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Charles Scott Sherrington

From Wikipedia, the free encyclopedia
English footballer, neurophysiologist and Nobel Prize recipient (1857–1952)

Sir Charles Scott Sherrington
43rd President of the Royal Society
In office
1920–1925
Preceded byJ. J. Thomson
Succeeded byErnest Rutherford
Personal details
Born(1857-11-27)27 November 1857
Died4 March 1952(1952-03-04) (aged 94)
Eastbourne, Sussex, England, United Kingdom
CitizenshipBritish
Scientific career
Alma mater
Awards
Fields
Institutions
Academic advisors
Doctoral students

Sir Charles Scott Sherrington (27 November 1857 – 4 March 1952) was a Britishneurophysiologist. His experimental research established many aspects of contemporary neuroscience, including the concept of the spinal reflex as a system involving connected neurons (the "neuron doctrine"), and the ways in which signal transmission between neurons can be potentiated or depotentiated. Sherrington himself coined the word "synapse" to define the connection between two neurons. His bookThe Integrative Action of the Nervous System (1906)[3] is a synthesis of this work, in recognition of which he was awarded theNobel Prize in Physiology or Medicine in 1932 (along withEdgar Adrian).[4][5][6][7]

In addition to his work in physiology, Sherrington did research inhistology,bacteriology, andpathology. He was president of theRoyal Society in the early 1920s.

Biography

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Early years and education

[edit]

Official biographies claim Charles Scott Sherrington was born inIslington, London, England, on 27 November 1857, and that he was the son of James Norton Sherrington, a country doctor, and his wife, Anne Thurtell.[8] However James Norton Sherrington was an ironmonger and artist's colourman in Great Yarmouth, not a doctor, and died in Yarmouth in 1848, nearly 9 years before Charles was born.[9][10]

In the 1861 census, Charles is recorded as Charles Scott (boarder, 4, born India) with Anne Sherrington (widow) as the head and Caleb Rose (visitor, married, surgeon).[11] He was brought up in this household with Caleb recorded as head in 1871,[12] although Anne and Caleb did not marry until after the death of his wife in 1880.[13]

The relationship between Charles and his childhood family is unknown. During the 1860s the whole family moved to Anglesea Road,Ipswich, reputedly because London exacerbated Caleb Rose's tendency to asthma.[14]

Sherrington's origins have been discussed in several published sources: Chris Moss and Susan Hunter, in theJournal of Medical Biography of January 2018, presented an article discussing the potential origins of Charles Sherrington, i.e. whether he was born in India of unknown parents, or was the illegitimate child of Caleb Rose and Anne Sherrington.[15]

Erling Norrby, PhD, inNobel Prizes and Notable Discoveries (2016) observed: "His family origin apparently is not properly given in his official biography. Considering that motherhood is a matter of fact and fatherhood a matter of opinion, it can be noted that his father was not James Norton Sherrington, from whom his family name was derived. Charles was born 9 years after the death of his presumed father. Instead Charles and his two brothers were the illegitimate sons of Caleb Rose, a highly regarded Ipswich surgeon."[16]

InIpswich Town: A History, Susan Gardiner writes: "George and William Sherrington, along with their older brother, Charles, were almost certainly the illegitimate sons of Anne Brookes, née Thurtell and Caleb Rose, a leading surgeon from Ipswich, with whom she was living in College Road, Islington at the time that all three boys were born. No father was named in the baptism register of St James' Church, Clerkenwell, and there is no official record of the registration of any of their births. It was claimed they were the sons of a country doctor, James Norton Sherrington. However, it was with Caleb Rose that Anne and the three Sherrington boys moved to Anglesea Road, Ipswich in 1860 and the couple were married in 1880 after Caleb's first wife had died."[17]

Judith Swazey, inReflexes and Motor Integration: Sherrington's Concept of Integrative Action (1969), quotes Charles Scott Sherrington's son, Carr Sherrington: "James N. Sherrington was always called Mr. and I have no knowledge that he was a Dr. either in law or in medicine... [He] was mainly interested in art and was a personal friend of J. B. Crone and other painters."[18]

Caleb Rose was noteworthy as both a classical scholar and an archaeologist. At the family's Edgehill House in Ipswich one could find a fine selection of paintings, books, and geological specimens.[1][19] Through Rose's interest in theNorwich School of Painters, Sherrington gained a love of art.[20] Intellectuals frequented the house regularly. It was this environment that fostered Sherrington's academic sense of wonder. Even before matriculation, the young Sherrington had read Johannes Müller'sElements of Physiology. The book was given to him by Caleb Rose.

Sherrington enteredIpswich School in 1871.[1]Thomas Ashe, a famous English poet, taught at the school. Ashe served as an inspiration to Sherrington, instilling a love of classics and the desire to travel.

Rose had pushed Sherrington towards medicine. Sherrington first began to study with theRoyal College of Surgeons of England. He also sought to study at Cambridge, but a bank failure had devastated the family's finances. Sherrington elected to enroll atSt Thomas' Hospital in September 1876 as a "perpetual pupil".[1] He did so in order to allow his two younger brothers to do so ahead of him. The two studied law there. Medical studies at St. Thomas's Hospital were intertwined with studies atGonville and Caius College, Cambridge.[19] Physiology was Sherrington's chosen major at Cambridge. There, he studied under the "father of British physiology,"Sir Michael Foster.[21]

Sherrington playedfootball for his grammar school, and forIpswich Town Football Club; he playedrugby for St. Thomas's, was on the rowing team at Oxford.[19][22] During June 1875, Sherrington passed his preliminary examination in general education at theRoyal College of Surgeons of England (RCS). This preliminary exam was required for Fellowship, and also exempted him from a similar exam for the Membership. In April 1878, he passed his Primary Examination for the Membership of the RCS, and twelve months later the Primary for Fellowship.

In October 1879, Sherrington entered Cambridge as anon-collegiate student.[23] The following year he enteredGonville and Caius College. Sherrington was a first-rate student. In June 1881, he took Part I in theNatural Sciences Tripos (NST) and was awarded aStarred first inphysiology; there were nine candidates in all (eight men, one woman), of whom five gainedFirst-class honours (Firsts); in June 1883, in Part II of the NST, he also gained a First, alongsideWilliam Bateson.[24]Walter Holbrook Gaskell, one of Sherrington's tutors, informed him in November 1881 that he had earned the highest marks for his year in botany, human anatomy, and physiology; second in zoology; and highest overall.[19]John Newport Langley was Sherrington's other tutor. The two were interested in how anatomical structure is expressed in physiological function.[21]

Sherrington earned hisMembership of the Royal College of Surgeons on 4 August 1884. In 1885, he obtained a First Class in the Natural Science Tripos with the mark of distinction. In the same year, Sherrington earned the degree ofM.B., Bachelor of Medicine and Surgery from Cambridge. In 1886, Sherrington added the title ofL.R.C.P., Licentiate of the Royal College of Physicians.[1]

Seventh International Medical Congress

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Charles Scott Sherrington

The 7thInternational Medical Congress was held in London in 1881. It was at this conference that Sherrington began his work in neurological research. At the conference controversy broke out.Friedrich Goltz ofStrasbourg argued that localized function in thecortex did not exist. Goltz came to this conclusion after observing dogs who had parts of their brains removed.David Ferrier, who became a hero of Sherrington's, disagreed. Ferrier maintained that there was localization of function in the brain. Ferrier's strongest evidence was a monkey who suffered fromhemiplegia, paralysis affecting one side of the body only, after a cerebral lesion.

A committee, including Langley, was made up to investigate. Both the dog and the monkey were chloroformed. The right hemisphere of the dog was delivered to Cambridge for examination. Sherrington performed a histological examination of the hemisphere, acting as a junior colleague to Langley. In 1884, Langley and Sherrington reported on their findings in a paper. The paper was the first for Sherrington.[1]

Travel

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In the winter of 1884–1885, Sherrington left England for Strasbourg. There, he worked with Goltz. Goltz, like many others, positively influenced Sherrington. Sherrington later said of Goltz that:"[h]e taught one that in all things only the best is good enough."[1]

A case ofasiatic cholera had broken out in Spain in 1885. A Spanish doctor claimed to have produced a vaccine to fight the outbreak. Under the auspices ofCambridge University, theRoyal Society of London, and the Association for Research in Medicine, a group was put together to travel to Spain to investigate.C.S. Roy, J. Graham Brown, and Sherrington formed the group. Roy was Sherrington's friend and the newly elected professor of pathology at Cambridge. As the three travelled toToledo, Sherrington was skeptical of the Spanish doctor.[19] Upon returning, the three presented a report to the Royal Society. The report discredited the Spaniard's claim.

Sherrington didnot meetSantiago Ramón y Cajal on this trip. While Sherrington and his group remained in Toledo, Cajal was hundreds of miles away inZaragoza.[19]

Later that year Sherrington travelled toRudolf Virchow in Berlin to inspect the cholera specimens he procured in Spain. Virchow later on sent Sherrington toRobert Koch for a six weeks' course in technique. Sherrington ended up staying with Koch for a year to do research in bacteriology. Under these two, Sherrington parted with a good foundation in physiology,morphology,histology, andpathology.[21] During this period he may have also studied withWaldeyer andZuntz.

In 1886, Sherrington went to Italy to again investigate a cholera outbreak. While in Italy, Sherrington spent much time in art galleries. It was in this country that Sherrington's love for rare books became an obsession.[19]

Employment

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C.S. Roy and Charles Scott Sherrington (right), at the door of the Old Pathological Laboratory, Cambridge, 1893

In 1891, Sherrington was appointed as superintendent of the Brown Institute for Advanced Physiological and Pathological Research of the University of London, a center for human and animal physiological and pathological research.[19][20] Sherrington succeededSir Victor Alexander Haden Horsley.[25] There, Sherrington worked on segmental distribution of the spinal dorsal and ventral roots, he mapped the sensory dermatomes, and in 1892 discovered that muscle spindles initiated thestretch reflex. The institute allowed Sherrington to study many animals, both small and large. The Brown Institute had enough space to work with large primates such as apes.

Liverpool

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Plaque reads "The Physiological Society. Sir Charles Sherrington (1857-1952). OM PRS. Physiologist and Nobel Laureate. George Holt Professor of Physiology, University of Liverpool from 1895-1913. Shared the Nobel Prize in Physiology or Medicine with Edgar Adrian in 1932 for their discoveries regarding the functions of neurons. Sherrington laid the foundation for our understanding of the central nervous system."
Plaque dedicated to Sherrington at the University of Liverpool

Sherrington's first job of full-professorship came with his appointment as Holt Professor of Physiology atLiverpool in 1895, succeeding Francis Gotch.[19] With his appointment to the Holt Chair, Sherrington ended his active work in pathology.[1] Working on cats, dogs, monkeys, and apes that had been bereaved of their cerebral hemispheres, he found that reflexes must be considered integrated activities of the total organism, not just the result of activities of the so-called reflex-arcs, a concept then generally accepted.[25] There he continued his work on reflexes andreciprocal innervation. His papers on the subject were synthesized into theCroonian lecture of 1897.

Sherrington showed that muscle excitation was inversely proportional to the inhibition of an opposing group of muscles. Speaking of the excitation-inhibition relationship, Sherrington said "desistence from action may be as truly active as is the taking of action." Sherrington continued his work on reciprocal innervation during his years at Liverpool. Come 1913, Sherrington was able to say that "the process of excitation and inhibition may be viewed as polar opposites [...] the one is able to neutralize the other." Sherrington's work on reciprocal innervation was a notable contribution to the knowledge of the spinal cord.[1]

Oxford

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As early as 1895, Sherrington had tried to gain employment at Oxford University. By 1913, the wait was over. Oxford offered Sherrington theWaynflete Chair of Physiology atMagdalen College.[1] The electors to that chair unanimously recommended Sherrington without considering any other candidates.[19] Sherrington enjoyed the honor of teaching many bright students at Oxford, includingWilder Penfield, who he introduced to the study of the brain. Several of his students were Rhodes scholars, three of whom –Sir John Eccles,Ragnar Granit, andHoward Florey – went on to be Nobel laureates.[26] Sherrington also influenced American pioneer brain surgeonHarvey Williams Cushing.

Sherrington's philosophy as a teacher can be seen in his response to the question of what was the real function of Oxford University in the world. Sherrington said:

"After some hundreds of years of experience we think that we have learned here in Oxford how to teach what is known. But now with the undeniable upsurge of scientific research, we cannot continue to rely on the mere fact that we have learned how to teach what is known. We must learn to teach the best attitude to what is not yet known. This also may take centuries to acquire but we cannot escape this new challenge, nor do we want to."[19]

Box of microscope slides carrying the plaque: "Sir Charles Sherrington's Histology Demonstration Slides: St Thomas's Hospital: 1886–1895; Liverpool University: 1895–1915; Oxford University: 1914–1935"

While at Oxford, Sherrington kept hundreds of microscope slides in a specially constructed box labelled "Sir Charles Sherrington's Histology Demonstration Slides". As well as histology demonstration slides, the box contains slides which may be related to original breakthroughs such as cortical localization in the brain; slides from contemporaries such asAngelo Ruffini andGustav Fritsch; and slides from colleagues at Oxford such asJohn Burdon-Sanderson – the firstWaynflete Chair of Physiology – andDerek Denny-Brown, who worked with Sherrington at Oxford (1924–1928)).[27]

Sherrington's teachings at Oxford were interrupted byWorld War I. When the war started, it left his classes with only nine students. During the war, he laboured at a shell factory to support the war and to study fatigue in general, but specifically industrial fatigue. His weekday work hours were from 7:30am to 8:30pm; and from 7:30am to 6:00pm on the weekends.[19]

In March 1916, Sherrington fought for women to be admitted to the medical school at Oxford.

He lived at 9Chadlington Road in north Oxford from 1916 to 1934, and on 28 April 2022 an Oxfordshire blue plaque in his honour was unveiled on this house.[28]

Retirement

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Charles Sherrington retired from Oxford in the year of 1936.[1] He then moved to his boyhood town of Ipswich, where he built a house.[21] There, he kept up a large correspondence with pupils and others from around the world. He also continued to work on his poetic, historical, and philosophical interests.[26] From 1944 until his death he was President of theIpswich Museum, on the committee he had previously served.[29]

Sherrington's mental faculties were crystal clear up to the time of his sudden death, which was caused by a sudden heart failure at age 94. His bodily health, however, did suffer in old age. Arthritis was a major burden.[21] Speaking of his condition, Sherrington said "old age isn't pleasant[,] one can't do things for oneself."[1] The arthritis put Sherrington in a nursing home in the year before his death, in 1951.[26]

Family

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On 27 August 1891, Sherrington married Ethel Mary Wright (died 1933), daughter of John Ely Wright of Preston Manor, Suffolk, England. They had one child, a son named Charles ("Carr") E.R. Sherrington, who was born in 1897.[21]

On weekends during the Oxford years the couple would frequently host a large group of friends and acquaintances at their house for an enjoyable afternoon.[1]

Publications

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The Integrative Action of the Nervous System

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Published in 1906,[3] this was a compendium of ten of Sherrington'sSilliman lectures, delivered atYale University in 1904.[30]

The book discussed neuron theory, the "synapse" (a term he had introduced in 1897, the word itself suggested byclassicistA. W. Verrall[31]), communication between neurons, and a mechanism for the reflex-arc function.[6] The work effectively resolved the debate between neuron andreticular theory in mammals, thereby shaping our understanding of the central nervous system.[30]

He theorized that the nervous system coordinates various parts of the body and that the reflexes are the simplest expressions of the interactive action of the nervous system, enabling the entire body to function toward a definite purpose. Sherrington pointed out that reflexes must be goal-directive and purposive. Furthermore, he established the nature of postural reflexes and their dependence on the anti-gravitystretch reflex and traced the afferent stimulus to theproprioceptive end organs, which he had previously shown to be sensory in nature ("proprioceptive" was another term he had coined[6]). The work was dedicated to Ferrier.[21]

Mammalian Physiology: a Course of Practical Exercises

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The textbook was published in 1919 at the first possible moment after Sherrington's arrival at Oxford and the end of the War.[1]

The Assaying of Brabantius and other Verse

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This collection of previously published war-time poems was Sherrington's first major poetic release, published in 1925. Sherrington's poetic side was inspired byJohann Wolfgang von Goethe. Sherrington was fond of Goethe the poet, but not Goethe the scientist. Speaking of Goethe's scientific writings, Sherrington said "to appraise them is not a congenial task."[1]

Man on His Nature

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A reflection on Sherrington's philosophical thought. Sherrington had long studied the 16th century French physicianJean Fernel, and grew so familiar with him that he considered him a friend. During the academic year 1937-38, Sherrington delivered theGifford lectures at the University of Edinburgh. They focused on Fernel and his times, and formed the basis ofMan on His Nature. The book was published in 1940, with a revised edition in 1951. It explores philosophical thoughts about the mind, human existence, and God, in accordance withnatural theology.[32] In the original edition, each of the twelve chapters begins with one of the twelvezodiac signs; Sherrington discusses astrology in Fernel's time in Chapter 2.[33] In his ideas on mind and cognition, Sherrington introduced the idea that neurons work as groups in a "million-fold democracy" to produce outcomes rather than with central control.[34]

Honours and awards

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Stained glass window in the dining hall of Gonville and Caius College, in Cambridge (UK), commemorating Charles Scott Sherrington

Sherrington was elected aFellow of the Royal Society (FRS) in 1893.[1]

At the time of his death Sherrington receivedhonoris causa Doctors from twenty-two universities: Oxford,Paris,Manchester,Strasbourg,Louvain,Uppsala,Lyon,Budapest,Athens,London,Toronto,Harvard,Dublin,Edinburgh,Montreal,Liverpool,Brussels,Sheffield,Bern,Birmingham,Glasgow, and theUniversity of Wales.[1]

Eponyms

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Liddell-Sherrington reflex
Associated with Edward George Tandy Liddell and Charles Scott Sherrington, the Liddell-Sherrington reflex is the tonic contraction of muscle in response to its being stretched. When a muscle lengthens beyond a certain point, the myotatic reflex causes it to tighten and attempt to shorten. This can be felt as tension during stretching exercises.
Schiff-Sherrington reflex
Associated with Moritz Schiff and Charles Scott Sherrington, describes a grave sign in animals: rigid extension of the forelimbs after damage to the spine. It may be accompanied by paradoxical respiration – the intercostal muscles are paralysed and the chest is drawn passively in and out by the diaphragm.
Sherrington's First Law
Every posterior spinal nerve root supplies a particular area of the skin, with a certain overlap of adjacent dermatomes.
Sherrington's Second Law
The law of reciprocal innervation. When contraction of a muscle is stimulated, there is a simultaneous inhibition of its antagonist. It is essential for coordinated movement.
Vulpian-Heidenhain-Sherrington phenomenon
Associated withRudolf Peter Heinrich Heidenhain,Edmé Félix Alfred Vulpian, and Charles Scott Sherrington. Describes the slow contraction of denervated skeletal muscle by stimulating autonomic cholinergic fibres innervating its blood vessels.

See also

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References

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  1. ^abcdefghijklmnopqLiddell, E. G. T. (1952). "Charles Scott Sherrington. 1857-1952".Obituary Notices of Fellows of the Royal Society.8 (21):241–270.doi:10.1098/rsbm.1952.0016.JSTOR 768811.S2CID 35977575.
  2. ^Todman, Donald (2008). "Howard Florey and research on the cerebral circulation".Journal of Clinical Neuroscience.15 (6):613–616.doi:10.1016/j.jocn.2007.04.017.PMID 18280740.S2CID 40353145.His mentor was the neurophysiologist and Nobel Laureate, Sir Charles Sherrington who directed him in neuroscience research. Florey's initial studies on the cerebral circulation represent an original contribution to medical knowledge and highlight his remarkable scientific method. The mentorship and close personal relationship with Sherrington was a crucial factor in Florey's early research career.
  3. ^abSherrington, Charles Scott (1906).The integrative action of the nervous system (1st ed.). Oxford University Press: H. Milford. pp. xvi, 411 p., [19] leaves of plates.
  4. ^""Sir Charles Sherrington – Nobel Lecture: Inhibition as a Coordinative Factor"". Retrieved31 July 2012.
  5. ^"Sir Charles Scott Sherrington". Encyclopædia Britannica, Inc. Retrieved31 July 2012.
  6. ^abcPearce, J. M. (2004)."Sir Charles Scott Sherrington (1857–1952) and the synapse".Journal of Neurology, Neurosurgery, and Psychiatry.75 (4): 544.doi:10.1136/jnnp.2003.017921.PMC 1739021.PMID 15026492.
  7. ^Penfield, W. (1962). "Sir Charles Sherrington, O.M., F.R.S. (1857–1952): An Appreciation".Notes and Records of the Royal Society.17 (2):163–168.doi:10.1098/rsnr.1962.0015.S2CID 145536063.
  8. ^Charles Scott Sherrington on Nobelprize.orgEdit this at Wikidata, accessed 11 October 2020
  9. ^GRO index: 1848 Dec, Yarmouth 13, 258
  10. ^Will of James Norton Sherrington, proved at London 5 March 1849, National Archives Catalogue Reference:Prob 11/2090, image 171
  11. ^"1861 England, Wales & Scotland Census Transcription". Findmypast. 1861. Retrieved26 November 2016.
  12. ^"1871 England, Wales & Scotland Census Transcription". Findmypast. 1871. Retrieved26 November 2016.
  13. ^GRO marriages index: 1880 Dec, Ipswich 4a, 1377
  14. ^Anon (1895). "Obituary: Mr. Caleb Rose".BMJ.2 (1820): 1266.doi:10.1136/bmj.2.1820.1266-a.S2CID 220183796.
  15. ^Moss, Chris; Hunter, Susan (2020)."Charles Scott Sherrington's origins".Journal of Medical Biography.28 (1):3–7.doi:10.1177/0967772017746271.PMID 29998781.S2CID 51622906.
  16. ^Nobel Prizes and Notable Discoveries, Erling Norrby, The Royal Swedish Academy of Sciences, Sweden, World Scientific Publishing, 2016, p. 24
  17. ^Ipswich Town: A History, Susan Gardiner, Amberley Publishing, 2009, Chapter 2: Origins.
  18. ^Reflexes and Motor Integration: Sherrington's Concept of Integrative Action, Judith P. Swazey, Harvard University Press, 1969, p. 211
  19. ^abcdefghijklmEccles, J.; Gibson, W. (1979).Sherrington: His Life and Thought. Berlin; New York: Springer International. pp. 1–6, 15, 24–25.ISBN 978-0-387-09063-4.
  20. ^abKarl Grandin, ed. (1932)."Sir Charles Sherrington Biography".Les Prix Nobel. The Nobel Foundation. Retrieved23 July 2008.
  21. ^abcdefgKusurkar, R. A. (2004). "Sir Charles Sherrington (1857–1952)".Journal of Postgraduate Medicine.50 (3):238–239.PMID 15377819.
  22. ^Granit, R. (1967).Charles Scott Sherrington: An Appraisal. Garden City, NY:Doubleday. p. 3.OCLC 573353.
  23. ^"Sherrington, Charles Scott (SHRN879CS)".A Cambridge Alumni Database. University of Cambridge.
  24. ^University of Cambridge Calendar, 1894-95, p. 330
  25. ^ab"Sir Charles Scott Sherrington".Who Named It?. 2008. Archived fromthe original on 7 October 2009. Retrieved23 July 2008.
  26. ^abcGibson, W.C. (2001)."Chapter 1: Sir Charles Sherrington, O.M., P.R.S. (1857–1952)"(PDF).Twentieth Century Neurology: The British Contribution. London: Imperial College Press. pp. 4–6.ISBN 978-1-86094-245-7. Archived fromthe original(PDF) on 26 March 2009. Retrieved27 April 2008.
  27. ^Molnar, Zoltan; Brown, Richard (June 2010). "Insights into the life and work of Sir Charles Sherrington".Nature Reviews Neuroscience.11 (6):429–436.doi:10.1038/nrn2835.PMID 20445541.S2CID 5930858.
  28. ^"Sir Charles Sherrington".
  29. ^Ipswich Museum Records.
  30. ^abBurke, RE (April 2007)."Sir Charles Sherrington's the integrative action of the nervous system: a centenary appreciation".Brain.130 (Pt 4):887–94.doi:10.1093/brain/awm022.PMID 17438014.
  31. ^Cowan, W. Maxwell; Südhof, Thomas C.; Stevens, Charles F. (2003).Synapses. JHU Press. p. 11.ISBN 9780801871184. Retrieved9 June 2020.
  32. ^Charles Scott Sherrington R. Scott Spurlock, University of Edinburgh
  33. ^Sherrington,Man on his Nature (Cambridge, 1940), pp. 51–61
  34. ^Quiroga, Rodrigo Quian (2013)."Gnostic cells in the 21st century".Acta Neurobiol. Exp.73 (4):1–9.doi:10.55782/ane-2013-1952.PMID 24457638.
  35. ^"No. 32563".The London Gazette (Supplement). 31 December 1921. p. 10716.
  36. ^Sherrington's Presidential Address to the British Association Meeting, held at Hull in 1922

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