Monday, November 11, 2013

William Harvey

William Harvey 2.jpg
William Harvey
Born 1 April 1578
Folkestone
Died 3 June 1657 (aged 79)
Roehampton
Nationality English
Fields Medicine
Anatomy
Doctoral advisor Hieronymus Fabricius
Known for Systemic circulation
Signature

William Harvey (1 April 1578 – 3 June 1657) was an English physician. He was the first to describe completely and in detail the systemic circulation and properties of blood being pumped to the brain and body by the heart, though earlier writers had provided precursors of the theory.[1][2] After his death the William Harvey Hospital was constructed in the town of Ashford, several miles from his birthplace of Folkestone.

Family

William's father, Thomas Harvey, was a jurat of Folkestone where he served the office of mayor in 1600. Records and personal descriptions delineate him as an overall calm, diligent and intelligent man whose "sons... revered, consulted and implicitly trusted in him... (they) made their father the treasurer of their wealth when they acquired great estates...(He) kept, employed and improved their gainings to their great advantage."[3] Thomas Harvey's portrait can still be seen in the central panel of a wall of the dining-room at Rolls Park, Chigwell, in Essex.

Life

Early life and the University of Padua

Harvey's initial education was carried out in Folkestone, where he learned Latin. He then entered the King's School (Canterbury). Harvey stayed at the King's School for five years, after which he joined Caius College in Cambridge.
Harvey graduated as a Bachelor of Arts from Caius College in 1597.[4] He then traveled through France and Germany to Italy, where he entered the University of Padua, in 1599.
During Harvey's years of study there, he developed a relationship with Fabricius and read Fabricius' De Venarum Ostiolis.
Harvey graduated as a Doctor of Medicine at the age of 24 from the University of Padua on 25 April 1602. It reports that Harvey had
"conducted himself so wonderfully well in the examination and had shown such skill, memory and learning that he had far surpassed even the great hopes which his examiners had formed of him."[5]

The College of Physicians, Marriage and Saint Bartholomew's Hospital

After graduating from Padua, Harvey immediately returned to England where he obtained the degree of Doctor of Medicine from the University of Cambridge that same year. Following this, Harvey established himself in London, joining the College of Physicians on 5 October 1604.
A few weeks after his admission, Harvey married Elizabeth Browne, "daughter of Lancelot Browne, (himself) Dr. Physic".[6][clarification needed] They had no children.
Elected a Fellow of the College of Physicians on 5 June 1607, Harvey accepted a position at St. Bartholomew's Hospital that he was to occupy for almost all the rest of his life. Succeeding a Dr Wilkinson on 14 October 1609, he became the Physician in charge at St. Bartholomew's Hospital, which enjoined him, "in God's most holy name" to "endeavor yourself to do the best of your knowledge in the profession of physic to the poor then present, or any other of the poor at any time of the week which shall be sent home unto you by the Hospitaller... You shall not, for favor, lucre or gain, appoint or write anything for the poor but such good and wholesome things as you shall think with your best advice will do the poor good, without any affection or respect to be had to the apothecary. And you shall take no gift or reward... for your counsel... This you will promise to do as you shall answer before God... "[7]
Harvey earned around thirty-three pounds a year and lived in a small house in Ludgate, although two houses in West Smithfield were attached as fringe benefits to the post of Physician. At this point, the physician's function consisted of a simple but thorough analysis of patients who were brought to the hospital once a week and the consequent writing of prescriptions.

Lumleian Lecturer

The next important phase of Harvey's life began with his appointment to the office of Lumleian lecturer on 4 August 1615. The Lumleian lectureship was founded by Lord Lumley and a Dr. Richard Caldwell in 1583, consisted in pronouncing lectures for a period of seven years, with the purpose of "spreading light" and increasing the general knowledge of anatomy throughout England.
Harvey began his lectures in April 1616. At this time, at the age of thirty-seven, he was described as "a man of lowest stature, round faced; his eyes small, round, very black and full of spirit; his hair as black as a raven and curling".[8] The notes which he used at the time are preserved in the British Museum.
At the beginning of his lectures, Harvey laid down the canons for his guidance:
  • "To show as much as may be at a glance, the whole belly for instance, and afterwards to subdivide the parts according to their positions and relations.
  • To point out what is peculiar to the actual body which is being dissected.
  • To supply only by speech what cannot be shown on your own credit and by authority.
  • To cut up as much as may be in the sight of the audience.
  • To enforce the right opinion by remarks drawn far and near, and to illustrate man by the structure of animals.
  • Not to praise or dispraise other anatomists, for all did well, and there was some excuse even for those who are in error.
  • Not to dispute with others, or attempt to confute them, except by the most obvious retort.
  • To state things briefly and plainly, yet not letting anything pass unmentioned which can be seen.
  • Not to speak of anything which can be as well explained without the body or can be read at home.
  • Not to enter into too much detail, or in too minute dissection, for the time does not permit.
  • To allot a definite time to each part of the body (i.e. first day's lectures dedicated to the abdomen, the second to the thorax, the third to the brain and so on."[9]

Physician to James I

Harvey continued to participate in the Lumleian lectures while also taking care of his patients at St. Bartholomew's Hospital; he thus soon attained an important and fairly lucrative practice, which climaxed with his appointment as 'Physician Extraordinary' to King James I on 3 February 1618. He seems to have similarly served various aristocrats, including Lord Chancellor Bacon. Also not may people know that William Harvey was also banished out of England with the future king at the time.
In 1628 he published in Frankfurt his completed treatise on the circulation of the blood, the De Motu Cordis. As a result of negative comments by other physicians Harvey "fell mightily in his practice",[10] but continued advancing his career. He was re-elected 'Censor' of the College of Physicians in 1629, having been elected for the first time in 1613 and the second time in 1625. Eventually, Harvey was also elected Treasurer of the College.

Excursions abroad, election as physician to Charles I and the English Civil War

At the age of fifty-two, Harvey received commands by the king to accompany the Duke of Lennox during his trip abroad. This voyage – the first after his return from Padua – lasted three years, taking Harvey through the countries of France and Spain during the Mantuan War and Plague. During this journey he wrote to Viscount Dorchester:
"I can complain that by the way we could scarce see a dog, crow, kite, raven or any other bird, or anything to anatomize, only some few miserable people, the relics of the war and the plague where famine had made anatomies before I came. It is scarce credible in so rich, populous, and plentiful countries as these were that so much misery and desolation, poverty and famine should in so short a time be, as we have seen. I interpret it well that it will be a great motive for all here to have and procure assurance of settled peace. It is time to leave fighting when there is nothing to eat, nothing to be keep, and nothing to be gotten".[11]
Having returned to England in 1632, Harvey accompanied King Charles I wherever he went as 'Physician in Ordinary.' In particular, Charles' hunting expeditions gave Harvey access to many deer carcasses; it was upon them that Harvey made many observations and consequent theories. Harvey returned to Italy in 1636, dining at the English College, Rome, as a guest of the Jesuits there, in October 1636. It is possible he met Galileo in Florence en route.[12]
During the English Civil War a mob of citizen-soldiers against the King entered Harvey's lodgings, stole his goods, and scattered his papers. The papers consisted of the records of a large number of dissections... of diseased bodies, with this observations on the development on insects, and a series of notes on comparative anatomy."[13] During this period, Harvey maintained his position, helped the wounded on several occasions and protected the King's children.
The conflicts of the Civil War soon led King Charles to Oxford, with Harvey attending, where the physician was made 'Doctor of Physic' in 1642 and later Warden of Merton College in 1645. "In Oxford he (Harvey) very soon settled down to his accustomed pursuits, unmindful of the clatter of arms and of the constant marching and countermarching around him, for the city remained the base of operations until its surrender... "[14]

Harvey's later years, death and burial

The surrender of Oxford in 1645 marks the beginning of Harvey's gradual retirement from public life and duties. Now sixty-eight years old and childless, Harvey had lost three brothers and wife at this time. He thus decided to return to London and live with his brothers Eliab and Daniel separately and in different periods of time. Having retired from St Bartholomew's Hospital and his various other aforementioned positions, he passed most of this time reading general literature. Several attempts to bring Harvey back into the 'working world' were made, however; here is an excerpt of one of Harvey's answers:
"Would you be the man who should recommend me to quit the peaceful haven where I now pass my life and launch again upon the faithless sea? You know full well what a storm my former lucubrations raised. Much better is it oftentimes to grow wise at home and in private, than by publishing what you have amassed with infinite labour, to stir up tempests that may rob you of peace and quiet for the rest of your days."[15]
Harvey died at Roehampton in the house of his brother Eliab on 3 June 1657. Descriptions of the event seem to show that he died of a cerebral hemorrhage from vessels long injured by gout: it is highly probable that the left middle cerebral artery malfunctioned, leading to a gradual accumulation of blood to the brain which eventually overwhelmed it. There exists a fairly detailed account of what happened on that day; according to the information at hand, Harvey:
"went to speak and found that he had the dead palsy in his tongue; then he saw what was to become of him. He knew there were then no hopes of his recovery, so presently he sends for his young nephews to come up to him. He then made signs (for seized with the dead palsy in his tongue he could not speak) to let him blood his tongue, which did him little or no good, and so ended his days, dying in the evening of the day on which he was stricken, the palsy giving him an easy passport."[16]
His will distributed his material goods and wealth throughout his extended family and also left a substantial amount of money to the College of Physicians.
Harvey was buried in Hempstead, Essex. The funeral procession started on the 26 June 1657 leading Harvey to be placed in the 'Harvey Chapel' built by Eliab. The conditions of Harvey's burial are also known: "Harvey was laid in the chapel between the bodies of his two nieces, and like them he was lept in lead, coffin less ".[17] On St.Luke's Day, October 18, 1883, Harvey's remains were reinterred, the leaden case carried from the vault by eight Fellows of the College of Physicians, and deposited in a sarcophagus containing his works and an inscription:
"The body of William Harvey lapt in lead, simply soldered, was laid without shell or enclosure of any kind in the Harvey vault of this Church of Hempstead, Essex, in June, 1657. In the course of time the lead enclosing the remains was, from expose and natural decay, so seriously damaged as to endanger its preservation, rendering some repair of it the duty of those interested in the memory of the illustrious discoverer of the circulation of the Blood. The Royal College of Physicians, of which corporate body Harvey was a munificent Benefactor did in the years 1882–1883, by permission of the Representatives of the Harvey family, undertake this duty. In accordance with this determination the leaden mortuary chest containing the remains of Harvey was repaired, and was, as far as possible, restored to its original state... "[18]

"De Motu Cordis" (otherwise known as "On the Motion of the Heart and Blood")

Published in 1628 in the city of Frankfurt (host to an annual book fair that Harvey knew would allow immediate dispersion of his work), this 72 page book contains the matured account of the circulation of the blood. Opening with a simple but clear dedication to King Charles I, the quarto has 17 chapters which give a perfectly clear and connected account of the action of the heart and the consequent movement of the blood around the body in a circuit. Having only a mere lens at his disposal, Harvey was not able to reach the adequate images that were attained through such microscopes used by Leeuwenhoek; thus he had to resort to theory – and not practical evidence – in certain parts of his book. After the first chapter, which simply outlines past ideas and accepted rules regarding the heart and lungs, Harvey moves on to a fundamental premise to his treatise, stating that it was extremely important to study the heart when it was active in order to truly comprehend its true movement; a task which even he found of great difficulty, as he says:
"...I found the task so truly arduous... that I was almost tempted to think... that the movement of the heart was only to be comprehended by God. For I could neither rightly perceive at first when the systole and when the diastole took place by reason of the rapidity of the movement..."[19]
This initial thought led Harvey's ambition and assiduousness to a detailed analysis of the overall structure of the heart (studied with less hindrances in cold-blooded animals). After this, Harvey goes on to an analysis of the arteries, showing how their pulsation depends upon the contraction of the left ventricle, while the contraction of the right ventricle propels its charge of blood into the pulmonary artery. Whilst doing this, the physician reiterates the fact that these two ventricles move together almost simultaneously and not independently like had been thought previously by his predecessors. This discovery was made while observing the heart of such animals as the eel and several other types of fish; indeed, the general study of countless animals was of utmost importance to the physician: among the ones already cited, one can add the study of the snail, the invisible shrimp, the chick before its hatching and even the pigeon. A digression to an experiment can be made to this note: using the inactive heart of a dead pigeon and placing upon it a finger wet with saliva, Harvey was able to witness a transitory and yet incontrovertible pulsation. He had just witnessed the heart's ability to recover from fatigue.
As early as the 17th century, William Harvey had already discerned the existence of the Ductus Arteriosus and explained its relative function. Here he says, "...in embryos, whilst the lungs are in a state of inaction, performing no function, subject to no movement any more than if they had not been present, Nature uses the two ventricles of the heart as if they formed but one for the transmission of the blood."[20] However, the apex of Harvey's work is probably the eighth chapter, in which he deals with the actual quantity of blood passing through the heart from the veins to the arteries. Coming into conflict with Galen's accepted view of the liver as the origin of venous blood, Harvey estimated the capacity of the heart, how much blood is expelled through each pump of the heart, and the amount of times the heart beats in a half an hour. All of these estimates were purposefully low, so that people could see the vast amount of blood Galen's theory required the liver to produce. He estimated that the capacity of the heart was 1.5 imperial fluid ounces (43 ml), and that every time the heart pumps, 1/8 of that blood is expelled. This led to Harvey's estimate that about 16 imperial fluid ounces (4.7 ml) of blood went through the heart every time it pumped. The next estimate he used was that the heart beats 1000 times every half an hour, which gave 10 pounds 6 ounces of blood in a half an hour, and when this number was multiplied by 48 half hours in a day he realized that the liver would have to produce 540 pounds of blood in a day.[contradictory]
Having this simple but essential mathematical proportion at hand – which proved the overall impossible aforementioned role of the liver – Harvey went on to prove how the blood circulated in a circle by means of countless experiments initially done on serpents and fish: tying their veins and arteries in separate periods of time, Harvey noticed the modifications which occurred; indeed, as he tied the veins, the heart would become empty, while as he did the same to the arteries, the organ would swell up.
Image of veins from Harvey's Exercitatio Anatomica de Motu Cordis et Sanguinis in Animalibus
This process was later performed on the human body (in the image on the right): the physician tied a tight ligature onto the upper arm of a person. This would cut off blood flow from the arteries and the veins. When this was done, the arm below the ligature was cool and pale, while above the ligature it was warm and swollen. The ligature was loosened slightly, which allowed blood from the arteries to come into the arm, since arteries are deeper in the flesh than the veins. When this was done, the opposite effect was seen in the lower arm. It was now warm and swollen. The veins were also more visible, since now they were full of blood. Harvey then noticed little bumps in the veins, which he realized were the valves of the veins, discovered by his teacher, Hieronymus Fabricius. Harvey tried to push blood in the vein down the arm, but to no avail. When he tried to push it up the arm, it moved quite easily. The same effect was seen in other veins of the body, except the veins in the neck. Those veins were different from the others – they did not allow blood to flow up, but only down. This led Harvey to believe that the veins allowed blood to flow to the heart, and the valves maintained the one way flow.
It is also important to state how Harvey had theorized the existence of capillaries: however, unable to discern them due to the aforementioned scarcity of instruments at this disposal, he was never truly capable of understanding how blood passed from the arterioles into the venules.

Views of the circulation of blood before Harvey

R.A. Young wrote: "Wiberg suggests that the early Greeks knew of the circulation, and quotes a passage from one of the Hippocratic writings which would bear that interpretation."[21]
At the time of Harvey's publication, Galen had been an influential medical authority for several centuries. Galen believed that blood passed between the ventricles by means of invisible pores. According to Galen's views, the venous system was quite separate from the arterial system, except when they came in contact through the unseen pores. Arabic scholar Ibn al-Nafis had disputed aspects of Galen's views, providing a model that seems to imply a form of pulmonary circulation in his Commentary on Anatomy in Avicenna's Canon (1242). Al-Nafis stated that blood moved from the heart to the lungs, where it mixed with air, and then back to the heart, from which it spread to the rest of the body.[22] Harvey's discoveries inevitably and historically came into conflict with Galen's teachings and the publication of his treatise De Motu Cordis incited considerable controversy within the medical community. Some doctors affirmed they would "rather err with Galen than proclaim the truth with Harvey."[23][24] Galen incompletely perceived the function of the heart, believing it a "productor of heat", while the function of its affluents, the arteries, was that of cooling the blood as the lungs "...fanned and cooled the heart itself".[25] Galen thought that during dilation the arteries sucked in air, while during their contraction they discharged vapours through pores in the flesh and skin.
Until the 17th century, two separate systems were thought to be involved in blood circulation: the natural system, containing venous blood which had its origin in the liver, and the vital system, containing arterial blood and the 'spirits' which flowed from the heart, distributing heat and life to all parts. Like bellows, the lungs fanned and cooled this vital blood.
Independently from Ibn Al-Nafis, Michael Servetus identified pulmonary circulation, but this discovery did not reach the public because it was written down for the first time in the Manuscript of Paris in 1546.[26] It was later published in the theological work which caused his execution in 1553, almost all copies of which were destroyed. Pulmonary circulation was described by Renaldus Columbus, Andrea Cesalpino and Vesalius, before Harvey would provide a refined and complete description of the circulatory system.

More facts on Harvey

In terms of his personality, information shows that William Harvey was seen as a "...humorous but extremely precise man...",[27] how he was often so immersed in his own thoughts that he would often suffer from insomnia (cured with a simple walk through the house), and how he was always ready for an open and direct conversation. He also loved the darkness, for it is said that it was there where "...he could best contemplate", thus sometimes hiding out in caves. A heavy drinker of coffee, Harvey would walk out combing his hair every morning full of energy and enthusiastic spirit through the fields. We have also come to understand Harvey's somewhat unorthodox method of dealing with his gout, here cited completely: "...his [Harvey's] cure was thus: he would sit with his legs bare...put them into a pail of water till he was almost dead with cold, then betake himself to his stove, and so 'twas gone".[28] Apart from the already mentioned love of literature, Harvey was also an intense and dedicated observer of birds during his free time: several long and detailed passages of citations could be written delineating his observations in such places as the "Pile of Boulders" (a small island in Lancashire), 'Bass Rock' (island of the East Coast of Scotland) and "The Bass" (another Scottish island).
A final allusion to the rules established and followed by the physician throughout his life can be made: general canons which are still followed in hospitals today, they embody a final representation of the man that was, William Harvey.
  1. "That none be taken into the Hospital but such as be curable, or but a certain number of such as are curable.
  2. That none lurk here for relief only or for slight causes.
  3. That the Chirurgions, in all difficult cases or where inward physic may be necessary, shall consult with the Doctor, at the times he sitteth once in the week and then the Surgeon himself relate to the Doctor what he conceiveth of the cure and what he hath done therein.
  4. That no Chirurgion or his man do trepan the head, pierce the body, dismember, or do any great operation on the body of any but with the approbation and the direction of the Doctor..."[29]

Saturday, October 12, 2013

Charles Darwin


Charles Darwin
Three quarter length studio photo showing Darwin's characteristic large forehead and bushy eyebrows with deep set eyes, pug nose and mouth set in a determined look. He is bald on top, with dark hair and long side whiskers but no beard or moustache. His jacket is dark, with very wide lapels, and his trousers are a light check pattern. His shirt has an upright wing collar, and his cravat is tucked into his waistcoat which is a light fine checked pattern.
Darwin, aged 45 in 1854, by then working towards publication of On the Origin of Species
Born Charles Robert Darwin
12 February 1809
The Mount, Shrewsbury, Shropshire, UK
Died 19 April 1882 (aged 73)
Down House, Downe, Kent, UK
Residence England
Citizenship British
Nationality British
Fields Natural history, Geology
Institutions tertiary education:
University of Edinburgh Medical School (medicine)
University of Cambridge (ordinary Bachelor of Arts)
professional institution:
Geological Society of London
Academic advisors John Stevens Henslow
Adam Sedgwick
Known for The Voyage of the Beagle
On the Origin of Species
evolution by
natural selection,
common descent
Influences Alexander von Humboldt
John Herschel
Charles Lyell
Influenced Joseph Dalton Hooker
Thomas Henry Huxley
George Romanes
Ernst Haeckel
Sir John Lubbock
Notable awards Royal Medal (1853)
Wollaston Medal (1859)
Copley Medal (1864)
Spouse Emma Darwin (married 1839)
Children 10 children (see list)
Signature
"Charles Darwin", with the surname underlined by a downward curve that mimics the curve of the initial "C"
Charles Robert Darwin, FRS (12 February 1809 – 19 April 1882) was an English naturalist and geologist,[1] best known for his contributions to evolutionary theory.[I] He established that all species of life have descended over time from common ancestors,[2] and in a joint publication with Alfred Russel Wallace introduced his scientific theory that this branching pattern of evolution resulted from a process that he called natural selection, in which the struggle for existence has a similar effect to the artificial selection involved in selective breeding.[3]
Darwin published his theory of evolution with compelling evidence in his 1859 book On the Origin of Species, overcoming scientific rejection of earlier concepts of transmutation of species.[4][5] By the 1870s the scientific community and much of the general public had accepted evolution as a fact. However, many favoured competing explanations and it was not until the emergence of the modern evolutionary synthesis from the 1930s to the 1950s that a broad consensus developed in which natural selection was the basic mechanism of evolution.[6][7] In modified form, Darwin's scientific discovery is the unifying theory of the life sciences, explaining the diversity of life.[8][9]
Darwin's early interest in nature led him to neglect his medical education at the University of Edinburgh; instead, he helped to investigate marine invertebrates. Studies at the University of Cambridge encouraged his passion for natural science.[10] His five-year voyage on HMS Beagle established him as an eminent geologist whose observations and theories supported Charles Lyell's uniformitarian ideas, and publication of his journal of the voyage made him famous as a popular author.[11]
Puzzled by the geographical distribution of wildlife and fossils he collected on the voyage, Darwin began detailed investigations and in 1838 conceived his theory of natural selection.[12] Although he discussed his ideas with several naturalists, he needed time for extensive research and his geological work had priority.[13] He was writing up his theory in 1858 when Alfred Russel Wallace sent him an essay which described the same idea, prompting immediate joint publication of both of their theories.[14] Darwin's work established evolutionary descent with modification as the dominant scientific explanation of diversification in nature.[6] In 1871 he examined human evolution and sexual selection in The Descent of Man, and Selection in Relation to Sex, followed by The Expression of the Emotions in Man and Animals. His research on plants was published in a series of books, and in his final book, he examined earthworms and their effect on soil.[15]
In recognition of Darwin's pre-eminence as a scientist, he was honoured with a major ceremonial funeral and buried in Westminster Abbey, close to John Herschel and Isaac Newton.[16] Darwin has been described as one of the most influential figures in human history.[17][18]

Biography

Early life and education

Charles Robert Darwin was born in Shrewsbury, Shropshire, England on 12 February 1809 at his family home, The Mount.[19] He was the fifth of six children of wealthy society doctor and financier Robert Darwin, and Susannah Darwin (née Wedgwood). He was the grandson of two prominent abolitionists: Erasmus Darwin on his father's side, and of Josiah Wedgwood on his mother's side.
Three quarter length portrait of seated boy smiling and looking at the viewer. He has straight mid brown hair, and wears dark clothes with a large frilly white collar. In his lap he holds a pot of flowering plants
The seven-year-old Charles Darwin in 1816.
Both families were largely Unitarian, though the Wedgwoods were adopting Anglicanism. Robert Darwin, himself quietly a freethinker, had baby Charles baptised in November 1809 in the Anglican St Chad's Church, Shrewsbury, but Charles and his siblings attended the Unitarian chapel with their mother. The eight-year-old Charles already had a taste for natural history and collecting when he joined the day school run by its preacher in 1817. That July, his mother died. From September 1818 he joined his older brother Erasmus attending the nearby Anglican Shrewsbury School as a boarder.[20]
Darwin spent the summer of 1825 as an apprentice doctor, helping his father treat the poor of Shropshire, before going to the University of Edinburgh Medical School, at the time the best medical school in the UK, with his brother Erasmus in October 1825. He found lectures dull and surgery distressing, so neglected his studies. He learned taxidermy from John Edmonstone, a freed black slave who had accompanied Charles Waterton in the South American rainforest, and often sat with this "very pleasant and intelligent man".[21]
In Darwin's second year he joined the Plinian Society, a student natural history group whose debates strayed into radical materialism. He assisted Robert Edmond Grant's investigations of the anatomy and life cycle of marine invertebrates in the Firth of Forth, and on 27 March 1827 presented at the Plinian his own discovery that black spores found in oyster shells were the eggs of a skate leech. One day, Grant praised Lamarck's evolutionary ideas. Darwin was astonished by Grant's audacity, but had recently read similar ideas in his grandfather Erasmus' journals.[22] Darwin was rather bored by Robert Jameson's natural history course which covered geology including the debate between Neptunism and Plutonism. He learned classification of plants, and assisted with work on the collections of the University Museum, one of the largest museums in Europe at the time.[23]
This neglect of medical studies annoyed his father, who shrewdly sent him to Christ's College, Cambridge, for a Bachelor of Arts degree as the first step towards becoming an Anglican parson. As Darwin was unqualified for the Tripos, he joined the ordinary degree course in January 1828.[24] He preferred riding and shooting to studying. His cousin William Darwin Fox introduced him to the popular craze for beetle collecting; Darwin pursued this zealously, getting some of his finds published in Stevens' Illustrations of British entomology. He became a close friend and follower of botany professor John Stevens Henslow and met other leading naturalists who saw scientific work as religious natural theology, becoming known to these dons as "the man who walks with Henslow". When his own exams drew near, Darwin focused on his studies and was delighted by the language and logic of William Paley's Evidences of Christianity.[25] In his final examination in January 1831 Darwin did well, coming tenth out of 178 candidates for the ordinary degree.[26]
Darwin had to stay at Cambridge until June. He studied Paley's Natural Theology, which made an argument for divine design in nature, explaining adaptation as God acting through laws of nature.[27] He read John Herschel's new book, which described the highest aim of natural philosophy as understanding such laws through inductive reasoning based on observation, and Alexander von Humboldt's Personal Narrative of scientific travels. Inspired with "a burning zeal" to contribute, Darwin planned to visit Tenerife with some classmates after graduation to study natural history in the tropics. In preparation, he joined Adam Sedgwick's geology course, then travelled with him in the summer for a fortnight, in order to map strata in Wales.[28]

Voyage of the Beagle

Route from Plymouth, England, south to Cape Verde then southwest across the Atlantic to Bahia, Brazil, south to Rio de Janeiro, Montevideo, the Falkland Islands, round the tip of South America then north to Valparaiso and Callao. Northwest to the Galapagos Islands before sailing west across the Pacific to New Zealand, Sydney, Hobart in Tasmania, and King George's Sound in Western Australia. Northwest to the Keeling Islands, southwest to Mauritius and Cape Town, then northwest to Bahia and northeast back to Plymouth.
The voyage of the Beagle
After a week with student friends at Barmouth, Darwin returned home on 29 August to find a letter from Henslow proposing him as a suitable (if unfinished) gentleman naturalist for a self-funded supernumerary place on HMS Beagle with captain Robert FitzRoy, more as a companion than a mere collector. The ship was to leave in four weeks on an expedition to chart the coastline of South America.[29] Robert Darwin objected to his son's planned two-year voyage, regarding it as a waste of time, but was persuaded by his brother-in-law, Josiah Wedgwood, to agree to (and fund) his son's participation.[30]
After delays, the voyage began on 27 December 1831; it lasted almost five years. As FitzRoy had intended, Darwin spent most of that time on land investigating geology and making natural history collections, while the Beagle surveyed and charted coasts.[6][31] He kept careful notes of his observations and theoretical speculations, and at intervals during the voyage his specimens were sent to Cambridge together with letters including a copy of his journal for his family.[32] He had some expertise in geology, beetle collecting and dissecting marine invertebrates, but in all other areas was a novice and ably collected specimens for expert appraisal.[33] Despite suffering badly from seasickness, Darwin wrote copious notes while on board the ship. Most of his zoology notes are about marine invertebrates, starting with plankton collected in a calm spell.[31][34]
On their first stop ashore at St. Jago, Darwin found that a white band high in the volcanic rock cliffs included seashells. FitzRoy had given him the first volume of Charles Lyell's Principles of Geology which set out uniformitarian concepts of land slowly rising or falling over immense periods,[II] and Darwin saw things Lyell's way, theorising and thinking of writing a book on geology.[35]
When they reached Brazil Darwin was delighted by the tropical forest,[36] but detested the sight of slavery.[37] The survey continued to the south in Patagonia. They stopped at Bahía Blanca, and in cliffs near Punta Alta Darwin made a major find of fossil bones of huge extinct mammals beside modern seashells, indicating recent extinction with no signs of change in climate or catastrophe. He identified the little-known Megatherium by a tooth and its association with bony armour which had at first seemed to him like a giant version of the armour on local armadillos. The finds brought great interest when they reached England.[38][39]
On rides with gauchos into the interior to explore geology and collect more fossils, Darwin gained social, political and anthropological insights into both native and colonial people at a time of revolution, and learnt that two types of rhea had separate but overlapping territories.[40][41] Further south he saw stepped plains of shingle and seashells as raised beaches showing a series of elevations. He read Lyell's second volume and accepted its view of "centres of creation" of species, but his discoveries and theorising challenged Lyell's ideas of smooth continuity and of extinction of species.[42][43]
On a sea inlet surrounded by steep hills, with high snow covered mountains in the distance, someone standing in an open canoe waves at a square-rigged sailing ship, seen from the front
As HMS Beagle surveyed the coasts of South America, Darwin theorised about geology and extinction of giant mammals.
Three Fuegians on board, who had been seized during the first Beagle voyage and had spent a year in England, were taken back to Tierra del Fuego as missionaries. Darwin found them friendly and civilised, yet their relatives seemed "miserable, degraded savages", as different as wild from domesticated animals.[44] To Darwin the difference showed cultural advances, not racial inferiority. Unlike his scientist friends, he now thought there was no unbridgeable gap between humans and animals.[45] A year on, the mission had been abandoned. The Fuegian they had named Jemmy Button lived like the other natives, had a wife, and had no wish to return to England.[46]
Darwin experienced an earthquake in Chile and saw signs that the land had just been raised, including mussel-beds stranded above high tide. High in the Andes he saw seashells, and several fossil trees that had grown on a sand beach. He theorised that as the land rose, oceanic islands sank, and coral reefs round them grew to form atolls.[47][48]
On the geologically new Galápagos Islands Darwin looked for evidence attaching wildlife to an older "centre of creation", and found mockingbirds allied to those in Chile but differing from island to island. He heard that slight variations in the shape of tortoise shells showed which island they came from, but failed to collect them, even after eating tortoises taken on board as food.[49][50] In Australia the marsupial rat-kangaroo and the platypus seemed so unusual that Darwin thought it was almost as though two distinct Creators had been at work.[51] He found the Aborigines "good-humoured & pleasant", and noted their depletion by European settlement.[52]
The Beagle investigated how the atolls of the Cocos (Keeling) Islands had formed, and the survey supported Darwin's theorising.[48] FitzRoy began writing the official Narrative of the Beagle voyages, and after reading Darwin's diary he proposed incorporating it into the account.[53] Darwin's Journal was eventually rewritten as a separate third volume, on natural history.[54]
In Cape Town Darwin and FitzRoy met John Herschel, who had recently written to Lyell praising his uniformitarianism as opening bold speculation on "that mystery of mysteries, the replacement of extinct species by others" as "a natural in contradistinction to a miraculous process".[55] When organising his notes as the ship sailed home, Darwin wrote that if his growing suspicions about the mockingbirds, the tortoises and the Falkland Islands Fox were correct, "such facts undermine the stability of Species", then cautiously added "would" before "undermine".[56] He later wrote that such facts "seemed to me to throw some light on the origin of species".[57]

Inception of Darwin's evolutionary theory

Three quarter length portrait of Darwin aged about 30, with straight brown hair receding from his high forehead and long side-whiskers, smiling quietly, in wide lapelled jacket, waistcoat and high collar with cravat.
While still a young man, Charles Darwin joined the scientific elite
When the Beagle reached Falmouth, Cornwall, on 2 October 1836, Darwin was already a celebrity in scientific circles as in December 1835 Henslow had fostered his former pupil's reputation by giving selected naturalists a pamphlet of Darwin's geological letters.[58] Darwin visited his home in Shrewsbury and saw relatives, then hurried to Cambridge to see Henslow, who advised on finding naturalists available to catalogue the collections and agreed to take on the botanical specimens. Darwin's father organised investments, enabling his son to be a self-funded gentleman scientist, and an excited Darwin went round the London institutions being fêted and seeking experts to describe the collections. Zoologists had a huge backlog of work, and there was a danger of specimens just being left in storage.[59]
Charles Lyell eagerly met Darwin for the first time on 29 October and soon introduced him to the up-and-coming anatomist Richard Owen, who had the facilities of the Royal College of Surgeons to work on the fossil bones collected by Darwin. Owen's surprising results included other gigantic extinct ground sloths as well as the Megatherium, a near complete skeleton of the unknown Scelidotherium and a hippopotamus-sized rodent-like skull named Toxodon resembling a giant capybara. The armour fragments were actually from Glyptodon, a huge armadillo-like creature as Darwin had initially thought.[60][39] These extinct creatures were related to living species in South America.[61]
In mid-December Darwin took lodgings in Cambridge to organise work on his collections and rewrite his Journal.[62] He wrote his first paper, showing that the South American landmass was slowly rising, and with Lyell's enthusiastic backing read it to the Geological Society of London on 4 January 1837. On the same day, he presented his mammal and bird specimens to the Zoological Society. The ornithologist John Gould soon announced that the Galapagos birds that Darwin had thought a mixture of blackbirds, "gros-beaks" and finches, were, in fact, twelve separate species of finches. On 17 February Darwin was elected to the Council of the Geological Society, and Lyell's presidential address presented Owen's findings on Darwin's fossils, stressing geographical continuity of species as supporting his uniformitarian ideas.[63]
Early in March, Darwin moved to London to be near this work, joining Lyell's social circle of scientists and experts such as Charles Babbage,[64] who described God as a programmer of laws. Darwin stayed with his freethinking brother Erasmus, part of this Whig circle and close friend of writer Harriet Martineau who promoted Malthusianism underlying the controversial Whig Poor Law reforms to stop welfare from causing overpopulation and more poverty. As a Unitarian she welcomed the radical implications of transmutation of species, promoted by Grant and younger surgeons influenced by Geoffroy. Transmutation was anathema to Anglicans defending social order,[65] but reputable scientists openly discussed the subject and there was wide interest in John Herschel's letter praising Lyell's approach as a way to find a natural cause of the origin of new species.[55]
Gould met Darwin and told him that the Galápagos mockingbirds from different islands were separate species, not just varieties, and what Darwin had thought was a "wren" was also in the finch group. Darwin had not labelled the finches by island, but from the notes of others on the Beagle, including FitzRoy, he allocated species to islands.[66] The two rheas were also distinct species, and on 14 March Darwin announced how their distribution changed going southwards.[67]
A page of hand-written notes, with a sketch of branching lines.
In mid-July 1837 Darwin started his "B" notebook on Transmutation of Species, and on page 36 wrote "I think" above his first evolutionary tree.
By mid-March, Darwin was speculating in his Red Notebook on the possibility that "one species does change into another" to explain the geographical distribution of living species such as the rheas, and extinct ones such as the strange Macrauchenia which resembled a giant guanaco. His thoughts on lifespan, asexual reproduction and sexual reproduction developed in his "B" notebook around mid-July on to variation in offspring "to adapt & alter the race to changing world" explaining the Galápagos tortoises, mockingbirds and rheas. He sketched branching descent, then a genealogical branching of a single evolutionary tree, in which "It is absurd to talk of one animal being higher than another", discarding Lamarck's independent lineages progressing to higher forms.[68]

Overwork, illness, and marriage

While developing this intensive study of transmutation, Darwin became mired in more work. Still rewriting his Journal, he took on editing and publishing the expert reports on his collections, and with Henslow's help obtained a Treasury grant of £1,000 to sponsor this multi-volume Zoology of the Voyage of H.M.S. Beagle, a sum equivalent to about £77,000 in 2011.[69] He stretched the funding to include his planned books on geology, and agreed unrealistic dates with the publisher.[70] As the Victorian era began, Darwin pressed on with writing his Journal, and in August 1837 began correcting printer's proofs.[71]
Darwin's health suffered from the pressure. On 20 September he had "an uncomfortable palpitation of the heart", so his doctors urged him to "knock off all work" and live in the country for a few weeks. After visiting Shrewsbury he joined his Wedgwood relatives at Maer Hall, Staffordshire, but found them too eager for tales of his travels to give him much rest. His charming, intelligent, and cultured cousin Emma Wedgwood, nine months older than Darwin, was nursing his invalid aunt. His uncle Jos pointed out an area of ground where cinders had disappeared under loam and suggested that this might have been the work of earthworms, inspiring "a new & important theory" on their role in soil formation which Darwin presented at the Geological Society on 1 November.[72]
William Whewell pushed Darwin to take on the duties of Secretary of the Geological Society. After initially declining the work, he accepted the post in March 1838.[73] Despite the grind of writing and editing the Beagle reports, Darwin made remarkable progress on transmutation, taking every opportunity to question expert naturalists and, unconventionally, people with practical experience such as farmers and pigeon fanciers.[6][74] Over time his research drew on information from his relatives and children, the family butler, neighbours, colonists and former shipmates.[75] He included mankind in his speculations from the outset, and on seeing an orangutan in the zoo on 28 March 1838 noted its childlike behaviour.[76]
Three quarter length portrait of woman aged about 30, with dark hair in centre parting straight on top, then falling in curls on each side. She smiles pleasantly and is wearing an open necked blouse with a large shawl pulled over her arms
Darwin chose to marry his cousin, Emma Wedgwood.
The strain took a toll, and by June he was being laid up for days on end with stomach problems, headaches and heart symptoms. For the rest of his life, he was repeatedly incapacitated with episodes of stomach pains, vomiting, severe boils, palpitations, trembling and other symptoms, particularly during times of stress such as attending meetings or making social visits. The cause of Darwin's illness remained unknown, and attempts at treatment had little success.[77]
On 23 June he took a break and went "geologising" in Scotland. He visited Glen Roy in glorious weather to see the parallel "roads" cut into the hillsides at three heights. He later published his view that these were marine raised beaches, but then had to accept that they were shorelines of a proglacial lake.[78]
Fully recuperated, he returned to Shrewsbury in July. Used to jotting down daily notes on animal breeding, he scrawled rambling thoughts about career and prospects on two scraps of paper, one with columns headed "Marry" and "Not Marry". Advantages included "constant companion and a friend in old age ... better than a dog anyhow", against points such as "less money for books" and "terrible loss of time."[79] Having decided in favour, he discussed it with his father, then went to visit Emma on 29 July. He did not get around to proposing, but against his father's advice he mentioned his ideas on transmutation.[80]

Malthus and natural selection

Continuing his research in London, Darwin's wide reading now included the sixth edition of Malthus's An Essay on the Principle of Population, and on 28 September 1838 he noted its assertion that human "population, when unchecked, goes on doubling itself every twenty five years, or increases in a geometrical ratio", a geometric progression so that population soon exceeds food supply in what is known as a Malthusian catastrophe. Darwin was well prepared to compare this to de Candolle's "warring of the species" of plants and the struggle for existence among wildlife, explaining how numbers of a species kept roughly stable. As species always breed beyond available resources, favourable variations would make organisms better at surviving and passing the variations on to their offspring, while unfavourable variations would be lost. He wrote that the "final cause of all this wedging, must be to sort out proper structure, & adapt it to changes", so that "One may say there is a force like a hundred thousand wedges trying force into every kind of adapted structure into the gaps of in the economy of nature, or rather forming gaps by thrusting out weaker ones."[6][81] This would result in the formation of new species.[6][82] As he later wrote in his Autobiography:
In October 1838, that is, fifteen months after I had begun my systematic enquiry, I happened to read for amusement Malthus on Population, and being well prepared to appreciate the struggle for existence which everywhere goes on from long-continued observation of the habits of animals and plants, it at once struck me that under these circumstances favourable variations would tend to be preserved, and unfavourable ones to be destroyed. The result of this would be the formation of new species. Here, then, I had at last got a theory by which to work..."[83]
By mid December Darwin saw a similarity between farmers picking the best stock in selective breeding, and a Malthusian Nature selecting from chance variants so that "every part of newly acquired structure is fully practical and perfected",[84] thinking this comparison "a beautiful part of my theory".[85] He later called his theory natural selection, an analogy with what he termed the artificial selection of selective breeding.[6]
On 11 November, he returned to Maer and proposed to Emma, once more telling her his ideas. She accepted, then in exchanges of loving letters she showed how she valued his openness in sharing their differences, also expressing her strong Unitarian beliefs and concerns that his honest doubts might separate them in the afterlife.[86] While he was house-hunting in London, bouts of illness continued and Emma wrote urging him to get some rest, almost prophetically remarking "So don't be ill any more my dear Charley till I can be with you to nurse you." He found what they called "Macaw Cottage" (because of its gaudy interiors) in Gower Street, then moved his "museum" in over Christmas. On 24 January 1839 Darwin was elected a Fellow of the Royal Society.[87]
On 29 January Darwin and Emma Wedgwood were married at Maer in an Anglican ceremony arranged to suit the Unitarians, then immediately caught the train to London and their new home.[88]

Geology books, Barnacles, evolutionary research

Darwin in his thirties, with his son dressed in a frock sitting on his knee.
Darwin in 1842 with his eldest son, William Erasmus Darwin
Darwin now had the framework of his theory of natural selection "by which to work",[83] as his "prime hobby".[89] His research included extensive experimental selective breeding of plants and animals, finding evidence that species were not fixed and investigating many detailed ideas to refine and substantiate his theory.[6] For fifteen years this work was in the background to his main occupation of writing on geology and publishing expert reports on the Beagle collections.[90]
When FitzRoy's Narrative was published in May 1839, Darwin's Journal and Remarks was such a success as the third volume that later that year it was published on its own.[91] Early in 1842, Darwin wrote about his ideas to Charles Lyell, who noted that his ally "denies seeing a beginning to each crop of species".[92]
Darwin's book The Structure and Distribution of Coral Reefs on his theory of atoll formation was published in May 1842 after more than three years of work, and he then wrote his first "pencil sketch" of his theory of natural selection.[93] To escape the pressures of London, the family moved to rural Down House in September.[94] On 11 January 1844 Darwin mentioned his theorising to the botanist Joseph Dalton Hooker, writing with melodramatic humour "it is like confessing a murder".[95][96] Hooker replied "There may in my opinion have been a series of productions on different spots, & also a gradual change of species. I shall be delighted to hear how you think that this change may have taken place, as no presently conceived opinions satisfy me on the subject."[97]
Path covered in sandy gravel winding through open woodland, with plants and shrubs growing on each side of the path.
Darwin's "sandwalk" at Down House was his usual "Thinking Path".[98]
By July, Darwin had expanded his "sketch" into a 230-page "Essay", to be expanded with his research results if he died prematurely.[99] In November the anonymously published sensational best-seller Vestiges of the Natural History of Creation brought wide interest in transmutation. Darwin scorned its amateurish geology and zoology, but carefully reviewed his own arguments. Controversy erupted, and it continued to sell well despite contemptuous dismissal by scientists.[100][101]
Darwin completed his third geological book in 1846. He now renewed a fascination and expertise in marine invertebrates, dating back to his student days with Grant, by dissecting and classifying the barnacles he had collected on the voyage, enjoying observing beautiful structures and thinking about comparisons with allied structures.[102] In 1847, Hooker read the "Essay" and sent notes that provided Darwin with the calm critical feedback that he needed, but would not commit himself and questioned Darwin's opposition to continuing acts of creation.[103]
In an attempt to improve his chronic ill health, Darwin went in 1849 to Dr. James Gully's Malvern spa and was surprised to find some benefit from hydrotherapy.[104] Then in 1851 his treasured daughter Annie fell ill, reawakening his fears that his illness might be hereditary, and after a long series of crises she died.[105]
In eight years of work on barnacles (Cirripedia), Darwin's theory helped him to find "homologies" showing that slightly changed body parts served different functions to meet new conditions, and in some genera he found minute males parasitic on hermaphrodites, showing an intermediate stage in evolution of distinct sexes.[106] In 1853 it earned him the Royal Society's Royal Medal, and it made his reputation as a biologist.[107] He resumed work on his theory of species in 1854, and in November realised that divergence in the character of descendants could be explained by them becoming adapted to "diversified places in the economy of nature".[108]

Publication of the theory of natural selection

Studio photo showing Darwin's characteristic large forehead and bushy eyebrows with deep set eyes, pug nose and mouth set in a determined look. He is bald on top, with dark hair and long side whiskers but no beard or moustache.
Charles Darwin, aged 46 in 1855, by then working towards publication of his theory of natural selection. He wrote to Hooker about this portrait, "if I really have as bad an expression, as my photograph gives me, how I can have one single friend is surprising."[109]
By the start of 1856, Darwin was investigating whether eggs and seeds could survive travel across seawater to spread species across oceans. Hooker increasingly doubted the traditional view that species were fixed, but their young friend Thomas Henry Huxley was firmly against transmutation of species. Lyell was intrigued by Darwin's speculations without realising their extent. When he read a paper by Alfred Russel Wallace, "On the Law which has Regulated the Introduction of New Species", he saw similarities with Darwin's thoughts and urged him to publish to establish precedence. Though Darwin saw no threat, he began work on a short paper. Finding answers to difficult questions held him up repeatedly, and he expanded his plans to a "big book on species" titled Natural Selection. He continued his researches, obtaining information and specimens from naturalists worldwide including Wallace who was working in Borneo. The American botanist Asa Gray showed similar interests, and on 5 September 1857 Darwin sent Gray a detailed outline of his ideas including an abstract of Natural Selection. In December, Darwin received a letter from Wallace asking if the book would examine human origins. He responded that he would avoid that subject, "so surrounded with prejudices", while encouraging Wallace's theorising and adding that "I go much further than you."[110]
Darwin's book was only partly written when, on 18 June 1858, he received a paper from Wallace describing natural selection. Shocked that he had been "forestalled", Darwin sent it on that day to Lyell, as requested by Wallace,[111][112] and although Wallace had not asked for publication, Darwin suggested he would send it to any journal that Wallace chose. His family was in crisis with children in the village dying of scarlet fever, and he put matters in the hands of Lyell and Hooker. After some discussion, they decided on a joint presentation at the Linnean Society on 1 July of On the Tendency of Species to form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection; however, Darwin's baby son died of the scarlet fever and he was too distraught to attend.[113]
There was little immediate attention to this announcement of the theory; the president of the Linnean Society remarked in May 1859 that the year had not been marked by any revolutionary discoveries.[114] Only one review rankled enough for Darwin to recall it later; Professor Samuel Haughton of Dublin claimed that "all that was new in them was false, and what was true was old."[115] Darwin struggled for thirteen months to produce an abstract of his "big book", suffering from ill health but getting constant encouragement from his scientific friends. Lyell arranged to have it published by John Murray.[116]
On the Origin of Species proved unexpectedly popular, with the entire stock of 1,250 copies oversubscribed when it went on sale to booksellers on 22 November 1859.[117] In the book, Darwin set out "one long argument" of detailed observations, inferences and consideration of anticipated objections.[118] His only allusion to human evolution was the understatement that "light will be thrown on the origin of man and his history".[119] His theory is simply stated in the introduction:
As many more individuals of each species are born than can possibly survive; and as, consequently, there is a frequently recurring struggle for existence, it follows that any being, if it vary however slightly in any manner profitable to itself, under the complex and sometimes varying conditions of life, will have a better chance of surviving, and thus be naturally selected. From the strong principle of inheritance, any selected variety will tend to propagate its new and modified form.[120]
He put a strong case for common descent, and at the end of the book concluded that:
There is grandeur in this view of life, with its several powers, having been originally breathed into a few forms or into one; and that, whilst this planet has gone cycling on according to the fixed law of gravity, from so simple a beginning endless forms most beautiful and most wonderful have been, and are being, evolved.[121]
The last word was the only variant of "evolved" in the first five editions of the book. "Evolutionism" at that time was associated with other concepts, most commonly with embryological development, and Darwin first used the word evolution in The Descent of Man in 1871, before adding it in 1872 to the 6th edition of The Origin of Species.[122]

Responses to publication

Three quarter length portrait of sixty-year-old man, balding, with white hair and long white bushy beard, with heavy eyebrows shading his eyes looking thoughtfully into the distance, wearing a wide lapelled jacket.
During the Darwin family's 1868 holiday in her Isle of Wight cottage, Julia Margaret Cameron took portraits showing the bushy beard Darwin grew between 1862 and 1866.
White bearded head of Darwin with the body of a crouching ape.
An 1871 caricature following publication of The Descent of Man was typical of many showing Darwin with an ape body, identifying him in popular culture as the leading author of evolutionary theory.
The book aroused international interest, with less controversy than had greeted the popular Vestiges of the Natural History of Creation. Though Darwin's illness kept him away from the public debates, he eagerly scrutinised the scientific response, commenting on press cuttings, reviews, articles, satires and caricatures, and corresponded on it with colleagues worldwide. Darwin had only said "Light will be thrown on the origin of man", but the first review claimed it made a creed of the "men from monkeys" idea from Vestiges.Amongst early favourable responses, Huxley's reviews swiped at Richard Owen, leader of the scientific establishment Huxley was trying to overthrow. In April, Owen's review attacked Darwin's friends and condescendingly dismissed his ideas, angering Darwin, but Owen and others began to promote ideas of supernaturally guided evolution.
The Church of England's response was mixed. Darwin's old Cambridge tutors Sedgwick and Henslow dismissed the ideas, but liberal clergymen interpreted natural selection as an instrument of God's design, with the cleric Charles Kingsley seeing it as "just as noble a conception of Deity". In 1860, the publication of Essays and Reviews by seven liberal Anglican theologians diverted clerical attention from Darwin, with its ideas including higher criticism attacked by church authorities as heresy. In it, Baden Powell argued that miracles broke God's laws, so belief in them was atheistic, and praised "Mr Darwin's masterly volume [supporting] the grand principle of the self-evolving powers of nature". Asa Gray discussed teleology with Darwin, who imported and distributed Gray's pamphlet on theistic evolution, Natural Selection is not inconsistent with Natural Theology. The most famous confrontation was at the public 1860 Oxford evolution debate during a meeting of the British Association for the Advancement of Science, where the Bishop of Oxford Samuel Wilberforce, though not opposed to transmutation of species, argued against Darwin's explanation and human descent from apes. Joseph Hooker argued strongly for Darwin, and Thomas Huxley's legendary retort, that he would rather be descended from an ape than a man who misused his gifts, came to symbolise a triumph of science over religion.
Even Darwin's close friends Gray, Hooker, Huxley and Lyell still expressed various reservations but gave strong support, as did many others, particularly younger naturalists. Gray and Lyell sought reconciliation with faith, while Huxley portrayed a polarisation between religion and science. He campaigned pugnaciously against the authority of the clergy in education,aiming to overturn the dominance of clergymen and aristocratic amateurs under Owen in favour of a new generation of professional scientists. Owen's claim that brain anatomy proved humans to be a separate biological order from apes was shown to be false by Huxley in a long running dispute parodied by Kingsley as the "Great Hippocampus Question", and discredited Owen.
Darwinism became a movement covering a wide range of evolutionary ideas. In 1863 Lyell's Geological Evidences of the Antiquity of Man popularised prehistory, though his caution on evolution disappointed Darwin. Weeks later Huxley's Evidence as to Man's Place in Nature showed that anatomically, humans are apes, then The Naturalist on the River Amazons by Henry Walter Bates provided empirical evidence of natural selection.[136] Lobbying brought Darwin Britain's highest scientific honour, the Royal Society's Copley Medal, awarded on 3 November 1864.That day, Huxley held the first meeting of what became the influential X Club devoted to "science, pure and free, untrammelled by religious dogmas".By the end of the decade most scientists agreed that evolution occurred, but only a minority supported Darwin's view that the chief mechanism was natural selection.
The Origin of Species was translated into many languages, becoming a staple scientific text attracting thoughtful attention from all walks of life, including the "working men" who flocked to Huxley's lectures. Darwin's theory also resonated with various movements at the time and became a key fixture of popular culture. Cartoonists parodied animal ancestry in an old tradition of showing humans with animal traits, and in Britain these droll images served to popularise Darwin's theory in an unthreatening way. While ill in 1862 Darwin began growing a beard, and when he reappeared in public in 1866 caricatures of him as an ape helped to identify all forms of evolutionism with Darwinism.

Descent of Man, sexual selection, and botany

Head and shoulders portrait, increasingly bald with rather uneven bushy white eyebrows and beard, his wrinkled forehead suggesting a puzzled frown
By 1878, an increasingly famous Darwin had suffered years of illness.
More detailed articles cover Darwin's life from Orchids to Variation, from Descent of Man to Emotions and from Insectivorous Plants to Worms
Despite repeated bouts of illness during the last twenty-two years of his life, Darwin's work continued. Having published On the Origin of Species as an abstract of his theory, he pressed on with experiments, research, and writing of his "big book". He covered human descent from earlier animals including evolution of society and of mental abilities, as well as explaining decorative beauty in wildlife and diversifying into innovative plant studies.
Enquiries about insect pollination led in 1861 to novel studies of wild orchids, showing adaptation of their flowers to attract specific moths to each species and ensure cross fertilisation. In 1862 Fertilisation of Orchids gave his first detailed demonstration of the power of natural selection to explain complex ecological relationships, making testable predictions. As his health declined, he lay on his sickbed in a room filled with inventive experiments to trace the movements of climbing plants.Admiring visitors included Ernst Haeckel, a zealous proponent of Darwinismus incorporating Lamarckism and Goethe's idealism.Wallace remained supportive, though he increasingly turned to Spiritualism.
The Variation of Animals and Plants under Domestication of 1868 was the first part of Darwin's planned "big book", and included his unsuccessful hypothesis of pangenesis attempting to explain heredity. It sold briskly at first, despite its size, and was translated into many languages. He wrote most of a second part, on natural selection, but it remained unpublished in his lifetime.
Darwin's figure is shown seated, dressed in a toga, in a circular frame labelled "TIME'S METER" around which a succession of figures spiral, starting with an earthworm emerging from the broken letters "CHAOS" then worms with head and limbs, followed by monkeys, apes, primitive men, a loin cloth clad hunter with a club, and a gentleman who tips his top hat to Darwin.
Punch's almanac for 1882, published shortly before Darwin's death, depicts him amidst evolution from chaos to Victorian gentleman with the title Man Is But A Worm.
Lyell had already popularised human prehistory, and Huxley had shown that anatomically humans are apes. With The Descent of Man, and Selection in Relation to Sex published in 1871, Darwin set out evidence from numerous sources that humans are animals, showing continuity of physical and mental attributes, and presented sexual selection to explain impractical animal features such as the peacock's plumage as well as human evolution of culture, differences between sexes, and physical and cultural racial characteristics, while emphasising that humans are all one species.[145] His research using images was expanded in his 1872 book The Expression of the Emotions in Man and Animals, one of the first books to feature printed photographs, which discussed the evolution of human psychology and its continuity with the behaviour of animals. Both books proved very popular, and Darwin was impressed by the general assent with which his views had been received, remarking that "everybody is talking about it without being shocked." His conclusion was "that man with all his noble qualities, with sympathy which feels for the most debased, with benevolence which extends not only to other men but to the humblest living creature, with his god-like intellect which has penetrated into the movements and constitution of the solar system–with all these exalted powers–Man still bears in his bodily frame the indelible stamp of his lowly origin."
His evolution-related experiments and investigations led to books on Insectivorous Plants, The Effects of Cross and Self Fertilisation in the Vegetable Kingdom, different forms of flowers on plants of the same species, and The Power of Movement in Plants. In his last book he returned to The Formation of Vegetable Mould through the Action of Worms.

Death and legacy

In 1882 he was diagnosed with what was called "angina pectoris" which then meant coronary thrombosis and disease of the heart. At the time of his death, the physicians diagnosed "anginal attacks", and "heart-failure".
He died at Down House on 19 April 1882. His last words were to his family, telling Emma "I am not the least afraid of death – Remember what a good wife you have been to me – Tell all my children to remember how good they have been to me", then while she rested, he repeatedly told Henrietta and Francis "It's almost worth while to be sick to be nursed by you".[149] He had expected to be buried in St Mary's churchyard at Downe, but at the request of Darwin's colleagues, after public and parliamentary petitioning, William Spottiswoode (President of the Royal Society) arranged for Darwin to be buried in Westminster Abbey, close to John Herschel and Isaac Newton.

Man dressed as Charles Darwin during Lyme Regis Fossil Festival.
Darwin had convinced most scientists that evolution as descent with modification was correct, and he was regarded as a great scientist who had revolutionised ideas. Though few agreed with his view that "natural selection has been the main but not the exclusive means of modification", he was honoured in June 1909 by more than 400 officials and scientists from across the world who met in Cambridge to commemorate his centenary and the fiftieth anniversary of On the Origin of Species. During this period, which has been called "the eclipse of Darwinism", scientists proposed various alternative evolutionary mechanisms which eventually proved untenable. The development of the modern evolutionary synthesis from the 1930s to the 1950s, incorporating natural selection with population genetics and Mendelian genetics, brought broad scientific consensus that natural selection was the basic mechanism of evolution. This synthesis set the frame of reference for modern debates and refinements of the theory.

Children

Darwin's children
William Erasmus Darwin (27 December 1839 – 1914)
Anne Elizabeth Darwin (2 March 1841 – 23 April 1851)
Mary Eleanor Darwin (23 September 1842 – 16 October 1842)
Henrietta Emma "Etty" Darwin (25 September 1843 – 1929)
George Howard Darwin (9 July 1845 – 7 December 1912)
Elizabeth "Bessy" Darwin (8 July 1847 – 1926)
Francis Darwin (16 August 1848 – 19 September 1925)
Leonard Darwin (15 January 1850 – 26 March 1943)
Horace Darwin (13 May 1851 – 29 September 1928)
Charles Waring Darwin (6 December 1856 – 28 June 1858)
The Darwins had ten children: two died in infancy, and Annie's death at the age of ten had a devastating effect on her parents. Charles was a devoted father and uncommonly attentive to his children.Whenever they fell ill, he feared that they might have inherited weaknesses from inbreeding due to the close family ties he shared with his wife and cousin, Emma Wedgwood. He examined this topic in his writings, contrasting it with the advantages of crossing amongst many organisms.[152] Despite his fears, most of the surviving children and many of their descendants went on to have distinguished careers (see Darwin-Wedgwood family).
Of his surviving children, George, Francis and Horace became Fellows of the Royal Society, distinguished as astronomer, botanist and civil engineer, respectively. His son Leonard went on to be a soldier, politician, economist, eugenicist and mentor of the statistician and evolutionary biologist Ronald Fisher.[156]

Views and opinions

Religious views

Three quarter length studio photo of seated girl about nine years old, looking slightly plump and rather solemn, in a striped dress, holding a basket of flowers on her lap.
In 1851 Darwin was devastated when his daughter Annie died. By then his faith in Christianity had dwindled, and he had stopped going to church.
Darwin's family tradition was nonconformist Unitarianism, while his father and grandfather were freethinkers, and his baptism and boarding school were Church of England. When going to Cambridge to become an Anglican clergyman, he did not doubt the literal truth of the Bible. He learned John Herschel's science which, like William Paley's natural theology, sought explanations in laws of nature rather than miracles and saw adaptation of species as evidence of design On board the Beagle, Darwin was quite orthodox and would quote the Bible as an authority on morality. He looked for "centres of creation" to explain distribution, and related the antlion found near kangaroos to distinct "periods of Creation".
By his return he was critical of the Bible as history, and wondered why all religions should not be equally valid. In the next few years, while intensively speculating on geology and transmutation of species, he gave much thought to religion and openly discussed this with his wife Emma, whose beliefs also came from intensive study and questioning. The theodicy of Paley and Thomas Malthus vindicated evils such as starvation as a result of a benevolent creator's laws which had an overall good effect. To Darwin, natural selection produced the good of adaptation but removed the need for design, and he could not see the work of an omnipotent deity in all the pain and suffering such as the ichneumon wasp paralysing caterpillars as live food for its eggs. He still viewed organisms as perfectly adapted, and On the Origin of Species reflects theological views. Though he thought of religion as a tribal survival strategy, Darwin was reluctant to give up the idea of God as an ultimate lawgiver. He was increasingly troubled by the problem of evil.
Darwin remained close friends with the vicar of Downe, John Innes, and continued to play a leading part in the parish work of the church, but from around 1849 would go for a walk on Sundays while his family attended church. He considered it "absurd to doubt that a man might be an ardent theist and an evolutionist" and, though reticent about his religious views, in 1879 he wrote that "I have never been an atheist in the sense of denying the existence of a God. – I think that generally ... an agnostic would be the most correct description of my state of mind."
The "Lady Hope Story", published in 1915, claimed that Darwin had reverted to Christianity on his sickbed. The claims were repudiated by Darwin's children and have been dismissed as false by historians.

Human society

Darwin's views on social and political issues reflected his time and social position. He thought men's eminence over women was the outcome of sexual selection, a view disputed by Antoinette Brown Blackwell in The Sexes Throughout Nature. He valued European civilisation and saw colonisation as spreading its benefits, with the sad but inevitable effect of extermination of savage peoples who did not become civilised. Darwin's theories presented this as natural, and were cited to promote policies which went against his humanitarian principles.Darwin was strongly against slavery, against "ranking the so-called races of man as distinct species", and against ill-treatment of native people.
Darwin was intrigued by his half-cousin Francis Galton's argument, introduced in 1865, that statistical analysis of heredity showed that moral and mental human traits could be inherited, and principles of animal breeding could apply to humans. In The Descent of Man Darwin noted that aiding the weak to survive and have families could lose the benefits of natural selection, but cautioned that withholding such aid would endanger the instinct of sympathy, "the noblest part of our nature", and factors such as education could be more important. When Galton suggested that publishing research could encourage intermarriage within a "caste" of "those who are naturally gifted", Darwin foresaw practical difficulties, and thought it "the sole feasible, yet I fear utopian, plan of procedure in improving the human race", preferring to simply publicise the importance of inheritance and leave decisions to individuals. Francis Galton named this field of study "eugenics" in 1883.

Evolutionary social movements

Full length portrait of a very thin white bearded Darwin, seated but leaning eagerly forward and smiling.
Caricature from 1871 Vanity Fair
Darwin's fame and popularity led to his name being associated with ideas and movements which at times had only an indirect relation to his writings, and sometimes went directly against his express comments.
Thomas Malthus had argued that population growth beyond resources was ordained by God to get humans to work productively and show restraint in getting families, this was used in the 1830s to justify workhouses and laissez-faire economics.Evolution was by then seen as having social implications, and Herbert Spencer's 1851 book Social Statics based ideas of human freedom and individual liberties on his Lamarckian evolutionary theory.
Soon after the Origin was published in 1859, critics derided his description of a struggle for existence as a Malthusian justification for the English industrial capitalism of the time. The term Darwinism was used for the evolutionary ideas of others, including Spencer's "survival of the fittest" as free-market progress, and Ernst Haeckel's racist ideas of human development. Writers used natural selection to argue for various, often contradictory, ideologies such as laissez-faire dog-eat dog capitalism, racism, warfare, colonialism and imperialism. However, Darwin's holistic view of nature included "dependence of one being on another"; thus pacifists, socialists, liberal social reformers and anarchists such as Peter Kropotkin stressed the value of co-operation over struggle within a species. Darwin himself insisted that social policy should not simply be guided by concepts of struggle and selection in nature.
After the 1880s a eugenics movement developed on ideas of biological inheritance, and for scientific justification of their ideas appealed to some concepts of Darwinism. In Britain, most shared Darwin's cautious views on voluntary improvement and sought to encourage those with good traits in "positive eugenics". During the "Eclipse of Darwinism" a scientific foundation for eugenics was provided by Mendelian genetics. Negative eugenics to remove the "feebleminded" were popular in America, Canada and Australia, and eugenics in the United States introduced compulsory sterilization laws, followed by several other countries. Subsequently, Nazi eugenics brought the field into disrepute.
The term "Social Darwinism" was used infrequently from around the 1890s, but became popular as a derogatory term in the 1940s when used by Richard Hofstadter to attack the laissez-faire conservatism of those like William Graham Sumner who opposed reform and socialism. Since then it has been used as a term of abuse by those opposed to what they think are the moral consequences of evolution.

Commemoration

Three-quarter portrait of a senior Darwin dressed in black before a black background. His face and six-inch white beard are dramatically lit from the side. His eyes are shaded by his brows and look directly and thoughtfully at the viewer.
In 1881 Darwin was an eminent figure, still working on his contributions to evolutionary thought that had an enormous effect on many fields of science.
During Darwin's lifetime, many geographical features were given his name. An expanse of water adjoining the Beagle Channel was named Darwin Sound by Robert FitzRoy after Darwin's prompt action, along with two or three of the men, saved them from being marooned on a nearby shore when a collapsing glacier caused a large wave that would have swept away their boats, and the nearby Mount Darwin in the Andes was named in celebration of Darwin's 25th birthday. When the Beagle was surveying Australia in 1839, Darwin's friend John Lort Stokes sighted a natural harbour which the ship's captain Wickham named Port Darwin: a nearby settlement was renamed Darwin in 1911, and it became the capital city of Australia's Northern Territory.
More than 120 species and nine genera have been named after Darwin. In one example, the group of tanagers related to those Darwin found in the Galápagos Islands became popularly known as "Darwin's finches" in 1947, fostering inaccurate legends about their significance to his work.
Darwin's work has continued to be celebrated by numerous publications and events. The Linnean Society of London has commemorated Darwin's achievements by the award of the Darwin–Wallace Medal since 1908. Darwin Day has become an annual celebration, and in 2009 worldwide events were arranged for the bicentenary of Darwin's birth and the 150th anniversary of the publication of On the Origin of Species.
Darwin has been commemorated in the UK, with his portrait printed on the reverse of £10 banknotes printed along with a hummingbird and HMS Beagle, issued by the Bank of England.
A life size seated statue of Darwin can be seen in the main hall of the Natural History Museum in London. . A seated statue of Darwin stands in front of Shrewsbury Library, the building that used to house Shrewsbury School, which Darwin attended as a boy.
Darwin College, a postgraduate college at Cambridge University, is named after the Darwin family.

Works

Darwin was a prolific writer. Even without publication of his works on evolution, he would have had a considerable reputation as the author of The Voyage of the Beagle, as a geologist who had published extensively on South America and had solved the puzzle of the formation of coral atolls, and as a biologist who had published the definitive work on barnacles. While On the Origin of Species dominates perceptions of his work, The Descent of Man and The Expression of the Emotions in Man and Animals had considerable impact, and his books on plants including The Power of Movement in Plants were innovative studies of great importance, as was his final work on The Formation of Vegetable Mould through the Action of Worms.