Ernest Henry Starling
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Ernest Henry Starling (17 April 1866 – 2 May 1927) was a British
physiologist Physiology (; ) is the scientific study of functions and mechanisms in a living system. As a sub-discipline of biology, physiology focuses on how organisms, organ systems, individual organs, cells, and biomolecules carry out the chemical a ...
who contributed many fundamental ideas to this subject. These ideas were important parts of the British contribution to physiology, which at that time led the world. He made at least four significant contributions: 1. In the capillary, water is forced out through the pores in the wall by hydrostatic pressure and driven in by the
osmotic pressure Osmotic pressure is the minimum pressure which needs to be applied to a solution to prevent the inward flow of its pure solvent across a semipermeable membrane. It is also defined as the measure of the tendency of a solution to take in a pure ...
of
plasma proteins Blood-proteins, also termed plasma proteins, are proteins present in blood plasma. They serve many different functions, including transport of lipids, hormones, vitamins and minerals in activity and functioning of the immune system. Other blood pr ...
(or
oncotic pressure Oncotic pressure, or colloid osmotic-pressure, is a form of osmotic pressure induced by the proteins, notably albumin, in a blood vessel's plasma (blood/liquid) that causes a pull on fluid back into the capillary. Participating colloids displace ...
). These opposing forces approximately balance; which is known as Starling's Principle. 2. The discovery of the hormone
secretin Secretin is a hormone that regulates water homeostasis throughout the body and influences the environment of the duodenum by regulating secretions in the stomach, pancreas, and liver. It is a peptide hormone produced in the S cells of the duode ...
with his brother-in-law William Baylissand the introduction of the word
hormone A hormone (from the Greek participle , "setting in motion") is a class of signaling molecules in multicellular organisms that are sent to distant organs by complex biological processes to regulate physiology and behavior. Hormones are required ...
. 3. The analysis of the heart's activity as a pump, which is known as the
Frank–Starling law The Frank–Starling law of the heart (also known as Starling's law and the Frank–Starling mechanism) represents the relationship between stroke volume and end diastolic volume.Widmaier, E. P., Hershel, R., & Strang, K. T. (2016).''Vander's Huma ...
. 4. Several fundamental observations on the action of the kidneys. These include evidence for the existence of
vasopressin Human vasopressin, also called antidiuretic hormone (ADH), arginine vasopressin (AVP) or argipressin, is a hormone synthesized from the AVP gene as a peptide prohormone in neurons in the hypothalamus, and is converted to AVP. It then travel ...
, the anti-diuretic hormone. He also wrote the leading textbook of physiology in English, which ran through 20 editions.


Rising to prominence

Ernest Starling became a medical student at
Guy's Hospital Guy's Hospital is an NHS hospital in the borough of Southwark in central London. It is part of Guy's and St Thomas' NHS Foundation Trust and one of the institutions that comprise the King's Health Partners, an academic health science centre. ...
, London, in 1882 (when he was 16). He had a brilliant career there and set his sights on becoming a
Harley Street Harley Street is a street in Marylebone, Central London, which has, since the 19th century housed a large number of private specialists in medicine and surgery. It was named after Edward Harley, 2nd Earl of Oxford and Earl Mortimer.< ...
physician. But the science behind medicinephysiologyattracted him much more; he spent a long vacation in Wilhelm Kühne's laboratory in
Heidelberg Heidelberg (; Palatine German language, Palatine German: ''Heidlberg'') is a city in the States of Germany, German state of Baden-Württemberg, situated on the river Neckar in south-west Germany. As of the 2016 census, its population was 159,914 ...
, studying the mechanisms of
lymph Lymph (from Latin, , meaning "water") is the fluid that flows through the lymphatic system, a system composed of lymph vessels (channels) and intervening lymph nodes whose function, like the venous system, is to return fluid from the tissues to ...
formation and convinced himself that he could become a
physiologist Physiology (; ) is the scientific study of functions and mechanisms in a living system. As a sub-discipline of biology, physiology focuses on how organisms, organ systems, individual organs, cells, and biomolecules carry out the chemical a ...
. At that time such a job description did not exist in Britain. Guy's had no physiological laboratories, but Starling's enthusiasm changed all this, and he published nine papers on
lymph Lymph (from Latin, , meaning "water") is the fluid that flows through the lymphatic system, a system composed of lymph vessels (channels) and intervening lymph nodes whose function, like the venous system, is to return fluid from the tissues to ...
and
capillary A capillary is a small blood vessel from 5 to 10 micrometres (μm) in diameter. Capillaries are composed of only the tunica intima, consisting of a thin wall of simple squamous endothelial cells. They are the smallest blood vessels in the body: ...
function between 1893 and 1897. He showed that there are opposing forces across the capillary wallan outward movement of water due to hydrostatic pressure (derived from the heart's contraction) and an inward movement, secondary to the osmotic pressure of the
plasma proteins Blood-proteins, also termed plasma proteins, are proteins present in blood plasma. They serve many different functions, including transport of lipids, hormones, vitamins and minerals in activity and functioning of the immune system. Other blood pr ...
within the capillary. Without awareness of these forces, the physician cannot begin to understand such conditions as
edema Edema, also spelled oedema, and also known as fluid retention, dropsy, hydropsy and swelling, is the build-up of fluid in the body's Tissue (biology), tissue. Most commonly, the legs or arms are affected. Symptoms may include skin which feels t ...
. The inward and outward forces are often referred to as "Starling forces". They established him as a serious contributor. He was elected a Fellow of the
Royal Society The Royal Society, formally The Royal Society of London for Improving Natural Knowledge, is a learned society and the United Kingdom's national academy of sciences. The society fulfils a number of roles: promoting science and its benefits, re ...
in 1899.


Hormones

Starling enjoyed collaborating with
William Bayliss Sir William Maddock Bayliss (2 May 1860 – 27 August 1924) was an English physiologist. Life He was born in Wednesbury, Staffordshire but shortly thereafter his father, a successful merchant of ornamental ironwork, moved his family to a ...
(1860–1924), who was on the staff of
University College London , mottoeng = Let all come who by merit deserve the most reward , established = , type = Public research university , endowment = £143 million (2020) , budget = ...
(UCL), and together they published on the electrical activity of the heart and on
peristalsis Peristalsis ( , ) is a radially symmetrical contraction and relaxation of muscles that propagate in a wave down a tube, in an anterograde direction. Peristalsis is progression of coordinated contraction of involuntary circular muscles, which ...
. In 1891, when he was 25, Starling married Florence Amelia Wooldridge, the widow of Leonard Charles Wooldridge, who had been his physiology teacher at Guy's and died at the age of 32. She was a great support to Starling as a sounding board, secretary, and manager of his affairs as well as mother of their four children. In 1893 Bayliss married Gertrude, Starling's beautiful sister, so the two were brothers-in-law. When Starling was appointed professor at UCL in 1899, the scientific family was even closer. Bayliss and Starling were in the newspaper's headlines when involved in the
Brown Dog affair The Brown Dog affair was a political controversy about vivisection that raged in Britain from 1903 until 1910. It involved the infiltration of University of London medical lectures by Swedish feminists, battles between medical students and the ...
, a controversy relating to
vivisection Vivisection () is surgery conducted for experimental purposes on a living organism, typically animals with a central nervous system, to view living internal structure. The word is, more broadly, used as a pejorative catch-all term for experiment ...
. Bayliss and Starling investigated pancreatic secretion, which at that time was believed to be entirely under nervous control. They showed that whenever food or acid was put into the
duodenum The duodenum is the first section of the small intestine in most higher vertebrates, including mammals, reptiles, and birds. In fish, the divisions of the small intestine are not as clear, and the terms anterior intestine or proximal intestine m ...
some blood-borne stimulus was released, causing the pancreas to secrete. They called this substance
secretin Secretin is a hormone that regulates water homeostasis throughout the body and influences the environment of the duodenum by regulating secretions in the stomach, pancreas, and liver. It is a peptide hormone produced in the S cells of the duode ...
and Starling proposed that the body produces many secretin-like molecules, and in 1905 proposed that these substances should be called
hormone A hormone (from the Greek participle , "setting in motion") is a class of signaling molecules in multicellular organisms that are sent to distant organs by complex biological processes to regulate physiology and behavior. Hormones are required ...
s. By doing this, he began a whole new biological subject, which became known as
endocrinology Endocrinology (from '' endocrine'' + '' -ology'') is a branch of biology and medicine dealing with the endocrine system, its diseases, and its specific secretions known as hormones. It is also concerned with the integration of developmental event ...
.


Medical education

Starling felt passionately about many subjects, one of which was medical education. He found the lack of science behind medical practice intolerable, and when in 1910 a Royal Commission (The Haldane Commission) was set up to improve medical education, Starling was an enthusiastic contributor. One of the consequences of the commission was the establishment of medical units in London teaching hospitals: clinical practice supported by laboratory research is now taken for granted in every large institution. For this, we must thank Starling's and William Osler's evidence given before the Haldane Commission.


The law of the heart

The physiological discovery most often associated with Starling is the Law of the Heart. Occupying two years of his life (1910–1912) his investigations examined how the heart increased its output in response to more blood entering the organ, which increases the size of the chambers during filling. For this work he made use of the anaesthetized dog, in an experimental arrangement known as the heart-lung preparation. This played an important role in subsequent experiments, when he was not primarily interested in blood flow. Starling was unaware of previous work by a German physiologist,
Otto Frank Otto Heinrich Frank (12 May 1889 – 19 August 1980) was a German businessman who later became a resident of the Netherlands and Switzerland. He was the father of Anne and Margot Frank and husband of Edith Frank, and was the sole member of ...
, using the isolated frog heart. Frank showed that the longer the heart-muscle fibres were stretched the stronger the contraction. He did this with no interest in the circulation as a whole, so his finding have to be taken in parallel with Starling's, so it became the
Frank–Starling law The Frank–Starling law of the heart (also known as Starling's law and the Frank–Starling mechanism) represents the relationship between stroke volume and end diastolic volume.Widmaier, E. P., Hershel, R., & Strang, K. T. (2016).''Vander's Huma ...
.


World War I

During the 1914–1918 war, Starling first was involved in research into poison gases. As a commissioned officer he found the organization of the matter chaotic and on several occasions was very outspoken to his war office superiors. This did his prospects no good at all. Many of his distinguished contemporaries received knighthoods. Starling was awarded a CMG. He resigned from the army in June 1917 and finally was able to undertake war work that utilized his abilities. As chairman of the Royal Society Food (War) Committee, he was instrumental in setting up rationing that provided needed calories and also the nutritional supplements then known. Rationing actually improved nutrition in wartime Britain. Germany had a similar food shortage during the war, but coped with it disastrously.


Post-war

Starling returned to UCL at the end of the war. His wartime experiences had left him with a scathing vision on how the country was run, and in particular the educational system. He was especially outspoken on public (the British designation for private) school education. And particularly the teaching of classics themes: "After nine years, nine-tenths of the boys can read neither Latin nor Greek. They may have acquired a few catchwords or allusions to classical mythology, but they can give no account of the manner in which the Greeks lived, or the part played by Greek philosophy in the evolution of modern ideas, or in the way in which western government has been founded on Roman inventions." During the 1920s he was very busy doing experiments and his reputation attracted distinguished collaborators. Many of these involved the heart-lung preparation. It was used to investigate the control of blood pressure (with G. V. Anrep), the activity of insulin (with F. P. Knowlton), and renal function (with E. B. Verney). In 1923–24 the American embryologist, George Washington Corner worked with Starling in his laboratory. In 1920, Starling was found to have colonic cancer, and the surgeon Arbuthnot Lane removed half of his colon. It seriously limited his exceptional physical activities: he gave up mountaineering, for example.


The Nobel Committee

Starling's relationship with the Nobel Prize is of interest. He was first proposed for the prize in 1913 by
Otto Loewi Otto Loewi (; 3 June 1873 – 25 December 1961) was a German-born pharmacologist and psychobiologist who discovered the role of acetylcholine as an endogenous neurotransmitter. For his discovery he was awarded the Nobel Prize in Physiology or Med ...
(who won a Prize himself in 1936). Starling's subject was hormones, with secretin being prominent. The assessor, J. E. Johansson, decided that Starling should receive the prize, but not yet. No prize was given during World War I. In 1920
August Krogh Schack August Steenberg Krogh (15 November 1874 – 13 September 1949) was a Danish professor at the department of zoophysiology at the University of Copenhagen from 1916 to 1945. He contributed a number of fundamental discoveries within several ...
, a Danish physiologist was rewarded for his work on capillaries (his findings were not actually as significant as Starling's had been, twenty years before: but the significance of 'Starling forces' in the capillary had not become apparent). In 1926, Starling was proposed again, this time for his work on the kidney. On this occasion, Johansson felt that Starling's hormone studies should have been rewarded. But by then the experiments had been done almost a quarter of a century before, and Johansson felt that the prize should be given for recent discoveries. He had forgotten that it was he who had put Starling's work on the back burner in 1913. Subsequent British Laureates (such as
Gowland Hopkins Sir Frederick Gowland Hopkins (20 June 1861 – 16 May 1947) was an English biochemist who was awarded the Nobel Prize in Physiology or Medicine in 1929, with Christiaan Eijkman, for the discovery of vitamins, even though Casimir Funk, a Po ...
and
Charles Sherrington Sir Charles Scott Sherrington (27 November 1857 – 4 March 1952) was an eminent English neurophysiologist. His experimental research established many aspects of contemporary neuroscience, including the concept of the spinal reflex as a system ...
) were given the prize for work they had done twenty or thirty years before. But by this time Johansson was no longer involved with the awarding of the prize.


Death

The exact circumstances of Starling's death are far from clear. He was on a pleasure cruise in the West Indies, but when his ship, the Elders & Fyffes
banana boat Banana Boat is a Polish a cappella sextet, authoring and performing original songs representing the genre of neo-shanties. Being one of the pioneers of the new genre, the group retains its simultaneous focus on contemporary interpretations of ...
''Ariguani'', tied up in Kingston harbour he was found to be dead. He was apparently travelling by himself, and there were no friends or relations at his funeralin pouring rainin
Kingston, Jamaica Kingston is the capital and largest city of Jamaica, located on the southeastern coast of the island. It faces a natural harbour protected by the Palisadoes, a long sand spit which connects the town of Port Royal and the Norman Manley Inter ...
. No autopsy was performed, so the cause of his death which one may presume to be cancer secondary to his colon tumourwas never established. His passing marked the end of an outstanding contributor to medical science. In the words of Henry Dale "All had found him a generous comrade and leader, and it is impossible to think of physiology in the last thirty years without Starling as the central figure of inspiration … his courage was indomitable, his energy and his passion for knowledge flouted all restraint."


Descendants

Two of his great-grandchildren,
Boris Starling Boris Starling (born 1969)Sohn, Amy (1999) "GETTING A HANDLE ON HOT 'MESSIAH' SCRIBE", ''New York Post'', 7 September 1999, p. 22, ("at 30 he's already been on endless European best-seller lists") is a British novelist, screenwriter and newspaper ...
(born 1969) and Belinda Starling (1972–2006) are writers.


References


Further reading

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External links

* {{DEFAULTSORT:Starling, Ernest 1866 births 1927 deaths Scientists from London English physiologists Fellows of the Royal Society Academics of University College London Royal Medal winners Jodrell Professors of Physiology