617}.


Arthur Lindo Patterson (NZ-US) developed an analytical method for determining interatomic spacings from x-ray data {Patterson, 1935 #14632}.


Rudolf Schoenheimer (DE-US) and David Rittenberg (US) introduced the use of deuterium as an isotopic tracer into biochemical research in animals {Schoenheimer, 1935 #2611}. This methodology marked a milestone in biochemistry because for the first time an isotope was systematically introduced into an organic compound so that a defined reaction or pathway could be studied. See, Georg Charles de Hevesy, 1923.


Wolfgang Schott (DE), inspecting findings of the German Meteor oceanographic expedition of 1925-27, realized that the species whose shells were found in the muck of the seabed depended sensitively on the temperature of the water where the creatures had lived. The mix of foraminifera species could serve as a thermometer of past climates {Schott, 1935 #14906}.

Harold Clayton Urey (US), Samuel Epstein (US), Heinz A. Lowenstam (US), and Charles R. McKinney (US) were able to prepare a history of changing ocean temperatures over long geologic periods. This work was based on the knowledge that heavy isotopes of oxygen react more slowly than normal atoms. Thus, the proportion of oxygen isotopes in a seashell depends upon the temperature of the ocean at the time the shell was formed {Urey, 1947 #14907;Urey, 1950 #14908;Urey, 1951 #14909}.


David Rockwell Goddard (US) and Leonor Michaelis (DE- US) found that reducing agents such as thioglycolic acid could break the disulfide linkages in keratin (hair, wool, and feathers). They did not patent this discovery {Goddard, 1935 #10583}. Thioglycolic acid and other thiol compounds became the basic ingredients of the permanent wave solutions used in the cosmetic industry.


Harry Bender (US) discovered that technical benzene hexachloride (BHC) is a potent insecticide. He added benzene to chlorine in a Dewar flask in the open air and noticed that part of the product which spilled on the ground “attracted and killed flies and bees” {Bender, 1935 #3646}. See Michael Faraday, 1825 and Van der Linden, 1912.


Eric John Underwood (AU), John Francis Filmer (AU), and Hedley Ralph Marston (AU) showed that cobalt is a necessary element in the diet of animals {Underwood, 1935 #3108;Marston, 1935 #3109}.


Ernst Klenk (DE) characterized a new type of acidic glycolipid—naming it substance X—from the brains of patients suffering from amaurotic familial idiocy {Klenk, 1935 #13686}.

Ernst Klenk (DE) coined the term ganglioside to name substance X that he characterized in 1935 {Klenk, 1942 #13693}.


Paul Rothemund (US) synthesized the simple chlorophyll-like substance protochlorophyll {Rothemund, 1935 #8429}.


Robert H. Sifferd (US) and Vincent du Vigneaud (US) synthesized carnosine, a naturally occurring small peptide {Sifferd, 1935 #11942}.


Charles Robert Harington (GB) and Thomas Hobson Mead (GB) were the first to synthesize glutathione, a small naturally occurring polypeptide {Harington, 1935b #11937}.


Richard H. McCoy (US), Curtis E. Meyer (US), and William Cumming Rose (US) isolated the amino acid threonine from hydrolysates of fibrin and demonstrated that it is one of the essential amino acids in rats {McCoy, 1935 #2757;Meyer, 1936 #2758}.

Herbert E. Carter (US) synthesized all isomers of threonine and found that the L-isomer is the essential form in rats {Carter, 1979 #10983}.


Charles Robert Harington (GB) reported that enlargement of the thyroid gland in hyperthyroidism was found to be associated with heart dysfunction, exophthalmos, and increased metabolic rate {Harington, 1935a #3096}.

Ya-Pin Lee (US), Akira E. Takemori (US), Henry Arnold Lardy (US), Ching-Yuan Su (US), Nancy Kneer (US), and Susan Wielgus (US) found that thyroid hormone and also dehydroepiandrosterone induced the synthesis of mitochondrial glycerol-3-phosphate dehydrogenase to as much as 20 times the normal concentration and formed part of the thermogenic system {Lardy, 1989 #19414;Lardy, 1999 #19415;Lee, 1959 #19412;Lee, 1965 #19413}.


Harold King (GB) purified and determined the structure of d-tubocurarine, one of the many alkaloids present in curare preparations {King, 1935a #16823;King, 1935b #16824}.

A.J. Everett (GB), L.A. Lowe (GB), and S. Wilkinson (GB) made one minor correction in its structure {Everett, 1970 #5276}.


Kurt Guenter Stern (GB-US) using optical methods (a spectrophotometer) made the first direct observation of an enzyme-substrate complex {Stern, 1935 #5551}.


Béla Tankó (HU) and Robert Robison (GB) announced the discovery of phosphohexose kinase {Tankó, 1935 #18994}.


Karl Zeile (DE) and Axel Hugo Theodor Theorell (SE) performed work, which led to the complete purification of cytochrome c and its characterization as a protein unit of molecular weight 13,000 with a porphyrin unit covalently linked to the protein by way of two cysteine residues {Zeile, 1935 #2506;Theorell, 1947 #2507}.


Otto Heinrich Warburg (DE), Walter Christian (DE), Alfred Griese (DE), Frank Dickens (GB), Gertrude E. Glock (GB), Bernard Leonard Horecker (US), Pauline Z. Smyrniotis (US), Jarvis Edwin Seegmiller (US), Paul A. Marks (US), Howard H. Hiatt (US), Hans Klenow (US), Efraim Racker (PL-AT-US), and Dan Couri (US) extensively studied and purified all of the enzymes found in the type of glycolysis called the phosphogluconate pathway, also known as the pentose phosphate pathway or hexose monophosphate shunt. This is a multifunctional pathway specialized to carry out four main functions: (1) generate reducing power in the form of NADPH, needed for the synthesis of fatty acids and steroids from acetyl-CoA, (2) to convert hexoses to pentoses, particularly D-ribose 5-phosphate, required for the synthesis of nucleic acids, (3) the oxidative degradation of pentoses by converting them into hexoses, which can then enter the glycolytic sequence, and (4) it is modified so as to participate in the formation of glucose from CO2 in the dark reactions of photosynthesis {Dickens, 1936 #16184;Dickens, 1938 #9270;Dickens, 1951 #9283;Horecker, 1951a #9271;Horecker, 1951b #9282;Horecker, 1953a #15971;Horecker, 1953b #12402;Horecker, 1955 #15973;Couri, 1959 #11061;Warburg, 1935c #2551;Warburg, 1937 #9281}. Horecker (US) coined the phrase hexose monophosphate shunt in his 1951b paper mentioned above.

Paul A. Srere (US), Jack R. Cooper (US), Vida Klybas (US) and Efraim Racker (PL-AT-US), discovered xylulose-5-phosphate, a new intermediate in the pentose phosphate cycle {Srere, 1955a #16019}.

Paul A. Srere (US), Hans Leo Kornberg (GB-US), and Efraim Racker (PL-AT-US) demonstrated that transaldolase and transketolase, highly purified, were able to affect the conversion of pentose phosphate to hexose phosphate {Srere, 1955b #19393}.

Hans Leo Kornberg (GB-US) and Efraim Racker (PL-AT-US) found that erythrose 4-phosphate is an intermediate in that process {Kornberg, 1955 #16020}.

Harland Goff Wood (US), Joseph Katz (US), and Bernard R. Landau (US) used carbon-14 to estimate the proportion of carbohydrate metabolized in the pentose pathway and glycolysis. These studies helped determine the stoichiometry of the pentose pathway {Wood, 1963 #11035}.


Percy W. Zimmerman (US) and Frank Wilcoxon (US) discovered several synthetic substances with hormone activity in plants {Zimmerman, 1935 #14633}.


Hermann James Almquist (US) and E.L. Robert Stockstad (US) demonstrated that fecal microorganisms are capable of synthesizing vitamin K. They purified the vitamin under a high vacuum {Almquist, 1935 #3080;Almquist, 1936b #3081;Almquist, 1936c #3082;Almquist, 1936a #3083;Almquist, 1936d #3084}.


Phoebus Aaron Theodor Levene (RU-US) and Robert Stuart Tipson (US) determined that “…in desoxy-ribose nucleic acid the positions of the phosphoric acid radicals are carbon atoms (3) and (5) of the desoxyribose” {Levene, 1935b #14634}.


Otto Fritz Meyerhof (DE-US) and Wilhelm Kiessling (DE) found that muscle extract contains an isomerase, which catalyzes the conversion of synthetic D-3-phosphoglyceraldehyde to dihydroxyacetone phosphate {Meyerhof, 1935b #9264}.


Otto Fritz Meyerhof (DE-US) and Wilhelm Kiessling (DE) showed that in glycolysis it is the oxidation of the aldehyde to the acid that balances the reduction of acetylaldehyde to ethanol (in alcoholic fermentation) or of pyruvic acid to lactic acid (in glycolysis), and that phosphoglycerol is not a necessary participant in the dismutation as Embden had proposed. They also demonstrated that in iodoacetate poisoned muscle extracts, the phosphoryl group of 2-phosphoenolpyruvic acid is transferred to glucose by way of ATP to yield hexose phosphates and pyruvic acid {Meyerhof, 1935a #2548}.


Harland Goff Wood (US), Chester Hamlin Werkman (US), Allen Hemingway (US) and Alfred Otto Carl Nier (US) demonstrated that many heterotrophic forms of life assimilate carbon dioxide. They proposed that carbon dioxide and pyruvate combine to form oxaloacetate, which subsequently is reduced to succinate—the Wood-Werkman reaction {Werkman, 1942 #2620;Wood, 1935 #7173;Wood, 1936 #8820;Wood, 1938 #14590;Wood, 1941 #11197;Wood, 1942 #11198}.

Arthur Kaskel Solomon (US), Birgit Vennesland (DE-US), Friedrich W. Klemperer (US), John Machlin Buchanan (US), and A. Baird Hastings (US) determined that higher organisms utilize carbon dioxide as a substrate of reactions {Solomon, 1941 #15961}.

Hutton D. Slade (US), Harland Goff Wood (US), Alfred Otto Carl Nier (US), Allen Hemingway (US), and Chester Hamlin Werkman (US) reported that fixation of carbon dioxide by C3 and C1 addition is apparently a very general reaction among the heterotrophic bacteria {Slade, 1942 #11199}. In 1942, it was dogma that carbon dioxide is an inert product of the metabolism of all living forms except the specially adapted chemosynthetic and photosynthetic autotrophs.

Henry Arnold Lardy (US) and Julius Adler (US) found that propionate could be metabolized by carbon dioxide addition to ultimately yield succinate {Lardy, 1956c #19409}.


Hans Adolf Krebs (DE-GB) found that the kidney contains separate oxidative deaminases for D- and L-amino acids and that both kidney and brain tissue will convert ammonium glutamate to glutamine and hydrolyze it back as well {Krebs, 1935a #2605;Krebs, 1935b #2606}.


Henry Borsook (US) and Geoffrey Keighley (US) concluded from nutritional studies that there is a continuing metabolism of protein, and that tissue proteins are constantly being synthesized from amino acids {Borsook, 1935 #2616}.


Sven Otto Hörstadius showed the existence of a double gradient of "animalization" and "vegetalization" in the echinoderm egg {Hörstadius, 1935 #25208}.


Charles Herbert Best (US-CA) and M. Elinor Huntsman (CA) reported that choline is a lipotropic agent that prevents deposition of fat in the liver {Best, 1935 #14769}.


Kenneth Bryan Raper (US) identified the slime mold, Dictyostelium discoideum, then with Theo M. Konijn (US) laid the groundwork for the later use of this organism as a model system for the study of intercellular communication {Raper, 1935 #9940;Konijn, 1961 #9943}.


Jakub (Jacob) Karol Parnas (PL) and Tadeusz (Tadeush) Baranowski (PL) found that in muscle extracts glycogen and inorganic phosphate can react to form hexose monophosphates if the oxidation-reduction process is blocked using iodoacetic acid and no ATP is being generated {Parnas, 1935 #14638}. This was of importance because it established beyond doubt the participation of inorganic phosphate in the splitting of glycogen and the discovery of phosphorylase activity in muscles.

Carl Ferdinand Cori (CZ-US) and Gerty Theresa Cori, née Radnitz (CZ -US) discovered that α-glucose-1-phosphate (Cori ester) is formed naturally in muscle. This was evidence for the presence of glycogen phosphorylase (Cori phosphorylase) and the realization that the formation of the 1-ester may be the result of a phosphorolytic split of glycogen{Cori, 1936 #2573}.

Carl Ferdinand Cori (CZ-US), Sidney P. Colowick (US), and Gerty Theresa Cori, née Radnitz (CZ -US) gave a full description of α-glucose-1-phosphate as a highly important ester. They found that by modifying the muscle extraction procedure another enzyme, phosphoglucomutase, catalyzed the conversion of α-glucose-1-phosphate to the very stable phosphoric ester that they identified as glucose-6-phosphate {Cori, 1937 #23148;Cori, 1938a #9259;Cori, 1938c #23155}.

Victor Assad Najjar (LB-US) obtained phosphoglucomutase in crystalline state {Najjar, 1948 #23156}.

Luis Federico Leloir (AR), Raul E. Trucco (AR), Carlos Eugenio Cardini (AR), Alejandro Paladini (AR), and Ranwel Caputto (AR) discovered that glucose-1,6-diphosphate acts as a coenzyme of phosphoglucomutase allowing a reversible conversion between α-glucose-1-phosphate and glucose-6-phosphate {Caputto, 1948 #23192;Cardini, 1949 #23162;Leloir, 1948 #23163}.

Carl Ferdinand Cori (CZ-US), Gerty Theresa Cori, née Radnitz (CZ-US), Sidney P. Colowick (US) and Gerhard Schmidt (DE-US) recognized that ATP is required to phosphorylate glucose and thus energize it for the biosynthesis of glycogen. They also discovered that if glycogen is broken down it is not hydrolyzed to glucose units but rather is converted to units of α-glucose-1-phosphate by phosphorylase in a readily reversible reaction {Caputto, 1950 #23165;Cardini, 1950 #23164;McElroy, 1951 #23167;Paladini, 1952 #23166}.

Carlos Eugenio Cardini (AR), Alejandro Paladini (AR), Ranwel Caputto (AR), and Luis Federico Leloir (AR) discovered that the glycogen synthesis reaction is uridine-5’-triphosphate + α-glucose-1-phosphate yields uridine diphosphate glucose (UDPG), which yields glycogen in the presence of glycogen synthetase {Caputto, 1950 #23165;Cardini, 1950 #23164;McElroy, 1951 #23167;Paladini, 1952 #23166}.

Luis Federico Leloir (AR), Maria A. Rongine de Fekete (AR), and Carlos Eugenio Cardini (AR) demonstrated the in vitro synthesis of amylose (the linear alpha 1,4-glycosidically linked polysaccharide) from α-glucose-1-phosphate {Leloir, 1961 #24372}.

Charles Samuel Hanes (CA), in 1940, was able to obtain starch synthesis in vitro with extracts from potato on α-glucose-1-phosphate as the glucosyl precursor {Hanes, 1940a #1847;Hanes, 1940b #23150}.

Arda Alden Green (US), Gerty Theresa Cori, née Radnitz (CZ-US), Carl Ferdinand Cori (CZ-US), and John L. Oncley (US) reported that they had purified and crystallized muscle phosphorylase {Green, 1942 #10710;Green, 1943 #2574;Cori, 1943a #23151;Cori, 1943c #23152}.

Gerty Theresa Cori, née Radnitz (CZ-US) and Arda Alden Green (US) showed that branched types of polysaccharides ensue because of the action of two enzymes {Cori, 1943b #23153}.

Earl Wilbur Sutherland, Jr. (US) and Carl Ferdinand Cori (CZ-US) found that epinephrine plays a crucial role in converting inactive forms of phosphorylase into active forms. This was found to be the case in liver as well as muscle {Sutherland, 1951a #23158;Sutherland, 1951b #23159}. Note: This was a sneek preview of the existence of regulator enzymes that constitute a whole network of cascade regulators.

Edwin Gerhard Krebs (US) and Edmond Henri Fischer (US) discovered that resting muscle contains largely phosphorylase b. They greatly clarified the issue by studying the conversion of phosphorylase b to a in muscle extracts {Krebs, 1955 #23160;Fischer, 1955 #16401}.

William H. Danforth (US), Ernst Helmreich (US), and Carl Ferdinand Cori (CZ-US) conclusively demonstrated that the conversion of phosphorylase b to a is a crucial reaction for the rapid breakdown of glycogen during muscular contraction and following epinephrine administration {Danforth, 1962 #23157}.


Ernö Annau (HU-BR-CA), Ilona Banga (HU), Bruckner Gozsy (HU), István Huszák (HU), Kámán Laki (HU-US), Frenc Bruno Staub (HU), A Blazsó (HU), and Albert Imre Szent-Györgyi (HU-US) drawing on their work and the earlier observations of Torsten Ludvig Thunberg (SE), Federico Battelli (IT), Lina Salomonovna Stern (LT-CH) and others assembled a logical sequence for the cellular oxidation of succinate: succinate—fumarate—malate—oxaloacetate. Especially significant was Szent-Györgyi’s observation that adding small amounts of oxaloacetate or malate to minced muscle suspensions evokes the utilization of an amount of oxygen far beyond that required to oxidize the added dicarboxylic acid to CO2 and water. From this and other experiments Szent-Györgyi concluded that these acids stimulate the oxidation of some endogenous substrate in the tissue, presumably glycogen, one molecule of malate or oxaloacetate promoting the oxidation of many molecules of the endogenous substrate {Annau, 1935 #23240;Annau, 1936 #23241;Szent-Györgyi, 1937 #2575}.

Frederik J. Stare (US) and Carl A. Baumann (US) noted that succinate normally formed is prevented by malonate from being oxidized to fumarate, and respiration fails owing to a lack of fumarate. They further attribute the decrease in respiration of muscle tissue in vitro to a loss of fumarate through diffusion. If, however, this loss is compensated for by the addition of fresh fumarate, respiration proceeds normally. The addition of fumarate therefore “ preserves ” the normal respiration of the tissue {Stare, 1936 #23242}.

Carl Martius (DE) and Georg Franz Knoop (DE), somewhat later, found that citrate is enzymatically oxidized to succinate by animal tissues in the sequence: citrate—alpha ketoglutarate—succinate {Martius, 1937a #8699}.

Hans Adolf Krebs (DE-GB), William Arthur Johnson (GB), and Leonard V. Eggleston (GB) observed that citric acid exerts a catalytic effect on the respiration of minced pigeon-breast muscle and that citrate is successively converted to alpha-ketoglutarate and succinate, and that oxaloacetate is converted into citrate by the addition of two carbon atoms from an unidentified source. From their conclusions and those of prior workers like Georg Franz Knoop (DE), Carl Martius (DE), and Albert Imre Szent-Györgyi (HU-US) they proposed a citric acid cycle: citrate—cis-aconitic acid— isocitrate — oxalosuccinate — alpha-ketoglutarate — succinate — fumarate — malate —oxaloacetate — citrate {Krebs, 1937a #2577;Krebs, 1940 #15963;Krebs, 1937b #23869}.

Severo Ochoa (ES-US), Feodor Felix Konrad Lynen (DE), and Helmut Scherer (DE) established oxalosuccinic acid as an intermediate, as already postulated by Martius and Knoop. They demonstrated the presence of a specific decarboxylase converting oxalosuccinate to alpha-ketoglutarate {Lynen, 1948 #23244;Ochoa, 1948 #23243}.

This cycle has been found to exist in virtually all plants, animals, and aerobic microorganisms {Krebs, 1981 #114} and has been called tricarboxylic acid cycle, citric acid cycle, and Krebs cycle.


Hugh A. Davson (GB) and James Frederic Danielli (GB) proposed a protein-lipid sandwich model for the structure of cell membranes. This Davson-Danielli model proposed the idea of two layers of phospholipids sandwiched in between two outer layers of protein. The phospholipids were oriented with their hydrophilic ends at the two surfaces and their hydrophobic tails towards the interior of the membrane. This structure explained the stability of plasma membranes because of the strong hydrophilic and hydrophobic interactions. It also accounted for the fact that lipid-soluble substances could pass through a plasma membrane easily. They proposed that this membrane was approximately eight nanometers thick and had small pores in the protein coat to allow the passage of certain molecules and ions {Danielli, 1935 #14161;Davson, 1952 #14162}.


John Burdon Sanderson Haldane (GB-IN) was the first to estimate the spontaneous mutation rate of a human gene {Haldane, 1935 #12335}.


Sterling Howard Emerson (US) and George Wells Beadle (US), using the attached-X chromosome in Drosophila melanogaster, were able to show that each crossover between chromosomes at the first division of meiosis could, with equal likelihood, involve either one of the two chromatids into which each chromosome is divided {Beadle, 1935a #22112;Beadle, 1935b #10654;Emerson, 1933 #10653}.

Royal Alexander Brink (CA-US) and Delmer C. Cooper (US) offered proof that crossing over involves an exchange of segments between homologous chromosomes in Zea mays {Brink, 1935 #22119}.


George Wells Beadle (US) and Boris Ephrussi (RU-FR) implanted embryonic eye tissue from larvae of vermilion and cinnabar mutants into larvae of normal Drosophila flies and observed that, upon metamorphosis of these larvae into mature flies, the implanted eye tissue developed into supernumerary eyes with normal eye color. It could be concluded, therefore, that the body tissues of the normal flies supply some substance that the vermilion and cinnabar mutant eye tissues are unable to synthesize, but one they can convert into the brown eye pigment. Beadle and Ephrussi then implanted the same embryonic mutant tissues into the larvae of vermilion and cinnabar mutant flies and observed that vermilion eye tissue implanted into cinnabar host larvae developed the normal eye color, whereas cinnabar eye tissue implanted into vermilion host larvae developed the mutant cinnabar eye color. Beadle and Ephrussi inferred from these observations that the synthesis of the brown eye pigment arises by the metabolic chain: Precursor to substance I to substance II to brown pigment.

The vermilion mutant would thus carry a block in the reaction that converts the precursor to substance I, whereas the cinnabar mutant would carry a block in the reaction that converts substance I to substance II. Thus, in the wild-type host larva, both mutant eye-tissue transplants are provided with substance II, which they can convert to the brown pigment. The vermilion mutant eye-tissue transplant in the cinnabar host larva is provided with substance I, which it can convert to substance II and to brown pigment. But the cinnabar mutant eye-tissue transplant in the vermilion host larva is not provided there with the substance II, which it lacks, and hence fails to form the brown pigment. Within a few years biochemical studies showed that the precursor is the amino acid tryptophan and that substances I and II are formylkynurenin and hydroxykynurenin, respectively. The genetically controlled metabolic eye color sequence could thus be written as: tryptophane to formylkynurenin (substance I) to hydroxykynurenin (substance II) to brown pigment.

The stage was now set for formulating more clearly the physiological role of genes. The normal, wild-type allele of the vermilion gene of Drosophila could be envisaged as presiding over the formation of an enzyme that catalyzes the conversion of tryptophan to formylkynurenin. The mutant allele, by contrast, has lost the capacity to form that enzyme. Hence the tissues of a homozygous mutant fly carrying the vermilion mutant gene on both of its homologous chromosomes lack the enzyme, and the metabolism of such flies is blocked at the reaction step normally catalyzed by that enzyme. The tissues of a heterozygous fly, carrying one mutant and one wild-type allele of the vermilion gene, would contain the enzyme, however, and hence can form formylkynurenin. Similarly, the mutant gene of the cinnabar gene has lost the capacity to form the enzyme that catalyzes the conversion of formylkynurenin to hydroxykynurenin, the enzyme that is normally formed under the direction of the wild-type allele. From this viewpoint, the recessive character of both the vermilion and cinnabar mutations is accounted for by the absence of an enzymatic function that the dominant, wild-type allele can supply {Beadle, 1935 #4976;Beadle, 1936a #112;Ephrussi, 1935 #14671;Ephrussi, 1942 #14672}.

George William Beadle (US) found that fat bodies of all eye color mutants studied, except those from vermilion larvae, are shown to produce v+ substance following transplantation to appropriate test animals. Malpighian tubes of wild type larvae release both v+ and cn+ substances following transplantation. Both substances can be extracted from the larval tubes with hot Ringer’s solution. It was possible to demonstrate the presence of v+ substance in the blood of wild type larvae during some part or all a 24-hour interval just prior to puparium formation {Beadle, 1937 #22113}.

Alfred Kuhn (DE), Adolf Friedrich Johann Butenandt (DE), Wolfhard Weidel (DE), and Erich Becker (DE) did very similar work in Ephestia, studying the biochemical genetics of eye-pigment synthesis {Butenandt, 1940 #14674;Kuhn, 1935 #14673}. These works represent the first step toward the one gene, one enzyme, hypothesis.


Nikolai Wladimirovich Timoféeff-Ressovsky (RU), Karl Günter Zimmer (DE), and Max Ludwig Henning Delbrück (DE-US) formulated a target theory of gene mutation, which says that a mutation can be induced if a single electron is detached by high-energy radiation {Timoféeff-Ressovsky, 1935 #14639}.


Calvin Blackman Bridges (US) showed that in Drosophila salivary gland chromosomes certain sequences recur in different regions and that these tend to be found paired in the form of synapsis of somatic chromosomes. He referred to these regions as repeats and assumed that they had arisen as duplications of the same segment (tandem repeats) {Bridges, 1935 #5029;Bridges, 1938a #12341}.

Calvin Blackman Bridges (US) and Philip N. Bridges (US) reported the accurate determination of the salivary characteristics of the Payne-R inversion (a supressor of crossing over) in Drosophila melanogaster {Bridges, 1938b #22115}.


Nagaharu U; Jang-choo Woo (KR-JP) discovered the evolutionary scheme of three allopolyploid species of Brassica based on three diploid species of genome donors. He used both the chromosome numbers and synthetic hybrids to prove the model and showed that rutabaga could be synthesized by crossing turnip and cabbage {Sageret, 1827 #17972;Nagaharu, 1935 #22255;de Candolle, 1822 #22256}.


Erwin Bünning (DE) proved the genetic origin of biological rhythms. He found that circadian rhythms persisted in the bean plant Phaseolus and the fruit fly Drosophila, even though generation after generation had been raised in environments completely lacking cues to the passage of time {Bünning, 1935 #9436;Bünning, 1936 #9437}.


J. Arthur Ramsay (GB) investigated water loss from the American cockroach, Periplaneta americana. He found that very high wind speeds significantly increased rates of water loss, attributing this result to eddies set up within the tracheal system, increasing water evaporation from inside the tracheal system itself. Ramsay provided evidence for what was later termed the critical or transition temperature for water loss. Water-loss rates increase rapidly above this temperature. Ramsay then performed the first biophysical measurements on surface (cuticle) waxes, finding that the surface tension of wax-coated droplets decreased dramatically at about 30°C, right where water loss began to increase. He concluded that melting of these same lipids on the cockroaches' cuticle was responsible for increased transpiration {Ramsay, 1935 #21251}.

In a striking example of convergent evolution, most terrestrial organisms have similar lipid waterproofing layers {Hadley, 1989 #21252}.


Wendell Meredith Stanley (US) reported the crystallization of pure tobacco mosaic virus to the world. This gave rise to the controversy of whether a virus is alive or dead. Most microbiologists today consider that they are not alive, because they are acellular {Stanley, 1935a #665;Stanley, 1935b #13985;Stanley, 1935c #13986;Stanley, 1935d #13987}.


James P. Cleary (US), Paul J. Beard (US), and Charles E. Clifton (US) concluded that the population of a bacterium in continuous culture will not exceed a specific maximum cell number even when the effects of inhibitory substances are reduced to a low level because the major population limiting factor appears to be the amount of energy and building material available per cell per unit time {Cleary, 1935 #14627}.


Frank Macfarlane Burnet (AU) and Diana H. Bull (AU) grew the influenza virus in chick embryos {Burnet, 1935 #6994;Burnet, 1943 #6995}.


St. Louis encephalitis was first recognized as a unique clinical entity in association with a large outbreak of the disease at St. Louis, MO in the summer of 1933 {Committee, 1935 #2376}.


Leslie T. Webster (US), Anna D. Clow (US), and Johannes H. Bauer (US) found that the Anopheles quadrimaculatus (mosquito) will harbor the virus of St. Louis encephalitis for 21-42 days following a blood meal from an infected animal {Webster, 1935 #2377}.


Kenneth M. Smith (GB) and John G. Bald (GB) were the first to describe tobacco necrosis disease as being caused by a virus {Smith, 1935 #2044}.


William Trager (US) performed the first successful infection of insect tissue in vitro by an insect virus (grasserie). He inoculated primary cultures of ovarian tissue from Bombyx mori with dilutions of hemolymph from diseased larvae {Trager, 1935 #8551}.


Meredith Hoskins (US) reported that when rhesus monkeys (Macacus rhesus) were inoculated simultaneously with neurotropic and viscerotropic strains of yellow fever virus the neurotropic strain appeared to have a very definite protective effect by reducing the virulence of the viscerotropic strain. This is referred to as the interference phenomenon and was first demonstrated in animals by Hoskins {Hoskins, 1935 #5728}.


Leonell C. Strong (US) established the C3H inbred strain of mice for the study of spontaneous carcinoma of the mammary gland {Strong, 1935 #11584}.


Emmy Klieneberger (GB) discovered L form bacteria when she found them growing in association with Streptobacillus moniliformis. Originally, she thought they represented a unique organism, but other workers subsequently showed them to be a bacterial variant of organisms such as Streptobacillus moniliformis {Klieneberger, 1935 #5747}. These organisms lack cell walls but are not mycoplasmas.


Gerhard J. Domagk (DE) discovered that a red dye compound, Prontosil rubrum (4-sulfonamido-2, 4-diamino-azobenzene hydrochloride) is very low in toxicity to animals but very active against streptococcal infections in animals {Domagk, 1935a #113}.

Jacques Gustave Marie Tréfouël (FR), Thérèse Tréfouël, née Boyer (FR), Filomena Nitti (IT-FR), and David Bovet (CH-FR), working at the Pasteur Institute, discovered that the antibacterial action of the drug prontosil is due to it being converted to sulfanilamide within the animal body {Tréfouël, 1935 #599}.

Leonard Colebrook (GB), Méave Kenny (GB), and Anthony W. Purdie (GB) provided overwhelming evidence of the efficacy of both Prontosil and sulfanilamide in streptococcal septicemia (bloodstream infection), thereby ushering in the sulfonamide era {Colebrook, 1936a #14640;Colebrook, 1936b #14641;Colebrook, 1937 #14642}.

In retrospect, the introduction of Prontosil marked a turning point in the history of medicine. As the first of the compounds called sulfonamides or, more familiarly, sulfa drugs, Prontosil initiated a revolution in the therapeutics and management of bacterial infections.


Gerhard J. Domagk (DE) discovered that the germicidal properties of quaternary ammonium compounds require that at least one of the four radicals consist of a long-chain aliphatic group {Domagk, 1935b #7966}.


Jerome T. Syverton (US), George Packer Berry (US), R.W. Harrison (US) and Elizabeth Moore (US) successfully cultured the virus of St. Louis encephalitis in tissue culture {Harrison, 1936 #2378;Syverton, 1935 #19483}.


Albert Spear Hitchcock (US) and Mary Agnes Chase (US) prepared the Manual of Grasses of the United States (1935). Its usefulness continues through today in the form of a 2nd edition in 1950 {Hitchcock, 1951 #8969;Hitchcock, 1971 #20276}.


Earl W. Flosdorf (US) and Stuart Mudd (US) promoted an easy way to store blood plasma. It was first frozen, and then dried into flakes in a vacuum. Packed in tiny ampullae, the plasma was shipped with a pint of sterile water and tubing {Flosdorf, 1935 #24373}. Note: this process is called lyophilization or freeze-drying.


Shoichi Imamura (JP) while examining the serum type of pigs, found the existence of the new agglutinin which had never been described, and named this agglutinin anti-QLandsteiner. All the types of human erythrocytes are divided into two classes, namely Q or what is agglutinated by anti-Q agglutinin and q or what is not. The Q agglutinogen is found at a rate of about 32% in all the blood types {Imamura, 1935 #24757}.


Helen S. Mitchell (US) and Warren M. Dodge (US) reported changes in the lens tissue of rats subsisting on a ration containing a high proportion of lactose {Mitchell, 1935 #23388}.


Dempsie B. Morrison (US), Alan Hisey (US), and Erich Peters (US) established that the combination of oxygen with hemoglobins takes place according to the ratio, Fe/O2, one gram-atom of pigmentary iron per gram-molecule of oxygen fixed {Morrison, 1935 #2422}.


Eli Kennerly Marshall, Jr. (US) and Morris Rosenfeld (US) observed that in respiratory depression anoxia provides a major ventilatory drive mediated through the sino-aortic mechanism. Their observation recognized that often when the mammal is threatened with anoxemia, “it may adapt itself … to a primitive type of respiratory control (the sino-aortic rather than central) which is normal for lower vertebrates” {Marshall, 1935 #10806;Rosenfeld, 1936 #10807}. See, Corneille and Jean Francois Heymans, 1927.

Note: It is a prime rule in accident rooms not to give oxygen to patients depressed with morphine, barbiturates, or allied drugs.


Mary Broadfoot Walker (GB) introduced the first effective treatment for myasthenia gravis, prostigmin {Walker, 1935 #22614}.


Carlyle F. Jacobsen (US) first discovered that damage to the primate prefrontal cortex (PF) appears to cause a short-term memory deficit {Jacobsen, 1935 #16271;Jacobsen, 1936 #16272}.

Shintaro Funahashi (US), Charles J. Bruce (US), and Patricia S. Goldman-Rakic (US) found evidence to strengthen the evidence that the dorsolateral prefrontal cortex of the monkey participates in the process of working or transient memory and further indicate that this area of the cortex contains a complete “memory” map of visual space {Funahashi, 1989 #16270}.


Armand James Quick (US), Margaret Stanley-Brown (US), and Frederic W. Bancroft (US) developed the one-stage prothrombin-time technique using rabbit brain extract. This test detects the amount of prothrombin present in blood plasma and determines prothrombin-clotting time {Quick, 1935a #11704;Quick, 1935b #13464}. The technique assumed that, given enough tissue, calcium, and fibrinogen, there is only one factor limiting the time course of clotting: prothrombin. It is now recognized that this result is limited by deficiencies of factors additional to prothrombin, but this does not diminish the importance of this technique in the control of coumarin therapy. See, Whipple, 1913.

The one-stage prothrombin-time made possible the immediate differentiation between the coagulation defect in hemophilia and that in obstructive jaundice.

Emory D. Warner (US), Kenneth M. Brinkhous (US), and Harry P. Smith (US) developed a method of measuring prothrombin, which became known as the two-stage technique {Warner, 1936 #11705}.


Hugh Leslie Marriott (GB) and Alan Kekwick (GB) introduced the continuous drip blood transfusion method, in which blood flows from a flask {Marriott, 1935 #7911}. This was made possible by Luis Agote’s discovery in 1914. Earlier transfusions were directly from donor to recipient.


Claude Schaeffer Beck (US) presented his technique for development of a blood supply to the heart by operation. It prescribed grafting a flap of the pectoralis muscle over the exposed epicardium to provide a new blood supply {Beck, 1935 #11325}.

Frank Mason Sones, Jr. (US) accidently performed the first selective coronary angiography when he accidently injected dye into the right coronary artery instead of into the cardiac valve as intended. Instead of fibrillating, the man's heart went into asystole, and Sones shouted at the patient to cough, which successfully restarted the heart beating {Dotter, 1958 #11280;Meyers, 2007 #20213}.

Arthur M. Vineberg (CA) introduced the Vineberg procedure, which consists of the revascularization of the entire heart by internal mammary artery implantation, epicardiectomy and free omental graft {Vineberg, 1946 #11326;Vineberg, 1966 #9815}.

Vasilii I. Kolessov (RU) performed the first internal mammary artery-coronary artery anastomosis {Kolessov, 1967 #11327}.

W. Dudley Johnson (US), Robert J. Flemma (US), Derward Lepley, Jr. (US), and Edwin H. Ellison (US) introduced modern coronary by-pass surgery with their method of myocardial revascularization. Veins are usually inserted into an area of normal artery; however, if a second area of atherosclerosis occurs (commonly in the mid-anterior descending artery), the arteriotomy extends across the plaque into normal artery on each end. The vein is sutured as a patch graft always extending the anastomosis to normal artery proximally and distally. With progressive atherosclerosis this maneuver preserves bidirectional flow {Johnson, 1969 #14522}.

Donald B. Effler (US), René Gerónimo Favaloro (AR), Laurence K. Groves (US), Chalit Cheanvechai (TH), Robert A. Quint (US), and Frank Mason Sones, Jr. (US), beginning in 1967, performed some of the first coronary by-pass operations using the patients’ native saphenous veins as autografts {Effler, 1970 #15197;Favaloro, 1971 #11328}.

H. Edward Garrett (US), Edward W. Dennis (US), and Michael Ellis DeBakey (US) performed an autogenous saphenous vein by-pass from the ascending aorta to the anterior descending coronary artery. The patient was a 42-year-old man who had extensive occlusive disease of the coronary artery and angina pectoris {Garrett, 1973 #11300}. This team first performed the operation in 1964.


Kenneth M. Lynch (US) and W. Atmar Smith (US) were the first to report carcinoma of the lung in a patient with “asbesto-silicosis” {Lynch, 1935 #23722}.


Allen Oldfather Whipple (US), William Barclay Parsons (US), and Clifton R. Mullins (US) introduced a two-stage radical pancreaticoduodenectomy to treat carcinoma of the ampulla of Vater {Whipple, 1935 #14486}. Twenty-eight months was the longest survival time course of the three cases reported. This was not the first time such an operation was performed.

Allen Oldfather Whipple (US) described the first reported case of a one-stage pancreaticoduodenectomy for a carcinoma of the head of the pancreas {Whipple, 1945 #18934}. See, Alessandro Codivilla, 1898.


Arnold Rice Rich (US) found carcinoma of the prostate in 14% of all autopsies and in 28% of those aged over 70 years {Rich, 1935a #11583}.


Pyotr Kuzmich Anokhin (RU) developed the concept of feedback. Furthermore, he elaborated the theory of functional systems (FS) which tied together subtle neurophysiological mechanisms and integral activity of an individual {Anokhin, 1935 #22960;Anokhin, 1937 #22961}.


Burrhus Frederic Skinner (US), a leading behaviorist and proponent of operant conditioning, invented the Skinner box for facilitating experimental observations. His main scientific works include The Behavior of Organisms (1938), and Verbal Behavior (1957). He founded behaviorism {Skinner, 1935 #7682;Skinner, 1938a #4913;Skinner, 1938b #7683;Skinner, 1957 #4912}.


John Ridley Stroop (US) was the first to think of combining a word with object/property dimensions, creating the now famous situation of response conflict {Stroop, 1935 #9417}. An example of this phenomenon is that when asked to name the color of ink in which an incompatible color word is printed (e.g., to say "red" aloud in response to the stimulus word GREEN printed in red ink), people take longer than to name the same ink color in a suitable control condition (e.g., to say "red" to the stimulus XXXXX printed in red ink). This has been called the Stroop effect.


Arthur George Tansley (GB) coined and defined the term ecosystem. "The weakness of Clements is. . . that vegetation is an organism and therefore must obey the laws of development of what we commonly know as organisms. . .. But the more fundamental conception is, as it seems to me, the whole system (in the sense of physics) including not only the organism-complex, but also the whole complex of physical factors forming what we call the environment of the biome…Though the organism may claim our primary interest, when we are trying to think fundamentally we cannot separate them [organisms] from their special environment, with which they form one physical system" {Tansley, 1935 #8295}.


Julia Anna Gardener (US), T. Wayland Vaughan (US), and Willis Parkison Popenoe (US) helped establish standard stratigraphic sections for Tertiary rocks in the Southern Caribbean, Coastal Plain of Texas, and the Rio Grande Embayment in Northeast Mexico {Gardner, 1935 #8976;Gardner, 1943 #8977}.



“If there is any doubt as to whether a person is or is not dead, apply lightly roasted onion to his nostrils, and if he is alive, he will immediately scratch his nose.” Johannes de Mirfield (GB) {Hartley, 1936 #11775}.


Henry Hallett Dale (GB) and Otto Loewi (DE-US) were awarded the Nobel Prize in Physiology or Medicine for their discoveries relating to chemical transmission of nerve impulses.


Harold Clayton Urey (US), Adriaan H.W. Aten, Jr. (US), Albert S. Keston (US), George B. Pegram (US), John R. Huffman (US), Harry G. Thode (US), and Marvin Fox (US) devised laboratory procedures for preparing high concentrations of such isotopes as carbon-13, oxygen-18, and nitrogen-15 {Urey, 1936a #10996;Urey, 1936b #10997;Urey, 1937 #10998}.


John Tileston Edsall (US) used Raman spectroscopy to show that both the amino and carboxyl groups of amino acids are charged at isoelectric pH {Edsall, 1936 #11245;Edsall, 1937a #11246;Edsall, 1937b #11247}.


Reynold C. Fuson (US), Robert E. Christ (US), Richard Johann Kuhn (DE) and Colin John Owen Rhonabwy Morris (GB) synthesized vitamin A (retinol) {Fuson, 1936 #2951;Kuhn, 1937 #2952}.


Thomas William Birch (AU-GB) and Paul György (GB) discovered vitamin B6 (pyridoxine hydrochloride) {Birch, 1936 #14637}.


Barend Coenraad Petrus Jansen (NL) authored a quantitative chemical test for vitamin B1 based on the oxidation of the vitamin into a yellow substance with intense blue fluorescence (thiochrome) as discovered by George Barger (GB), Franz Bergel (AT-GB), Alexander Robertus Todd (GB), and Rudolph Albert Peters (GB) {Jansen, 1936 #2750;Barger, 1935 #2965;Peters, 1935 #2966}.


Hans Christian Hagedorn (DK), B. Norman Jensen (DK), Niels B. Krarup (DK), and Inger Woodstrup (DK) developed protamine insulinate, one of the first successful longer-acting exogenous insulins {Hagedorn, 1936 #11550}.


Linus Carl Pauling (US) and Charles Dubois Coryell (US) demonstrated that in deoxyhemoglobin the bonds between iron and nitrogen are ionic and weak whereas in oxyhemoglobin they are covalent and stronger {Pauling, 1936 #940}.


Daniel Bovet (CH-FR-IT) and Anne Marie Staub (FR) synthesized the first histamine receptor antagonists (popularly referred to as the classical antihistamines but now called H1-receptor antagonists). These early studies of the antihistamines were qualitative, for example, the demonstration of their effectiveness in protecting against bronchospasm produced in guinea pigs by anaphylaxis or administration of histamine. Though qualitative, these studies yielded compounds that remain major ligands to define histamine receptors, e.g., mepyramine (pyrilamine) {Bovet, 1936 #11356;Staub, 1939 #9177}.


André Michel Lwoff (FR), Marguerite Lwoff (FR), Bert Cyril James Gabriel Knight (FR), and John Howard Mueller (FR) demonstrated that nicotinic acid is an essential growth factor for some bacteria {Knight, 1937 #16816;Lwoff, 1936a #16815;Lwoff, 1936b #17485;Mueller, 1937 #16817}. This led to the use of bacterial growth as a quantitative assay of growth factors.


Klaas Tammo Wieringa (NL) discovered Clostridium aceticum, the first acetogenic bacterium found to grow with hydrogen as an energy source and carbon dioxide as a carbon source {Wieringa, 1936 #15539}.


Chester Hamlin Werkman (US), E.A. Zoellner (US), Henry Gilman (US), and Howard Reynolds (US) were the first to isolate phosphoglyceric acid from bacteria {Werkman, 1936 #11210}. Along with Robert William Stone (US) and Wilbur Paul Wiggert (US) they reported the formation of phosphoglyceric acid and related intermediary compounds in the dissimilation of glucose by a variety of heterotrophic bacteria {Stone, 1936a #11211;Stone, 1936b #11212;Stone, 1937 #11213;Wiggert, 1938 #11214}.


Georg Franz Knoop (DE) and Carl Martius (DE) established some of the intermediate stages of the oxidation of citric acid when they discovered the following sequence of reactions in liver tissue: citrate — cis-aconitate — isocitrate — oxalosuccinate — alpha-ketoglutarate.

The end-product of these sequences, alpha-oxoglutaric acid (alpha-ketoglutaric acid), was already known as an intermediate product because it is formed from the amino acid glutamate and had been shown to be oxidized to succinic acid {Knoop, 1936 #2576}.

Carl Martius (DE) and Georg Franz Knoop (DE) demonstrated that citric acid could be formed from pyruvic acid {Martius, 1937a #8699}. Carl Martius (DE) worked out the sequence citric-aconitic-isocitric-oxalosuccinic acid by the action of aconitase and isocitrate dehydrogenase {Martius, 1937b #8700}.

Erich Adler (SE), Hans Karl August Simon von Euler-Chelpin (DE-SE), Gunnar Günther (SE), and Marianne Plass (SE) determined that triphosphopyridinenucleotide (TPN) is the coenzyme for isocitrate dehydrogenase {Adler, 1939 #20840}. Synonyms for TPN are Warburg's coenzyme II and codehydrogenase II.


Edward Lawrie Tatum (US), Harland Goff Wood (US), William Harold Peterson (US), Milton Silverman (US) and Chester Hamlin Werkman (US) were among the first to demonstrate the role of vitamin B1 and co-carboxylase in bacterial metabolism {Tatum, 1936 #11215;Silverman, 1938 #11205;Silverman, 1939 #11206}.


Béla Tankó (HU), Gladys Macphail James (GB), William Owen James (GB), Arthur Hugh Bunting (GB), C.R.C. Heard (GB), and Charles Samuel Hanes (GB) demonstrated that glycolytic systems in higher plants are essentially the same as those discovered in animals, and yeast {Tankó, 1936 #1843;Hanes, 1940a #1847;James, 1940 #1846;James, 1941 #1844;James, 1944 #1845}.


David Bodian (US) developed a new method for staining nerve fibers and nerve endings in mounted paraffin sections {Bodian, 1936 #9410}.


Peter A. Gorer (GB), using inbred strains of mice, discovered four blood group antigens. The growth and rejection of a tumor correlated with the expression of antigen II. Gorer formulated the concept of tissue transplantation as, “normal and neoplastic tissues contain iso-antigenic factors which are genetically determined. Iso-antigenic factors present in the grafted tissue and absent in the host are capable of eliciting a response which results in the destruction of the graft.” This represents the discovery of what came to be known as the H-2 genetic region in mice {Gorer, 1936 #6147;Gorer, 1937 #2188}.


Clara J. Lynch (US), Thomas P. Hughes (US), Leslie T. Webster (US), and Anna D. Clow (US) were the first to demonstrate that a host gene can control resistance to disease induced by an animal virus {Lynch, 1936 #8687;Webster, 1936 #8688}. This resistance was directed against flaviviruses.


Lewis John Stadler (US) and George F. Sprague (US) discovered that ultraviolet light is mutagenic in maize {Stadler, 1936 #15802}.


Cyril Norman Hugh Long (GB-US) and Francis D.W. Lukens (US) found that removing the adrenal gland and pituitary gland of a diabetic animal reduced the blood level of glucose and thus the severity of its diabetic state {Long, 1936 #10957}. This would later be linked to adrenocorticotropic hormone (ACTH) of the anterior pituitary gland. ACTH stimulates the release of other hormones, which encourage the conversion of amino acids to glucose.


Torbjörn Oskar Caspersson (SE) demonstrated that the so-called euchromatin bands on chromosomes represent areas, which are very rich in nucleic acids {Caspersson, 1936 #2475}.


Calvin Blackman Bridges (US), Eleanor Nichols Skoog (US), and Ju-chi Li (CN) chose the notopleural deficiency in Drosophila melanogaster to establish a close correspondence between the loci on the genetic maps, deduced from linkage studies, and the chromosome localities, deduced from study of the transverse bands which form a diversified series along the salivary chromosome {Bridges, 1936 #22117}.


Carl D. LaRue (US) successfully established embryo cultures of different gymnosperms {LaRue, 1936 #15257}.


George Henry Hepting (US) and Dorothy J. Blaisdell (US) described the mechanisms with which trees restrict the development of decay and discoloration in stems to “tissues extant at time of wounding.” This phenomenon is now known as compartmentalization {Hepting, 1936 #11140}.


John Zachary Young (GB) discovered the giant nerve fibers of the squid, Loligo forbesi. This made possible many important experiments in neurophysiology {Young, 1936 #3290;Young, 1938 #21269;Young, 1939 #21254}.


William Frederick Windle (US) and Marvin F. Austin (US) described the routes of the earliest axons in the central nervous system of the chick embryo, assessed at several stages between the second and sixth days of incubation. They focused on the descending, or reticulospinal, axons arising from hindbrain nuclei {Windle, 1936 #20389}.

William L.R. Cruce (US), Sherry L. Stuesse (US), and R. Glenn Northcutt (US) used retrogradely transported axonal tracers (horseradish peroxidase and Fluoro-Gold) to identify groups of brainstem neurons that projected to the spinal cord, in two cartilaginous fishes, the Thornback Guitarfish, and the Horn Shark. They identified numerous distinct reticular nuclei in these elasmobranchs, consistent with a complex organization like the reticular formation in other vertebrates {Cruce, 1999 #20390}.

John G. New (US), Bethany D. Snyder (US), and Katherine L. Woodson (US) traced axons descending to the spinal cord in the Channel Catfish and found that most neurons projecting to the spinal cord are located in the reticular formation of the hindbrain. Both ascending and descending reticular formation projections are of great clinical importance in humans, as they can be damaged or destroyed by strokes, spinal cord injuries, and astrocytomas {New, 1998 #20391}.


Herald Rea Cox (US), Peter Kosciusko Olitsky (US), Joseph W. Beard (US), and Harold Finkelstein (US) developed vaccines for equine encephalomyelitis {Cox, 1936 #2382;Beard, 1937 #2383}.


Curt Stern (DE-US), while working with several sex-linked alleles of Drosophila, revealed a previously unrecognized phenomenon, mitotic crossing over {Stern, 1936 #10821}.


Frederick Charles Bawden (GB), Norman Wingate Pirie (GB), John Desmond Bernal (GB), and Isadore Fankuchen (US) demonstrated that tobacco mosaic virus contains phosphorus as a component of a phospho-ribonucleic acid {Bawden, 1936 #5730;Bernal, 1941 #13992}.


Frederick Charles Bawden (GB), Norman Wingate Pirie (GB), Hubert S. Loring (US), Wendell Meredith Stanley (US), and Max Augustus Lauffer (US) demonstrated that all plant viruses tested up to this time were pure nucleoprotein. This was the first indication that nucleic acids, found in all cells, is found also in acellular "life" {Bawden, 1936 #5730;Bawden, 1937a #1822;Bawden, 1937b #14709;Bawden, 1943 #1824;Loring, 1939 #1823;Stanley, 1939 #1825}.


George P. Berry (US) and Helen M. Dedrick (US) reported the changing of rabbit fibroma virus (Shope) into infectious myxomatosis virus (Sanarelli). This Berry-Dedrick phenomenon has been referred to variously as transformation, recombination, multiplicity of reactivation, and non-genetic reactivation {Berry, 1936 #5731}.


Frank Macfarlane Burnet (AU) and Dora Lush (AU) published a paper showing that bacteriophages can sport mutants whose plaques have a distinctly different appearance from those of the ordinary wild-type {Burnet, 1936 #12141}.


Felix G. Gustafson (US) obtained the first successful growth of the parthenocarpic fruits (tomato, grape, and fig) by applying auxin on unfertilized ovaries {Gustafson, 1936 #14690}.


K. Kanazawa (JP) was the first to grow the rabies virus in tissue culture {Kanazawa, 1936 #7005}.


Henry Edward Shortt (GB), T. Ramachandra Rao (IN), and C.S. Swaminath (IN) reported culturing the virus of dengue fever (breakbone fever) on the chorioallantoic membrane of chick embryos {Shortt, 1936 #2371}.


Henry Edward Shortt (GB), S. Ramachandra Rao (IN), C.S. Swaminath (IN), and Chintaman Govind Pandit (IN) reported culturing the virus of pappataci fever (phlebotomus fever, sand-fly fever, three-day fever) on the chorioallantoic membrane of chick embryos {Shortt, 1936 #2371;Shortt, 1938 #2372}.


Jacob Traum (US) discovered the first calicivirus, vesicular exanthema virus of swine {Traum, 1936 #25951}.


Jean Cuillé (FR) and Paul-Louis Chelle (FR) reported that scrapie can be serially transmitted to sheep and that it passes bacterial filters. They noted that signs of the disease did not appear until more than one year had elapsed from the time of exposure to the contagious material {Cuillé, 1936a #6194;Cuillé, 1938 #8277;Cuillé, 1939 #14742}. Note: Scrapie gets its name from the tendency of affected animals to rub against fence posts during the excitable phase of the disease. See, Leopoldt, 1759

Josef Gerstmann (AT-US), Ernst Sträussler (AT), and Ilya Mark Scheinker (AT) described a disease characterized by degeneration of the nervous system, starting usually in the fourth or fifth decade of life with slowly developing dysarthria and cerebellar ataxia and then dementia, accompanied by spinocerebellar and corticospinal tract degeneration, and absence of leg reflexes. Death follows 2 to 10 years after the onset of symptoms {Gerstmann, 1936 #13274}. Note: Today this is known as Gerstmann-Sträussler-Scheinker syndrome, one of the amyloid dependent subacute spongioform encephalopathies associated with prion infection. It behaves in an autosomal dominant pattern.

Björn Sigurdsson (IS), Halldór Grimsson (IS), Páll A. Pálsson (IS), and Anna Tryggvadóttir (IS) developed the concept of slow viruses (maedi-visna virus, scrapie prion) {Sigurdsson, 1952 #25882;Sigurdsson, 1953 #25883;Sigurdsson, 1957 #8669;Sigurdsson, 1958 #25885}.

Bjorn Sigurdsson (IS), Páll Agnar Pálsson (IS), and Halldór Grímsson (IS) determined that both visna and maedi are slow virus infections of sheep {Sigurdsson, 1954a #8668;Sigurdsson, 1957 #8669}. Note: Visna means wasting and is characterized by progressive neurologic impairment and inanition. Maedi means shortness of breath and is characterized as a chronic pneumonia. Sigurdsson developed the concept of slow infection and described it as: (a) a long but rather predictable incubation period of months to years, during which the infectious agent produces clinically unapparent but progressive pathologic damage; and (b) a protracted course, once clinical signs have appeared, generally ending in serious disease or death {Sigurdsson, 1954b #8670}.

Daniel Carleton Gajdusek (US) and Vincent Zigas (AU) were the first to describe kuru (shivering or trembling), a human spongiform encephalopathy discovered among the Fore people of New Guinea {Gajdusek, 1957 #6193}.

Wlliam J. Hadlow (US) pointed out the similarity between the neuropathology of scrapie in sheep with that of kuru in man. He arrived at this conclusion after observing brain sections of kuru victims prepared by Igor Klatzo (DE-US) {Hadlow, 1959 #8271;Klatzo, 1959 #8272}.

Herbert B. Parry (GB) suggested that scrapie, a spongiform encephalopathy of sheep, is an inherited but also transmissible disease {Parry, 1962 #8273}.

Daniel Carleton Gajdusek (US), Clarence Joseph Gibbs, Jr. (US), and Michael P. Alpers (US) were the first to transmit a human prion disease, kuru, to experimental animals {Gajdusek, 1966 #5685;Gajdusek, 1977a #5686}.

Colin L. Masters (AU), Michael P. Alpers (AU), Daniel Carleton Gajdusek (US), Clarence Joseph Gibbs, Jr. (US), and Byron A. Kakulas (AU) induced experimental kuru in the gibbon and sooty mangabey and Creutzfeldt-Jakob disease in the pigtailed macaque {Masters, 1976 #25063}.

Tikvah Alper (ZA-GB), William A. Cramp (GB), David A. Haig (GB), Matthew C. Clarke (GB), and John Stanley Griffith (GB) developed the hypothesis that some transmissible spongiform encephalopathies are caused by an infectious agent consisting solely of proteins {Alper, 1967 #21828;Griffith, 1967 #21829}.

Elizabeth S. Williams (US) and Stephanie Ming Young (US) discovered that chronic wasting disease of deer is a transmissible spongiform encephalopathy caused by a prion {Williams, 1980 #25746}.

Patricia A. Merz (US), Robert A. Somerville (GB), Henry M. Wisniewski (US), and Khalid Igbal (US) used electron microscopy to find and name scrapie-associated fibrils {Merz, 1981 #25062}.

David C. Bolton (US), Michael P. McKinley (US), and Stanley Benjamin Prusiner (US) argued that a protein, not a virus or any other known parasite, was the infectious agent responsible for scrapie {Bolton, 1982 #1363}. Prusiner named the infectious agents prions (proteinaceous infectious particle) {Prusiner, 1982 #5674}.

Patricia A. Merz (US), Robert A. Somerville (GB), Henry M. Wisniewski (US), Laura Manuelidis (US), and Elias E. Manuelidis (US) found fibrils associated with Creutzfeldt-Jakob disease {Merz, 1983 #25061}.

Stanley Benjamin Prusiner (US), Michael P. McKinley (US), Karen A. Bowman (US), David C. Bolton (US), Paul E. Bendheim (US), Darlene F. Groth (US), George G. Glenner (US), Hansruedl Büeler (CH), Adriano Aguzzi (CH), Andreas Sailer (CH), R.A. Greiner (CH), Peter Autenried (US), Michel Aguet (CH), Charles Weissmann (CH), Ana Serban (US), Ruth Koehler (US), Dallas Foster (US), Marilyn Torchia (US), Dennis R. Burton (US), Shu-Lian Yang (CN-US), Stephen J. DeArmond (US), Neil Stahl (US), and Ruth Gabizon (IL) found that a modified form of the normal prion protein (PrP), called protein prion scrapie (PrPsc), is essential for infectivity {Prusiner, 1983 #5682;Prusiner, 1991 #5681;Büeler, 1993 #5680;Prusiner, 1993a #5684;Prusiner, 1993b #5683}. Note: It now appears that kuru, Creutzfeldt-Jakob disease (CJD), Gerstmann-Sträussler-Scheinker disease (GSS), and fatal familial insomnia (FFI) are all caused by a mutation of the normal gene PrPc to the abnormal gene PrPsc. The various disease phenotypes most likely arise from differences in familial genotypes and environmental factors.

Bruce Chesebro (US), Richard E. Race (US), Kathy Wehrly (US), Jane Nishio (US), Marshall Bloom (US), David Lechner (US), Sven Bergstrom (US), Ken Robbins (US), Leonard Mayer (US), and Jerry M. Keith (US) identified scrapie prion protein-specific mRNA in scrapie-infected and uninfected brain {Chesebro, 1985 #25064}.

Robert S. Sparkes (US), Melvin I. Simon (US), Vivian H. Cohn (US), R.E. Keith Fournier (US), Janice Lem (US), Ivana Klisak (US), Camilla Heinzmann (US), Cila Blatt (US), Michael Lucero (US), Thuluvancheri K. Mohandas (US), Stephen J. DeArmond (US), David Westaway (US), Stanley Benjamin Prusiner (US), and Leslie P. Weiner (US) determined that prion protein (PrP) is encoded by a gene on the short arm of chromosome 20 in humans {Sparkes, 1986 #5679}.

Yu-Cheng Liao (US), Roger V. Lebo (US), Gary A. Clawson (US), and Edward A. Smuckler (US) identified and characterized a human complementary DNA whose protein product is the major component of scrapie-associated fibrils in Creutzfeldt-Jakob disease, kuru, and Gerstmann-Straussler syndrome. {Liao, 1986 #5895}.

Gerald A.H. Wells (GB), Anthony Charles Scott (GB), Charles Todd Johnson (GB), Robert F. Gunning (GB), Rodney Douglas Hancock (GB), Moffitt D. Jeffrey (GB), Michael Dawson (GB), and Raymond Bradley (GB) reported a new spongiform encephalopathy in cattle {Wells, 1987 #8276}.

James Hope (GB), Laura J.D. Reekie (GB), and Nora Hunter (GB), Gerd Multhaup (DE), Konrad Beyreuther (DE), Heather White (GB), Anthony C. Scott (GB), Michael J. Stack (GB), Michael Dawson (GB), and Gerald A.H. Wells (GB) showed that bovine spongiform encephalopathy (BSE), commonly called mad cow disease, is caused by a prion {Hope, 1988 #5894}.

Karen Hsiao (US), Harry F. Baker (GB), Tim J. Crow (GB), Mark Poulter (GB), Frank Owen (GB), Joseph D. Terwilliger (US), David Westaway (US), Jurg Ott (US), and Stanley Benjamin Prusiner (US) showed that in Gerstmann-Straüssler-Scheinker (GSS) syndrome prion protein (PrP) codon 102 is linked to the putative gene for the syndrome in two pedigrees, providing the best evidence to date that this familial condition is inherited despite also being infectious, and that substitution of leucine for proline at prion protein (PrP) codon 102 may lead to the development of Gerstmann-Straüssler-Scheinker (GSS) syndrome {Hsiao, 1989 #5889}.

Dmitry Goldgaber (US), Lev G. Goldfarb (US), Paul Brown (US), David M. Asher (US), W. Ted Brown (US), Scott Lin (US), James W. Teener (US), Stephen M. Feinstone (US), Richard Rubenstein (US), Richard J. Kascsak (US), John W. Boellaard (DE) and Daniel Carlton Gajdusek (US) found a mutation in the prion protein (PrP) gene of two patients with Creutzfeldt-Jakob disease (CJD) from one family and a second mutation in the same gene in three patients with Gerstmann-Straüssler-Scheinker (GSS) from another family. The mutation in two related familial CJD patients changed glutamine in position 200 to lysine {Goldgaber, 1989 #5890}.

Matti Haltia (FI), Jussi Kovanen (FI), Lev G. Goldfarb (US), Paul Brown (US), and Daniel Carleton Gajdusek (US) found a new G-to-A mutation in codon 178 of the PRNP gene (resulting in a substitution of asparagine for aspartic acid) in the DNA of eight family members with typical Creutzfeldt-Jakob disease (CJD) {Haltia, 1991 #5891}.

Rossella Medori (US), Pasquale Montagna (IT), Hans-Juergen Tritschler (DE), Andrea C. LeBlanc (CA), Pietro Cortelli (IT), Paolo Tinuper (IT), Elio Lugaresi (IT), Pierluigi Gambetti (US), Federico Villare (US), Valeria Manetto (US), Hsiao Ying Chen (US), Run Xue (US), Suzanne M. Leal (US), Patrizia Avoni (IT), Mirella Mochi (IT), Agostino Baruzzi (IT), Jean Jacques Hauw (FR), Jurg Ott (US), and Lucila Autilio-Gambetti (US) identified fatal familial insomnia (FFI) as a disease caused by a prion linked to a mutation in codon 178 of the prion protein gene (PRNP) resulting in the substitution of aspartic acid with asparagine (D178N) {Medori, 1992a #5675;Medori, 1992b #5676}. Note: This disease is characterized by marked decrease or loss of the ability to sleep, dysautonomia and motor signs, and pathologically by preferential atrophy of thalamic nuclei.

Lev G. Goldfarb (US), Robert B. Petersen (US), Massimo Tabaton (IT), Paul Brown (US), Andrea C. LeBlanc (CA), Pasquale Montagna (IT), Pietro Cortelli (IT), Jean Julien (FR), Claude Vital (FR), William W. Pendelbury (US), Matti Haltia (FI), Peter R. Willis (US), Jean Jacques Hauw (FR), Paul E. McKeever (US), Lucia Monari (IT), Bertold Schrank (DE), Gary D. Swergold (US), Lucila Autilio-Gambetti (US), Daniel Carleton Gajdusek (US), Elio Lugaresi (IT), Pierluigi Gambetti (US), and Michael Steel (US) determined that familial Creutzfeldt-Jakob disease (CJD), subtype CJD 178, shares the D178N mutation with FFI although the clinical and pathological findings between the two are quite distinct. This is believed to be an example of phenotypic heterogeneity where both penetrance and expressivity are influenced by the familial genotypes and the environment {Goldfarb, 1992 #5677;Steel, 1993 #5678}.

Stephen J. Sawcer (GB), Gerald M. Yuill (GB), Thomas F. Esmonde (GB), Peter Estibeiro (GB), James W. Ironside (GB), John E. Bell (GB), and Robert G. Will (GB) reported that two farmers who had cattle with bovine spongiform encephalopathy (BSE) or mad cow disease died of Creutzfeldt-Jakob Disease (CJD) {Sawcer, 1993 #8275}.

Herbert Budka (AT), Adriano Aguzzi (CH), Paul Brown (FR), Jean‐Marie Brucher (FR), Orso Bugiani (IT), John Collinge (UK), Heino Diringer (DE), Filippo Gullotta (DE), Matti Haltia (FI), Jean‐Jacques Hauw (FR), James W. Ironside (GB), Hans A. Kretzschmar (DE), Peter L. Lantos (GB), Carlo Masullo (IT), Maurizio Pocchiari (IT), Wolfgang Schlote (DE), Jun Tateishi (JP), and Robert G. Will (GB) discovered that the bovine spongiform encephaopathy prion is the cause of Creutzfeldt-Jakob disease in humans {Budka, 1995 #25714}.

James A. Mastrianni (US), C. Iannicola (US), Richard M. Myers (US), Stephen J. DeArmond (US), and Stanley Benjamin Prusiner US) found a new mutation of the PrP gene in a patient with pathologically confirmed Creutzfeldt-Jakob disease and a negative family history for dementia. The same PrP mutation was identified in another younger member of the pedigree but was not present in more than 200 alleles tested. Such findings suggest that the frequency of inherited prion disease might be higher than ascertained by clinical history alone {Mastrianni, 1996 #22530}.

Stanley Benjamin Prusiner (US) and Michael R. Scott (US) suggested that mutant prions have the capacity to reshape normal prions into versions of themselves by causing a change in their folding pattern {Prusiner, 1997 #5892}.

Peter Chien (US), Jonathan S. Weissman (US), Angela H. DePace (US), Motomasa Tanaka (US), Nariman Naber (US), and Roger Cooke (US) found that the single protein of a prion can misfold into several different conformations each having specific growth dynamics, stabilities, pathologies, and cross-species infectivity {Chien, 2004 #27696;Tanaka, 2004 #27697}.

Ilia V. Baskakov (US), Byron Caughey (US), Jesús R. Requena (ES), Alejandro M. Sevillano (US), Witold K. Surewicz (US), and Holger Wille (CA) concluded that while numerous studies have now proposed various possible PrP structures as the authentic infectious prion assembly none of these have been convincingly correlated with high specific prion infectivity and no international consensus has been reached on their in vivo relevance {Baskakov, 2019 #27692}.


John Joseph Bittner (US), in mice, discovered a naturally transferable cancer agent that is transmitted through the milk of the mother to her offspring {Bittner, 1936 #8577;Bittner, 1939a #8575;Bittner, 1939b #8576}.

Samuel Graff (US), Dan H. Moore (US), Wendell Meredith Stanley (US), Henry T. Randall (US), and Cushman D. Haagensen (US) isolated and characterized this agent as a virus {Graff, 1948 #8578;Graff, 1949 #8579}. Today it is called the Bittner mammary tumor virus.


Albert Jan Kluyver (NL) and Cornelis Bernardus Kees van Niel (NL-US) proposed that bacterial genera be defined both morphologically and biochemically {Kluyver, 1936 #5099}.


Henry Hallett Dale (GB), Wilhelm Siegmund Feldberg (DE-GB), and Marthe Louise Vogt (DE-GB) found that in the presence of eserine (it blocks the action of acetylcholinesterase), curare does not block acetylcholine release from the motor nerve terminals upon electrical stimulation but, of course, blocks the effect of acetylcholine on the muscle membrane {Dale, 1936 #5285}.


Alister Clavering Hardy (GB) and Nora Ennis (GB) reported on the use of their plankton continuous recorder. Their aim was to develop a technique for estimating the numbers or weights of planktonic organisms beneath a unit area of sea surface or in a unit volume of water {Hardy, 1936 #8984}.


Max Walker De Laubenfels (US) wrote an overview of all sponge genera. Presented as a monograph on the sponges of the Florida Keys, he revised almost casually all extant supraspecific taxa, erecting in the process several hundreds of new taxa. These paper taxa, i.e., erected by simply reading the descriptions in the previous literature, were mostly insufficiently established to be of use in sponge classification. Nevertheless, his book presented the first comprehensive overview of the genera and families of sponges, and it formed the basis for modern sponge classification {De Laubenfels, 1936 #14691}. He is commemorated by Delaubenfelsia Dickinson, 1945; Endectyon delaubenfelsi Burton, 1930; Holoplocamia delaubenfelsi Little, 1963; Rhaphidophlus delaubenfelsi Lévi, 1963; and Xestospongia delaubenfelsi Riveros, 1951.


Karl von Frisch (AT) was the first to suggest that the tightly coupled otic gas bladder in the mormyrid fishes can transmit the sound pressure component of the acoustic signal into the inner ear to enhance overall hearing ability {von Frisch, 1936 #6134;von Frisch, 1938 #12394}.


Richard Edwin Shope (US) found evidence that the virus, which caused the 1918 influenza pandemic in humans, and the swine influenza virus are one and the same {Shope, 1936f #10920}.


M. Robert Irwin (US) and Leon J. Cole (US) established that antigens are inherited using the Ringdove, Streptopelia risoria, and the Pearlneck dove, Spilopelia chinensis, as their experimental material. The term immunogenetics was coined in this article {Irwin, 1936 #7301}.


Ronald Aylmer Fisher (GB-AU) produced an article that represents a milestone in numerical taxonomy {Fisher, 1936 #9409}.


Perrin Hamilton Long (US) and Eleanor A. Bliss (US) are credited with having introduced sulfonamides, which were the first effective antibacterial agents, to the United States. They used sulfanilamide in clinical applications at Johns Hopkins University in 1936 {Long, 1937a #17839;Long, 1937b #17841;Long, 1938 #17840}.

Leonard Colebrook (GB), showed that Prontosil was effective against hemolytic Streptococcus in childbirth and hence a cure for puerperal fever {Colebrook, 1936a #14640;Colebrook, 1936b #14641}.

Francis F. Schwentker (US), Sidney Gelman (US), Perrin Hamilton Long (US), and Eleanor A. Bliss (US) were the first to use antimicrobial therapy against meningococcal infections in demonstrating the efficacy of sulfonamides against meningococcus {Long, 1937c #17838;Schwentker, 1937 #11552}.


Walter Thomas James Morgan (GB) and Stanley Miles Partridge (NZ-GB) were the first to describe the somatic antigen (endotoxin) of the Enterobacteriaceae as a toxin {Morgan, 1936 #15916;Morgan, 1940 #15915;Morgan, 1942 #141}.


William Augustus Hinton (US) published the first major text on syphilis entitled Syphilis and Its Treatment {Hinton, 1936 #8816}. Hinton developed the Hinton Test to detect syphilis in blood and spinal fluids, which reduced the number of false positive diagnoses of the disease. John A.V. Davies (US) improved the test to make it applicable to the cerebrospinal fluid. It then became known as the Davies-Hinton Test.


Arthur J. Patek, Jr. (US), Richard P. Stetson (US), and F.H. Laskey Taylor (US) found that patients with hemophilia are lacking a factor present in normal plasma. They called it anti-hemophilic factor (AHF) or anti-hemophilic globulin (AHG) {Patek, 1936 #16073;Patek, 1937 #16074}. This deficiency now called hemophilia A (or factor VIII deficiency) is found almost exclusively in males and is one of the most common of the hereditary coagulation disorders.


Yngve Zotterman (SE) isolated the single nerve units in the taste receptors of the tongue {Diamant, 1963 #15532;Zotterman, 1936 #15533}.


Edward Alfred Cockayne (GB) described a symptom complex (Cockayne’s syndrome) with long list of clinical features, including dwarfism with disproportionately long extremities and large hands and feet, kyphosis, cold blue extremities, beak-like nose, and mental retardation. This is a condition of unknown pathogenesis, which usually presents in the 2nd year of life after normal infancy. Inheritance is autosomal recessive {Cockayne, 1936 #22881;Cockayne, 1946 #22882}.


Harold P. Himsworth (GB) noted that there are two main types of diabetes, the insulin-depleted (type 1) and the insulin-resistant form (type 2). “Insulin resistance” is a term and concept he coined {Himsworth, 1936 #21635}.

Jerry M. Radziuk (CA), Kenneth H. Norwich (CA), Mladen Vranic (HR-CA), Diana T. Finegood (CA), and Richard N. Bergman (CA) collaborated in the first clinical tracer studies on insulin resistance, hypertriglyceridemia, and the Cori cycle {Radziuk, 1978 #24815;Finegood, 1987 #24816}.


Wilder Graves Penfield (US-CA) found that stimulation anywhere on the cerebral cortex could bring responses of one kind or another, but he found that only by stimulating the temporal lobes (the lower parts of the brain on each side) could he elicit meaningful, integrated responses such as memory, including sound, movement, and color. These memories were much more distinct than usual memory and were often about things unremembered under ordinary circumstances. It seemed he had found a physical basis for memory, an engram. Penfield said, “there is hidden away in the brain, a record of the stream of consciousness” {Penfield, 1936-1937 #18234;Penfield, 1937 #17692;Penfield, 1958 #3338;Penfield, 1960 #6600}. However, the reported episodes of recall occurred in less than five percent of his patients, and these results have not been replicated by modern surgeons {Jensen, 2005 #20234}.


János Hugo Bruno “Hans” Selye (AT-HU-CA) proposed the concept of the stress syndrome after a twelve-year study of the physiological effects of stress on animals. It included thymicolymphatic involution, gastric ulcers, lipid discharge from the adrenal, and a loss of chromaffinity in the medulla that he established as a non-specific adaptive response to various kinds of agents He promoted the complex topic of stress as applied to every aspect of daily life or medicine {Selye, 1936 #13494;Selye, 1949 #22580;Selye, 1954 #13495;Selye, 1956 #13496;Selye, 1974 #13497;Selye, 1975 #13498;Selye, 1976 #13499;Selye, 1979 #6718;Selye, 1943b #23546}.

Hans Hugo Bruno Selye (AT-HU-CA) explained his stress model based on physiology and psychobiology as general adaptation syndrome (GAS). His model states that an event that threatens an organism’s well-being (a stressor) leads to a three-stage bodily response: stage 1: alarm, stage 2: resistance, and stage 3: exhaustion {Selye, 1946 #22740}.


Howard Bishop Lewis (US), Barker H. Brown (US), and Florence R. White (US) administered either cystine, cysteine hydrochloride, or dl-methionine, combined with a high or low protein diet to a cystinuric subject and measured the amount of extra cystine and sulfur excreted in the subject’s urine. They found that the administration of cystine did not affect cystine excretion, but it did induce a large increase in the sulfur content of the urine. Cysteine hydrochloride and dl-methionine, on the other hand, led to increases in the excreted amounts of both cystine and sulfate. Surprisingly, less extra cystine was excreted after methionine was administered to the subject when he consumed a high protein diet. These results led them to conclude, “the utilization of the precursor of the urinary cystine in cystinuria occurs more readily under conditions of a high level of protein metabolism." These experiments also confirmed an earlier hypothesis by Erwin Brand (US) who postulated that cysteine is a product of the catabolism of methionine and that the error in cystinuria is a failure of the proper utilization of cysteine and not of cystine. Thus, the extra cystine excreted after the feeding of methionine is derived directly from the degradation of methionine, with cysteine as an intermediary product {Lewis, 1936 #21461}.


J. Arnold Bargen (US), Jesse L. Bollman (US), and Edwin J. Kepler (US) described autonomic diabetic neuropathy causing diarrhea {Bargen, 1936 #22606}. Diabetic autonomic neuropathy (DAN) is a serious and common complication of diabetes.


W. Ritchie Russell (GB) used intraspinal alcohol injections in cases of int