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 subdiscipline of biology, physiology focuses on how organisms, organ systems, individual organs, cells, and biomolecules carry out chemical and ...

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 Hydrostatics is the branch of fluid mechanics that studies fluids at hydrostatic equilibrium and "the pressure in a fluid or exerted by a fluid on an immersed body". The word "hydrostatics" is sometimes used to refer specifically to water and o ...

and driven in by the

osmotic pressure Osmotic pressure is the minimum pressure which needs to be applied to a Solution (chemistry), 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 soluti ...

plasma proteins Plasma proteins, sometimes referred to as blood proteins, are proteins present in blood plasma. They perform many different functions, including transport of hormones, vitamins and minerals in activity and functioning of the immune system. Other b ...

(or

oncotic pressure Oncotic pressure, or colloid osmotic-pressure, is a type of osmotic pressure induced by the plasma proteins, notably albumin, in a blood vessel's plasma (or any other body fluid such as blood and lymph) that causes a pull on fluid back into ...

). 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 duodenum ...

with his brother-in-law

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 ...

and the introduction of the word

hormone A hormone (from the Ancient Greek, Greek participle , "setting in motion") is a class of cell signaling, signaling molecules in multicellular organisms that are sent to distant organs or tissues by complex biological processes to regulate physio ...

. 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 ...

. 4. Several fundamental observations on the action of the kidneys. These include evidence for the existence of

vasopressin Mammalian 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 ...

, 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 founded by philanthropist Thomas Guy in 1721, located 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 Kin ...

, 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, named after Edward Harley, 2nd Earl of Oxford and Earl Mortimer.Wilhelm Kühne's laboratory in

Heidelberg Heidelberg (; ; ) is the List of cities in Baden-Württemberg by population, fifth-largest city in the States of Germany, German state of Baden-Württemberg, and with a population of about 163,000, of which roughly a quarter consists of studen ...

, studying the mechanisms of

lymph Lymph () 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 be recirculated. At the ori ...

formation and convinced himself that he could become 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

and

capillary A capillary is a small blood vessel, from 5 to 10 micrometres in diameter, and is part of the microcirculation system. Capillaries are microvessels and the smallest blood vessels in the body. They are composed of only the tunica intima (the inn ...

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

within the capillary. Without awareness of these forces, the physician cannot begin to understand such conditions as

edema Edema (American English), also spelled oedema (British English), and also known as fluid retention, swelling, dropsy and hydropsy, is the build-up of fluid in the body's tissue (biology), tissue. Most commonly, the legs or arms are affected. S ...

. 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

(1860–1924), who was on the staff of

University College London University College London (Trade name, branded as UCL) is a Public university, public research university in London, England. It is a Member institutions of the University of London, member institution of the Federal university, federal Uni ...

(UCL), and together they published on the electrical activity of the heart and on

peristalsis Peristalsis ( , ) is a type of intestinal motility, characterized by symmetry in biology#Radial symmetry, radially symmetrical contraction and relaxation of muscles that propagate in a wave down a tube, in an wikt:anterograde, anterograde dir ...

. 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 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 Animal test ...

. 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 vertebrates, including mammals, reptiles, and birds. In mammals, it may be the principal site for iron absorption. The duodenum precedes the jejunum and ileum and is the shortest p ...

some blood-borne stimulus was released, causing the pancreas to secrete. They called this substance

and Starling proposed that the body produces many secretin-like molecules, and in 1905 proposed that these substances should be called

s. By doing this, he began a whole new biological subject, which became known as

endocrinology Endocrinology (from ''endocrine system, endocrine'' + ''wikt:-logy#Suffix, -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 ...

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, and the father of Anne Frank. He edited and published the first edition of her diary in 1947 (subsequently known in English as ''The Diary of a Young Girl'') and adv ...

, using the isolated

frog A frog is any member of a diverse and largely semiaquatic group of short-bodied, tailless amphibian vertebrates composing the order (biology), order Anura (coming from the Ancient Greek , literally 'without tail'). Frog species with rough ski ...

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 findings have to be taken in parallel with Starling's, thus it became the

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 George Washington Corner FRS FRSE (12 December 1889 – 28 September 1981) was an American physician, embryologist and pioneer of the contraceptive pill. He received an outstanding ten honorary degrees from various universities. He played a criti ...

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 Germany, German-born pharmacology, pharmacologist and psychobiologist who discovered the role of acetylcholine as an endogenous neurotransmitter. For this discovery, he was awarded the Nobel ...

(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 seve ...

, 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 In humans, the kidneys are two reddish-brown bean-shaped blood-filtering organ (anatomy), organs that are a multilobar, multipapillary form of mammalian kidneys, usually without signs of external lobulation. They are located on the left and rig ...

. 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. He also discovered the amino ...

and

Charles Sherrington Sir Charles Scott Sherrington (27 November 1857 – 4 March 1952) was a British neurophysiologist. His experimental research established many aspects of contemporary neuroscience, including the concept of the spinal reflex as a system involving ...

) 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 The West Indies is an island subregion of the Americas, surrounded by the Atlantic Ocean, North Atlantic Ocean and the Caribbean Sea, which comprises 13 independent island country, island countries and 19 dependent territory, dependencies in thr ...

, 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 Kingston may refer to: Places * List of places called Kingston, including the six most populated: ** Kingston, Jamaica ** Kingston upon Hull, England ** City of Kingston, Victoria, Australia ** Kingston, Ontario, Canada ** Kingston upon Thames, ...

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 (political), 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 spit (landform), sand spit which connects the town of Por ...

. No

autopsy An autopsy (also referred to as post-mortem examination, obduction, necropsy, or autopsia cadaverum) is a surgical procedure that consists of a thorough examination of a corpse by dissection to determine the cause, mode, and manner of deat ...

was performed, so the cause of his death which one may presume to be

cancer Cancer is a group of diseases involving Cell growth#Disorders, abnormal cell growth with the potential to Invasion (cancer), invade or Metastasis, spread to other parts of the body. These contrast with benign tumors, which do not spread. Po ...

secondary to his colon tumourwas never established. His death 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. His daughter Muriel (born c1893), married

Sydney Patterson Sydney Wentworth Patterson MB BS, MD, DSc, FRCP (born 1882 in Melbourne, Australia, died 1960 in London, England) was a physician, medical researcher and first director of the Walter and Eliza Hall Institute of Medical Research in Melbourne, A ...

at St. Mary's Kilburn, London in 1919.

References

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 Artists' Rifles soldiers Companions_of_the_Order_of_St_Michael_and_St_George