Charles Scott Sherrington

Sir Charles Scott Sherrington OM PRS FRCP FRCS[1][10] (27 November 1857 – 4 March 1952) was an English neurophysiologist, histologist, bacteriologist, and a pathologist, Nobel laureate and president of the Royal Society in the early 1920s. He received the Nobel Prize in Physiology or Medicine with Edgar Adrian, 1st Baron Adrian, in 1932 for their work on the functions of neurons.[11][12] Prior to the work of Sherrington and Adrian, it was widely accepted that reflexes occurred as isolated activity within a reflex arc. Sherrington received the prize for showing that reflexes require integrated activation and demonstrated reciprocal innervation of muscles (Sherrington's law).[13][14] Through his seminal 1906 publication, The Integrative Action of the Nervous System,[15] he had effectively laid to rest the theory that the nervous system, including the brain, can be understood as a single interlinking network. His alternative explanation of synaptic communication between neurons helped shape our understanding of the central nervous system.

Sir Charles Scott Sherrington

Charles Scott Sherrington2
Born27 November 1857
Died4 March 1952 (aged 94)
Eastbourne, Sussex, England, United Kingdom
CitizenshipBritish
Alma mater
Awards
Scientific career
Fields
Academic advisors
Doctoral students
Influences
Influenced

Biography

Early years and education

Official biographies claim Charles Scott Sherrington was born in Islington, London, England, on 27 November 1857 and that he was the son of James Norton Sherrington, a country doctor, and his wife Anne Thurtell.[16] 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.[17][18] 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).[19] He was brought up in this household with Caleb recorded as head in 1871,[20] although Ann and Caleb did not marry until after the death of his wife in 1880.[21] 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.[22]

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][23] Through Rose's interest in the Norwich School of Painters, Sherrington gained a love of art.[24] 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's Elements of Physiology. The book was given to him by Caleb Rose.

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

Rose had pushed Sherrington towards medicine. Sherrington first began to study with the Royal 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 at St 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 at Gonville and Caius College, Cambridge.[23] Physiology was Sherrington's chosen major at Cambridge. There, he studied under the "father of British physiology," Sir Michael Foster.[25]

Sherrington played football for his grammar school, and for Ipswich Town Football Club, rugby St. Thomas's, was on the rowing team at Oxford.[23][26] During June 1875, Sherrington passed his preliminary examination in general education at the Royal College. 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 Royal College of Surgeons, and 12 months later the Primary for Fellowship.

In October 1879, Sherrington entered Cambridge as a non-collegiate student.[27] The following year he entered Gonville and Caius College. Sherrington was quite the student. In June 1881 he took Part I in the Natural Sciences Tripos and was awarded a starred First in physiology; there were 9 candidates in all (8 men, 1 woman), of whom five gained Firsts; in June 1883 in Part II of the Tripos he also gained a First, alongside William Bateson.[28] 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.[23] John Newport Langley was Sherrington's other tutor. The two were interested in how anatomical structure is expressed in physiological function.[25]

Sherrington earned his Membership 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 of M.B., Bachelor of Medicine and Surgery from Cambridge. In 1886, Sherrington added the title of L.R.C.P., Licentiate of the Royal College of Physicians.[1]

Seventh International Medical Congress

Prof. Charles Scott Sherrington
Charles Scott Sherrington

The conference 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 of Strasbourg argued that localized function in the cortex 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 from hemiplegia, 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

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 of asiatic cholera had broken out in Spain in 1885. A Spanish physician claimed to have produced a vaccine to fight the outbreak. Under the auspices of Cambridge University, the Royal 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 traveled to Toledo, Sherrington was skeptical of the Spanish physician.[23] Upon returning, the three presented a report to the Royal Society. The report discredited the Spaniard's claim.

It should be mentioned that Sherrington did not meet Santiago Ramón y Cajal on this trip. While Sherrington and his group remained in Toledo, Cajal was hundreds of miles away in Zaragoza.[23]

Later that year Sherrington traveled to Rudolf Virchow in Berlin to inspect the cholera specimens he procured in Spain. Virchow later on sent Sherrington to Robert 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, and pathology.[25] During this period he may have also studied with Waldeyer and Zuntz.

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 addiction.[23]

Employment

Roy Sherrington 1893 01
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.[23][24] Sherrington succeeded Sir Victor Alexander Haden Horsley.[29] 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 the stretch 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

Sherrington's first job of full-professorship came with his appointment as Holt Professor of Physiology at Liverpool in 1895, succeeding Francis Gotch.[23] 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.[29] There he continued his work on reflexes and reciprocal innervation. His papers on the subject were synthesized into the Croonian 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

As early as 1895, Sherrington had tried to gain employment at Oxford University. By 1913, the wait was over. Oxford offered Sherrington the Waynflete Chair of Physiology.[1] The electors to that chair unanimously recommended Sherrington without considering any other candidates.[23] Sherrington enjoyed the honor of teaching many bright students at Oxford, including Wilder 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, and Howard Florey - went on to be Nobel laureates.[30] Sherrington also influenced American pioneer brain surgeon Harvey 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."[23]

Sir Charles Sherrington's histology demonstration slides box
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 as Angelo Ruffini and Gustav Fritsch; and slides from colleagues at Oxford such as John Burdon-Sanderson – the first Waynflete Chair of Physiology – and Derek Denny-Brown, who worked with Sherrington at Oxford (1924–1928)).[31]

Sherrington's teachings at Oxford were interrupted by World 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.[23]

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

Retirement

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.[25] 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.[30] From 1944 until his own death he was President of the Ipswich Museum, on the committee of which he had previously served.[32]

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 of his.[25] 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.[30]

Family

On 27 August 1891, Sherrington married Ethel Mary Wright (d.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.[25] His wife was both loyal and lively. She was a great host. 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]

Noted publications

The Integrative Action of the Nervous System
Published in 1906,[15] this was a compendium of ten of Sherrington's Silliman lectures, delivered at Yale University in 1904.[33] In the publication, he discussed neuron theory, the "synapse" (a term he had coined in 1897) and communication between neurons, and a mechanism for the reflex-arc function.[11] The work effectively resolved the debate between neuron and reticular theory in mammals, thereby shaping our understanding of the central nervous system.[33] He expressed his theory that the nervous system acts as the coordinator of 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 one definite end at a time. He also pointed out that reflexes have to be goal-directive and purposive. Furthermore, he established the nature of postural reflexes and their dependence on the anti-gravity stretch reflex and traced the afferent stimulus to the proprioceptive end organs, which he had previously shown to be sensory in nature ("proprioceptive" was another term he had coined[11]). The work was dedicated to Ferrier.[25]
Man on His Nature
A reflection of Sherrington's philosophical thought. Sherrington had long studied the 16th century French physician Jean Fernel, and grew so familiar with him that he considered him a friend. In the years of 1937 and 1938, Sherrington delivered the Gifford lectures at the University of Edinburgh; these focused on Fernel and his times, and came to form the principal content of Man on His Nature. The book was released in 1940, and a revised edition came out in 1951. It explores philosophical thoughts about the mind, the human existence, and God, in connection with natural theology.[34] Chapters of the book align with the twelve zodiac signs.[35] In his ideas on the 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.[36]
The Assaying of Brabantius and other Verse
A collection of previously published war-time poems. This was Sherrington's first major poetic release. The Assaying was published in 1925. Sherrington's poetic side was inspired by Johann 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]
Mammalian Physiology: a Course of Practical Exercises
The textbook was released in 1919 at the first possible moment after Sherrington's coming to Oxford and the end of the War.[1]

Honours and awards

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

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

At the time of his death Sherrington received honoris 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 the University of Wales.[1]

Eponyms

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 is the tension you feel 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 with Rudolf 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.

References

  1. ^ a b c d e f g h i j k l m n o p q r Liddell, 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. PMC 1392451. PMID 13000699.
  2. ^ Neurotree profile: Charles Scott Sherrington
  3. ^ Eccles, J. C.; Gibson, W. C. (1979). Sherrington: His Life and Thought. Springer Science+Business Media. ISBN 9783642618642. His library was housed mainly in one large room with open shelves reaching to the ceiling and a couple of turntable bookcases, one of them completely filled with editions of his favourite among all books, Sir Thomas Browne's Religio Medici
  4. ^ Eccles, J. (1968). "Two Hitherto Unrecognized Publications by Sir Charles Sherrington, O.M., F.R.S". Notes and Records of the Royal Society. 23: 86–100. doi:10.1098/rsnr.1968.0012.
  5. ^ Eccles, J. C. (1957). "Some Aspects of Sherrington's Contribution to Neurophysiology". Notes and Records of the Royal Society. 12 (2): 216–225. doi:10.1098/rsnr.1957.0012.
  6. ^ 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. 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.
  7. ^ Tansey, E. M. (2008). "Working with C. S. Sherrington, 1918-24". Notes and Records of the Royal Society. 62 (1): 123–130. doi:10.1098/rsnr.2007.0037. PMC 2628577. PMID 18548907.
  8. ^ Hill, A. V. (1975). "Jewels in My Acquaintance with C. S. Sherrington, F.R.S". Notes and Records of the Royal Society. 30 (1): 65–68. doi:10.1098/rsnr.1975.0006. PMID 11615581.
  9. ^ Todman, D. (2008). "Wilder Penfield (1891–1976)". Journal of Neurology. 255 (7): 1104–1105. doi:10.1007/s00415-008-0915-6. PMID 18500490.
  10. ^ Sherrington, C. E. (1975). "Charles Scott Sherrington (1857-1952)". Notes and Records of the Royal Society of London. 30 (1): 45–63. doi:10.1098/rsnr.1975.0005. PMC 1392451. PMID 11615580.
  11. ^ a b c Pearce, 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 (inactive 2018-08-22). PMC 1739021. PMID 15026492.
  12. ^ 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.
  13. ^ ""Sir Charles Sherrington – Nobel Lecture: Inhibition as a Coordinative Factor"". Retrieved 31 July 2012.
  14. ^ "Sir Charles Scott Sherrington". Encyclopædia Britannica, Inc. Retrieved 31 July 2012.
  15. ^ a b Sherrington, 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.
  16. ^ "Sir Charles Sherrington - Biographical". Nobel Prize official Website. 1932. Retrieved Nov 26, 2016.
  17. ^ GRO index: 1848 Dec, Yarmouth 13, 258
  18. ^ Will of James Norton Sherrington, proved at London 5 March 1849, National Archives Catalogue Reference:Prob 11/2090, image 171
  19. ^ "1861 England, Wales & Scotland Census Transcription". Findmypast. 1861. Retrieved Nov 26, 2016. (Subscription required (help)).
  20. ^ "1871 England, Wales & Scotland Census Transcription". Findmypast. 1871. Retrieved Nov 26, 2016. (Subscription required (help)).
  21. ^ GRO marriages index: 1880 Dec, Ipswich 4a, 1377
  22. ^ Anon (1895). "Obituary: Mr. Caleb Rose". BMJ. 2 (1820): 1266. doi:10.1136/bmj.2.1820.1266-a.
  23. ^ a b c d e f g h i j k l m Eccles, 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.
  24. ^ a b Karl Grandin, ed. (1932). "Sir Charles Sherrington Biography". Les Prix Nobel. The Nobel Foundation. Retrieved 2008-07-23.CS1 maint: Extra text: authors list (link)
  25. ^ a b c d e f g Kusurkar, R. A. (2004). "Sir Charles Sherrington (1857 - 1952)". Journal of Postgraduate Medicine. 50 (3): 238–239. PMID 15377819.
  26. ^ Granit, R. (1967). Charles Scott Sherrington: An Appraisal. Garden City, NY: Double Day & Company. p. 3. OCLC 573353.
  27. ^ "Sherrington, Charles Scott (SHRN879CS)". A Cambridge Alumni Database. University of Cambridge.
  28. ^ University of Cambridge Calendar, 1894-95, p. 330
  29. ^ a b "Sir Charles Scott Sherrington". Who Named It?. 2008. Archived from the original on 7 October 2009. Retrieved 23 July 2008.
  30. ^ a b c Gibson, 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.
  31. ^ 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. Retrieved 22 February 2013.
  32. ^ Ipswich Museum Records.
  33. ^ a b Burke, 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.
  34. ^ Charles Scott Sherrington R. Scott Spurlock, University of Edinburgh
  35. ^ Man on his Nature, Sherrington
  36. ^ Quiroga, Rodrigo Quian (2013). "Gnostic cells in the 21st century". Acta Neurobiol. Exp. 73: 1–9.
  37. ^ "No. 32563". The London Gazette (Supplement). 31 December 1921. p. 10716.
  38. ^ Sherrington's Presidential Address to the British Association Meeting, held at Hull in 1922

External links

1857 in the United Kingdom

Events from the year 1857 in the United Kingdom.

Alfred Fröhlich

Alfred Fröhlich (August 15, 1871 – March 22, 1953) was an Austrian-American pharmacologist and neurologist born in Vienna.

Charles Smart Roy

Charles Smart Roy (21 January 1854 - 4 October 1897) was a British professor of pathology who worked at the University of Cambridge.

Roy was born at Arbroath, Forfarshire to Adam Roy, a shipowner. His early education was at his birthplace of Arbroath and later at the St. Andrews. He studied medicine at the University of Edinburgh, graduating with distinction in 1875 and joining as a Resident Physician at the Edinburgh Royal Infirmary.

He moved to the Brown Institution in London to conduct research in the physiological aspects of pleuro-pneumonia. During the Turko-Serbian war of 1876 he was in charge of a hospital at Janina in Turkey. After the war he went to Berlin to study under Emil du Bois-Reymond and Rudolf Virchow working on aspects of heart physiology. He obtained an M.D. from Edinburgh with a gold medal. He was invited to the Strasburg Physiological Institute where he worked with F.L. Goltz on blood circulation before moving to Leipzig in 1879 where he worked under Julius Cohnheim.In 1880 he moved to Cambridge as George Henry Lewes' student, working in the laboratory of Dr. Michael Foster. He taught advanced physiology to students. He succeeded Dr W.S. Greenfield as the director of the Brown Institution.

In 1884 he was elected professor of pathology at the University of Cambridge and Fellow of the Royal Society. He worked with others at the Pathological Laboratory such as Charles Scott Sherrington and several students became eminent pathologists including Ernest Hanbury Hankin, John George Adami and James Lorrain Smith.He died in Cambridge at the age of 43.

Chemical synapse

Chemical synapses are biological junctions through which neurons' signals can be exchanged to each other and to non-neuronal cells such as those in muscles or glands. Chemical synapses allow neurons to form circuits within the central nervous system. They are crucial to the biological computations that underlie perception and thought. They allow the nervous system to connect to and control other systems of the body.

At a chemical synapse, one neuron releases neurotransmitter molecules into a small space (the synaptic cleft) that is adjacent to another neuron. The neurotransmitters are kept within small sacs called synaptic vesicles, and are released into the synaptic cleft by exocytosis. These molecules then bind to neurotransmitter receptors on the postsynaptic cell's side of the synaptic cleft. Finally, the neurotransmitters must be cleared from the synapse through one of several potential mechanisms including enzymatic degradation or re-uptake by specific transporters either on the presynaptic cell or possibly by neuroglia to terminate the action of the transmitter.

The adult human brain is estimated to contain from 1014 to 5 × 1014 (100–500 trillion) synapses. Every cubic millimeter of cerebral cortex contains roughly a billion (short scale, i.e. 109) of them. The number of synapses in the human cerebral cortex has separately been estimated at 0.15 quadrillion (150 trillion)The word "synapse" comes from "synaptein", which Sir Charles Scott Sherrington and colleagues coined from the Greek "syn-" ("together") and "haptein" ("to clasp"). Chemical synapses are not the only type of biological synapse: electrical and immunological synapses also exist. Without a qualifier, however, "synapse" commonly means chemical synapse.

Ferrier Lecture

The Ferrier Lecture is a Royal Society lectureship given every three years "on a subject related to the advancement of natural knowledge on the structure and function of the nervous system". It was created in 1928 to honour the memory of Sir David Ferrier, a neurologist who was the first British scientist to electronically stimulate the brain for the purpose of scientific study.In its 90-year history, the Lecture has been given 30 times. It has never been given more than once by the same person. The first female to be awarded the honour was Prof. Christine Holt in 2017. The first lecture was given in 1929 by Charles Scott Sherrington, and was titled "Some functional problems attaching to convergence". The most recent lecturer was provided by Prof. Christine Holt, who presented a lecture in 2017 titled "understanding of the key molecular mechanisms involved in nerve growth, guidance and targeting which has revolutionised our knowledge of growing axon tips". In 1971, the lecture was given by two individuals (David Hunter Hubel and Torsten Nils Wiesel) on the same topic, with the title "The function and architecture of the visual cortex".

Florence Buchanan

Florence Buchanan (21 April 1867 — 13 March 1931) was a zoologist. She was awarded a London D.Sc. in 1902, was appointed as a Fellow of the University College London in 1904, and was awarded the American Association of Collegiate Alumnae's prize in 1910 for her research.

Fullerian Professor of Physiology

The Fullerian Chairs at the Royal Institution in London, England, were established by John 'Mad Jack' Fuller.

List of Fellows of the Royal Society elected in 1893

Fellows of the Royal Society elected in 1893.

List of people from the London Borough of Islington

Among those who were born in the London Borough of Islington, or have dwelt within the borders of the modern borough are (alphabetical order):

Douglas Adams, writer, lived on Arlington Avenue and Duncan Terrace, later renting his house to comedian Angus Deayton.

Adewale Akinnuoye-Agbaje, actor was born there

Lily Allen, singer and daughter of actor Keith Allen

Nadia Almada, first transsexual winner of Big Brother

Tash Aw, Whitbread Book Award-winning author

Julian Barratt and Julia Davis

Nina Bawden, author, has lived in Islington for many years

James Beck, actor, was born there

Tony Blair, former Prime Minister of the UK, lived at 1 Richmond Crescent before moving to Downing Street.

Helena Bonham Carter, actress

Jay Bothroyd, footballer

Jim Broadbent, actor, lives in the area

Jonny Buckland, lead guitarist of the band Coldplay

Alexandra Burke, singer and winner of The X Factor

Kathy Burke, actor and director, lives in Islington

Elaine Lordan, actor, appeared on EastEnders as Lynne Slater

Asa Butterfield, actor

Neve Campbell, Canadian actor

Jimmy Carr, comedian

Natalie Cassidy, actress, was born and raised in Islington

John Chapple, one of the last Field Marshals of Great Britain and Governor of Gibraltar

J. Smeaton Chase (1864–1923), travel writer and photographer who wrote about California; buried in Palm Springs, California

Caroline Chisholm, lived at 32 Charlton Place.

Sorcha Cusack, actor

Phil Daniels, actor

Alan Davies, actor and comedian, Jonathan Creek and Bob and Rose, lives in Highbury

Dido, singer, born in Islington and owns a property there

Colin Firth, Academy Award-winning actor.

Edwin Flack (1873–1935), athlete and tennis player

Jonathan Fortune, Sheffield United F.C. footballer, born in Islington

John Foxx, electronic musician and first Ultravox singer, lived there in the 1970s

Peaches Geldof, daughter of Bob Geldof and Paula Yates

John Glascock (1951–1979), (musician), bassist of Carmen from 1971–1974 and Jethro Tull from 1975–1979, born and raised in Islington

Kate Greenaway, children's writer and book illustrator, lived at 147 Upper Street for 20 years before moving to Holloway

Jonas Grimås, film and television director

Teriy Keys, music executive, entrepreneur, founder and co-chief executive officer of R.O.A.D. Group

Tony Hadley, lead singer, Spandau Ballet, born in Islington in 1960

Edmund Halley, Astronomer Royal and discoverer of Halley's Comet lived in Islington (exact location unknown) from 1665

Charlie G. Hawkins, actor, Darren Miller in EastEnders

Isabel Hilton, journalist and broadcaster

William Hogarth, artist, was born in Bartholomew Close in 1697 and spent his early years in Islington

Edward Irving, founder of the Catholic Apostolic Church, lived in Claremont Square

Yusuf Islam, aka Cat Stevens, musician

Ian Jack, writer and journalist

Boris Johnson, MP Former Mayor of London

Semothy Jones, songwriter/record producer, grew up in Holloway

Churchill Julius, lived at 44 Milner Square in 1881, vicar of Holy Trinity Islington – went on to become Archbishop of New Zealand.

George Julius, lived at 44 Milner Square in 1881, invented the world's first automatic totalisator.

Gary Kemp and Martin Kemp of Spandau Ballet, born in Islington, lived on Elmore Street

Skandar Keynes, actor The Chronicles of Narnia

Danny King, wrote The Burglar Diaries and Thieves Like Us

Charles Lamb, writer, lived in Chapel Street from 1796, at 64 Duncan Terrace and also in Colebrook Row

Edward Lear, writer, poet, artist, born in Islington

Heath Ledger, lived in Roman way, Islington while filming his final film in 2007 before his death

Vladimir Lenin, lived at 30 Holford Square from 1902 and later at 16 Percy Circus

Leona Lewis, singer

Louise Lombard, actor

Arthur Louis, singer, lived at 12 Richmond Avenue in the 1970s

Louisa Lytton, actor

Marianne Majerus, photographer

Dean Mason, association football player

Princess Martha Louise of Norway lived in Islington 2012-2014.

James McAvoy, actor

Cameron McKenna, television announcer and radio broadcaster

Scott Mills, Radio 1 DJ, lives here

Andrew Mitchell, Secretary of State for International Development in the British Government (from May 2010)

Ugo Monye, Harlequins and English rugby union player

Robert Muchamore, author of the CHERUB series

Sheree Murphy, actor, was born here

Scott Neal, actor, Beautiful Thing and PC Luke Ashton in The Bill

Edmund John Niemann, 19th century landscape artist born in Islington

Ed O'Brien, guitarist, Radiohead

Joe Orton, playwright, lived and was murdered in a flat at 25 Noel Road

George Orwell, writer, lived at 50 Lawford Road and in a flat at 27B Canonbury Square

Nicholas Owen, newsreader and broadcaster, was born in Islington and raised in Reigate, Surrey

David Oyelowo, actor, grew up in Islington and attended City and Islington College.

Stephen Poliakoff, playwright

Su Pollard, actor, Hi Di Hi

Anna Popplewell, actress The Chronicles of Narnia

Jacob Post, religious writer.

Sir Walter Raleigh, writer, poet, courtier and explorer lived in Upper Street between 1575 and 1581

Simon Rattle, conductor who lived in Islington for a period

Linda Robson, actor, Birds of a Feather

Francis Ronalds, inventor of the electric telegraph lived in Canonbury and then Highbury Terrace in the period 1789-1813.

Ronnie Ronalde, music hall performer famous for his singing, whistling, yodelling and imitations of bird song was born and raised in Islington

Jon Ronson, author, columnist, documentary maker

Salman Rushdie, writer who lived in Islington for a period

Dana, winner of the 1970 Eurovision Song Contest

Kaya Scodelario, actress

Andy Serkis, actor, The Lord of the Rings film trilogy

Martin Shaw, actor, Ray Doyle and Judge John Deed, lived in Noel Road, including while starring in The Professionals

Ben Shephard, TV presenter, lived in Islington 2001–2004

Sir Charles Scott Sherrington, neurologist, pathologist, bacteriologist, born in Islington

Sid Smith, novelist, journalist, lives in Islington

Nicky Spesh, rapper, lives in Islington

Mark Strong, actor born in Islington

Shana and Joe Swash, Eastenders actors

Ann Taylor (1782–1866), poet, writer, born in Islington

Sir Charles Todd FRS (1826–1910), astronomer

Laura Trevelyan, international BBC newsreader and correspondent, born in Islington, now resides in New York City

Ms. Triniti (born 1974), recording artist, born in Islington

Peter Vowell, schoolteacher, executed for high treason

Frank Warren, boxing promoter, born in Islington

Emma Watson, actor.

Samuel West, actor

Kenneth Williams (1926–1988), actor and comedian, born at 11 Bingfield Street, lived in Cromer Street.

Kate Winslet, actress

Elizabeth Wilkinson (1700s), English bare-knuckle boxing champion, known to be the first female boxer.

Edgar Wright, film director.

Hugo Young, journalist, lived in Milner Square from the late 1960s until the mid-1980s

Michael Foster (physiologist)

Sir Michael Foster (8 March 1836 – 29 January 1907) was an English physiologist. He was instrumental in organizing the Cambridge Biological School and acted as Secretary of the Royal Society.

Motor unit

A motor unit is made up of a motor neuron and the skeletal muscle fibers innervated by that motor neuron's axonal terminals. Groups of motor units often work together to coordinate the contractions of a single muscle; all of the motor units within a muscle are considered a motor pool. The concept was proposed by Charles Scott Sherrington.All muscle fibres in a motor unit are of the same fibre type. When a motor unit is activated, all of its fibres contract. In vertebrates, the force of a muscle contraction is controlled by the number of activated motor units.

The number of muscle fibers within each unit can vary within a particular muscle and even more from muscle to muscle; the muscles that act on the largest body masses have motor units that contain more muscle fibers, whereas smaller muscles contain fewer muscle fibers in each motor unit. For instance, thigh muscles can have a thousand fibers in each unit, while extraocular muscles might have ten. Muscles which possess more motor units (and thus have greater individual motor neuron innervation) are able to control force output more finely.

Motor units are organized slightly differently in invertebrates; each muscle has few motor units (typically less than 10), and each muscle fiber is innervated by multiple neurons, including excitatory and inhibitory neurons. Thus, while in vertebrates the force of contraction of muscles is regulated by how many motor units are activated, in invertebrates it is controlled by regulating the balance between excitatory and inhibitory signals.

Revue neurologique

The Revue neurologique (Neurological Review) is a French neurological and psychiatric medical journal. It was established in 1893 with Jean-Martin Charcot as adviser. Today it is the official journal of the Société Française de Neurologie (French Neurology Society). According to the Journal Citation Reports, the journal has a 2016 impact factor of 1.039.The monthly journal became the main periodical in France devoted to psychiatry and nervous diseases. It was made the official journal of the Société de Neurologie de Paris when the society was founded in 1899. It gradually displaced the quarterly Archives de Neurologie, publishing original articles by prominent French neurologists such as Charcot, Joseph Jules Dejerine, Édouard Brissaud and Fulgence Raymond, as well as work from foreign researchers including Charles Scott Sherrington of Great Britain, Alexander E. Scherbak of Russia, and João Baptista de Lacerda of Argentine. In November 1901 the Revue Neurologique published a report by Ladislav Haškovec of Prague describing two cases of akathisia. The journal also published summary articles that went into great detail on a particular topic.

Sherrington's law of reciprocal innervation

Sherrington's law of reciprocal innervation, also called Sherrington's law II explains how a muscle will relax when its opposite muscle (e.g., biceps/triceps) is activated. René Descartes had hypothesized as much in 1662.

Sherrington's law of reciprocal innervation states that: When a muscle contracts, its direct antagonist relaxes to an equal extent allowing smooth movement.

St Thomas's Hospital Medical School

St Thomas's Hospital Medical School in London was one of the oldest and most prestigious medical schools in the UK. The school was absorbed to form part of King's College London.

Sybil Cooper

Sybil Cooper (January 1900 – 1970), was a British physiologist.

Thomas Graham Brown

Thomas Graham Brown FRS (27 March 1882 – 28 October 1965) (usually known as T. Graham Brown) was a Scottish mountaineer and physiologist.

Vulpian–Heidenhain–Sherrington phenomenon

Vulpian–Heidenhain–Sherrington phenomenon is a term given for slow contraction of denervated skeletal muscle by stimulating the autonomic cholinergic fibers innervating its blood vessels. It is named after French neurologist Alfred Vulpian (1826–87), German physiologist Rudolf Heidenhain (1834–1897) and English neurophysiologist Charles Scott Sherrington (1857–1952).

Waynflete Professorship

The Waynflete Professorships are four professorial fellowships at the University of Oxford endowed by Magdalen College and named in honour of the college founder William of Waynflete, who had a great interest in science. These professorships are statutory professorships of the University, that is, they are professorships established in the university's regulations, and which are by those regulations attached to Magdalen College in particular. The oldest professorship is the Waynflete Professor of Metaphysical Philosophy. The three science professorships were created following the recommendation of the University Commission in 1857, in recognition of William of Waynflete's lifetime support of science. The professorships are the Waynflete Professor of Chemistry, the Waynflete Professor of Physiology, and the Waynflete Professor of Pure Mathematics.

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