Great Hippocampus Question
Encyclopedia
The Great Hippocampus Question was a 19th century scientific controversy about the anatomy
Anatomy
Anatomy is a branch of biology and medicine that is the consideration of the structure of living things. It is a general term that includes human anatomy, animal anatomy , and plant anatomy...

 of ape
Ape
Apes are Old World anthropoid mammals, more specifically a clade of tailless catarrhine primates, belonging to the biological superfamily Hominoidea. The apes are native to Africa and South-east Asia, although in relatively recent times humans have spread all over the world...

s and human uniqueness. The dispute between Thomas Henry Huxley and Richard Owen
Richard Owen
Sir Richard Owen, FRS KCB was an English biologist, comparative anatomist and palaeontologist.Owen is probably best remembered today for coining the word Dinosauria and for his outspoken opposition to Charles Darwin's theory of evolution by natural selection...

 became central to the scientific debate on human evolution
Human evolution
Human evolution refers to the evolutionary history of the genus Homo, including the emergence of Homo sapiens as a distinct species and as a unique category of hominids and mammals...

 that followed Charles Darwin
Charles Darwin
Charles Robert Darwin FRS was an English naturalist. He established that all species of life have descended over time from common ancestry, and proposed the scientific theory that this branching pattern of evolution resulted from a process that he called natural selection.He published his theory...

's publication of On the Origin of Species. The name comes from the title of a satire the Reverend Charles Kingsley
Charles Kingsley
Charles Kingsley was an English priest of the Church of England, university professor, historian and novelist, particularly associated with the West Country and northeast Hampshire.-Life and character:...

 wrote about the arguments, which in modified form appeared as "the great hippopotamus test" in Kingsley's book for children, The Water-Babies, A Fairy Tale for a Land Baby
The Water-Babies, A Fairy Tale for a Land Baby
The Water-Babies, A Fairy Tale for a Land Baby is a children's novel by the Reverend Charles Kingsley. Written in 1862–1863 as a serial for Macmillan's Magazine, it was first published in its entirety in 1863...

. Together with other humorous skits on the topic, this helped to spread and popularise Darwin's ideas on evolution.

Background

In 1836 Charles Darwin
Charles Darwin
Charles Robert Darwin FRS was an English naturalist. He established that all species of life have descended over time from common ancestry, and proposed the scientific theory that this branching pattern of evolution resulted from a process that he called natural selection.He published his theory...

 returned from the Beagle voyage
Second voyage of HMS Beagle
The second voyage of HMS Beagle, from 27 December 1831 to 2 October 1836, was the second survey expedition of HMS Beagle, under captain Robert FitzRoy who had taken over command of the ship on its first voyage after her previous captain committed suicide...

 with fossil collections which the anatomist Richard Owen
Richard Owen
Sir Richard Owen, FRS KCB was an English biologist, comparative anatomist and palaeontologist.Owen is probably best remembered today for coining the word Dinosauria and for his outspoken opposition to Charles Darwin's theory of evolution by natural selection...

 described, contributing to the inception of Darwin's theory
Inception of Darwin's theory
The inception of Darwin's theory occurred during an intensively busy period which began when Charles Darwin returned from the survey voyage of the Beagle, with his reputation as a fossil collector and geologist already established...

 of natural selection
Natural selection
Natural selection is the nonrandom process by which biologic traits become either more or less common in a population as a function of differential reproduction of their bearers. It is a key mechanism of evolution....

. Darwin outlined his theory in an Essay of 1844, and discussed transmutation with his friend Joseph Dalton Hooker
Joseph Dalton Hooker
Sir Joseph Dalton Hooker OM, GCSI, CB, MD, FRS was one of the greatest British botanists and explorers of the 19th century. Hooker was a founder of geographical botany, and Charles Darwin's closest friend...

. He did not tell Owen, who as the up-and-coming "English Cuvier
Georges Cuvier
Georges Chrétien Léopold Dagobert Cuvier or Jean Léopold Nicolas Frédéric Cuvier , known as Georges Cuvier, was a French naturalist and zoologist...

" held the conventional belief that every species was uniquely created perfectly adapted. Owen's brilliance and political skills made him a leading figure in the scientific establishment, developing ideas of divine archetypes produced by vague secondary laws similar to a form of theistic evolution
Theistic evolution
Theistic evolution or evolutionary creation is a concept that asserts that classical religious teachings about God are compatible with the modern scientific understanding about biological evolution...

, while emphasising the differences separating man from ape. At the end of 1844 Vestiges of the Natural History of Creation
Vestiges of the Natural History of Creation
Vestiges of the Natural History of Creation is a unique work of speculative natural history published anonymously in England in 1844. It brought together various ideas of stellar evolution with the progressive transmutation of species in an accessible narrative which tied together numerous...

brought wide public interest in transmutation of species
Transmutation of species
Transmutation of species was a term used by Jean Baptiste Lamarck in 1809 for his theory that described the altering of one species into another, and the term is often used to describe 19th century evolutionary ideas that preceded Charles Darwin's theory of natural selection...

 and the idea that humans were descended from apes, and after a slow initial response, strong condemnation from the scientific establishment.

Darwin discussed his interest in transmutation with friends including Charles Lyell
Charles Lyell
Sir Charles Lyell, 1st Baronet, Kt FRS was a British lawyer and the foremost geologist of his day. He is best known as the author of Principles of Geology, which popularised James Hutton's concepts of uniformitarianism – the idea that the earth was shaped by slow-moving forces still in operation...

, and Hooker eventually read Darwin's Essay in 1847. When Thomas Henry Huxley savagely reviewed the latest edition of Vestiges in 1854, Darwin wrote to him, making friends while cautiously admitting to being "almost as unorthodox about species". Huxley had become increasingly irritated by Owen's condescension and manipulation, and having gained a teaching position at the school of mining, began openly attacking Owen's work.

Hippocampus minor

In 1564 a prominent feature on the floor of the Lateral ventricles
Lateral ventricles
The lateral ventricles are part of the ventricular system of the brain. Classified as part of the telencephalon, they are the largest of the ventricles....

 of the brain was named the hippocampus
Hippocampus
The hippocampus is a major component of the brains of humans and other vertebrates. It belongs to the limbic system and plays important roles in the consolidation of information from short-term memory to long-term memory and spatial navigation. Humans and other mammals have two hippocampi, one in...

 by Aranzi
Julius Caesar Aranzi
Julius Caesar Aranzi was a leading figure in the history of the science of human anatomy.He was born in Bologna, the son of Ottaviano di Jacopo and Maria Maggi...

 as its curved shape on each side supposedly reminded him of a seahorse, the Hippocampus (apart from a spell after Mayer mistakenly used the term hippopotamus
Hippopotamus
The hippopotamus , or hippo, from the ancient Greek for "river horse" , is a large, mostly herbivorous mammal in sub-Saharan Africa, and one of only two extant species in the family Hippopotamidae After the elephant and rhinoceros, the hippopotamus is the third largest land mammal and the heaviest...

 in 1779, and was followed by others until 1829). At that same time a ridge on the occipital horn
Posterior horn of lateral ventricle
The posterior horn of the lateral ventricle passes into the occipital lobe, its direction being backward and lateralward, and then medialward....

 was named the calcar avis, but in 1786 this was renamed the hippocampus minor, with the hippocampus being called the hippocampus major.
Richard Owen
Richard Owen
Sir Richard Owen, FRS KCB was an English biologist, comparative anatomist and palaeontologist.Owen is probably best remembered today for coining the word Dinosauria and for his outspoken opposition to Charles Darwin's theory of evolution by natural selection...

 presented several papers on the anatomical differences between apes and humans, arguing that they had been created separately and stressing the impossibility of apes being transmuted into men. In 1857 he went even further, presenting an authoritative paper to the Linnean Society of London
Linnean Society of London
The Linnean Society of London is the world's premier society for the study and dissemination of taxonomy and natural history. It publishes a zoological journal, as well as botanical and biological journals...

 on his anatomical studies of primate
Primate
A primate is a mammal of the order Primates , which contains prosimians and simians. Primates arose from ancestors that lived in the trees of tropical forests; many primate characteristics represent adaptations to life in this challenging three-dimensional environment...

 brains and asserting that humans were not merely a distinct biological order
Order (biology)
In scientific classification used in biology, the order is# a taxonomic rank used in the classification of organisms. Other well-known ranks are life, domain, kingdom, phylum, class, family, genus, and species, with order fitting in between class and family...

 of primates, as had been accepted by great anatomists such as Carl Linnaeus and Georges Cuvier
Georges Cuvier
Georges Chrétien Léopold Dagobert Cuvier or Jean Léopold Nicolas Frédéric Cuvier , known as Georges Cuvier, was a French naturalist and zoologist...

, but a separate sub-class
Class (biology)
In biological classification, class is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, order, family, genus, and species, with class fitting between phylum and order...

 of mammal
Mammal
Mammals are members of a class of air-breathing vertebrate animals characterised by the possession of endothermy, hair, three middle ear bones, and mammary glands functional in mothers with young...

ia, distinct from all the other primates and mammals generally. Owen supported his argument with a figure by himself of a South American monkey, a figure of a negro's brain by Friedrich Tiedemann
Friedrich Tiedemann
Friedrich Tiedemann was a German anatomist and physiologist.He was born at Cassel, the eldest son of Dietrich Tiedemann , a philosopher and psychologist of considerable repute. He graduated in medicine at Marburg in 1804, but soon abandoned practice...

, and of a chimpanzee's brain by Jacobus Schroeder van der Kolk
Jacobus Schroeder van der Kolk
Jacobus Ludovicus Conradus Schroeder van der Kolk was a Dutch anatomist and physiologist, and an influential researcher into the causes of epilepsy and mental illness....

 and Willem Vrolik
Willem Vrolik
Willem Vrolik was a Dutch anatomist and pathologist who was a native of Amsterdam. He was a pioneer in the field of vertebrate teratology....

.

While Owen conceded the "all-pervading similitude of structure—every tooth, every bone, strictly homologous" which made it difficult for anatomists to determine the difference between man and ape, he based his new classification on three characteristics which to him distinguished mankind's "highest form of brain", the most important being his claim that only the human brain
Human brain
The human brain has the same general structure as the brains of other mammals, but is over three times larger than the brain of a typical mammal with an equivalent body size. Estimates for the number of neurons in the human brain range from 80 to 120 billion...

 has a hippocampus minor. To Owen in 1857, this feature together with the extent to which the "posterior lobe"
Occipital lobe
The occipital lobe is the visual processing center of the mammalian brain containing most of the anatomical region of the visual cortex. The primary visual cortex is Brodmann area 17, commonly called V1...

 projected beyond the cerebellum
Cerebellum
The cerebellum is a region of the brain that plays an important role in motor control. It may also be involved in some cognitive functions such as attention and language, and in regulating fear and pleasure responses, but its movement-related functions are the most solidly established...

 and the presence of the posterior horn were how man "fulfills his destiny as the supreme master of this earth and of the lower creation." Charles Darwin
Charles Darwin
Charles Robert Darwin FRS was an English naturalist. He established that all species of life have descended over time from common ancestry, and proposed the scientific theory that this branching pattern of evolution resulted from a process that he called natural selection.He published his theory...

 commented, "Owen's is a grand Paper; but I cannot swallow Man making a division as distinct from a Chimpanzee, as an ornithorhynchus
Platypus
The platypus is a semi-aquatic mammal endemic to eastern Australia, including Tasmania. Together with the four species of echidna, it is one of the five extant species of monotremes, the only mammals that lay eggs instead of giving birth to live young...

 from a Horse: I wonder what a Chimpanzee wd. say to this?". Owen repeated the paper as the Rede Lecture
Rede Lecture
The Sir Robert Rede's Lecturer is an annual appointment to give a public lecture, the Sir Robert Rede's Lecture at the University of Cambridge. It is named for Sir Robert Rede, who was Chief Justice of the Common Pleas in the sixteenth century.-Initial series:The initial series of lectures ranges...

 at the University of Cambridge
University of Cambridge
The University of Cambridge is a public research university located in Cambridge, United Kingdom. It is the second-oldest university in both the United Kingdom and the English-speaking world , and the seventh-oldest globally...

 on 10 May 1859 when he was the first to be given an honorary degree by the university.

To Thomas Henry Huxley the claim about the hippocampus minor appeared to be a significant blunder by Owen, and Huxley began systematically dissecting the brains of monkeys, determined that "before I have done with that mendacious humbug I will nail him out, like a kite to a barn door, an example to all evil doers." He did not discuss this in public at this stage, but continued to attack Owen's other ideas, aiming to undermine Owen's status. At his 17 June 1858 Royal Institution
Royal Institution
The Royal Institution of Great Britain is an organization devoted to scientific education and research, based in London.-Overview:...

 Croonian Lecture
Croonian Lecture
The Croonian Lectures are prestigious lectureships given at the invitation of the Royal Society and the Royal College of Physicians.Among the papers of William Croone at his death in 1684, was a plan to endow one lectureship at both the Royal Society and the Royal College of Physicians...

 "On the Theory of the Vertebrate Skull", Huxley directly challenged Owen's central idea of archetypes shown by homology, with Owen in the audience. Huxley's aim was to overcome the domination of science by wealthy clergymen led by Owen, in order to create a professional salaried scientific civil service and make science secular
Secularism
Secularism is the principle of separation between government institutions and the persons mandated to represent the State from religious institutions and religious dignitaries...

. Under Darwin's influence he took up transmutation as a way of dividing science from theology, and in January 1859 argued that "it is as respectable to be modified monkey as modified dirt".

Owen and Huxley debate human and ape brain structure

Huxley was among the friends rallying round the publication of Darwin's On the Origin of Species, and was sharpening his "beak and claws" to disembowel "the curs who will bark and yelp". Charles Kingsley
Charles Kingsley
Charles Kingsley was an English priest of the Church of England, university professor, historian and novelist, particularly associated with the West Country and northeast Hampshire.-Life and character:...

 was sent a review copy, and told Darwin that he had "long since, from watching the crossing of domesticated animals and plants, learnt to disbelieve the dogma of the permanence of species." Darwin was delighted that this "celebrated author and divine" had "gradually learnt to see that it is just as noble a conception of the Deity to believe that He created a few original forms capable of self-development into other and needful forms, as to believe that He required a fresh act of creation to supply the voids caused by the action of His laws."

While reviews were by custom anonymous, their authors were usually known. Huxley's reviews of On the Origin of Species irritated Owen, whose own anonymous review in April praised himself and his own axiom of the continuous operation of the ordained becoming of living things, took offence at the way the creationist position had been depicted, and complained that his own pre-eminence had been ignored. Owen bitterly attacked Huxley, Hooker and Darwin, but also signalled acceptance of a kind of evolution as a teleological
Teleology
A teleology is any philosophical account which holds that final causes exist in nature, meaning that design and purpose analogous to that found in human actions are inherent also in the rest of nature. The word comes from the Greek τέλος, telos; root: τελε-, "end, purpose...

 plan in a continuous "ordained becoming", with new species appearing by natural birth.

The dispute between Huxley and Owen over human uniqueness began in public at the 1860 Oxford evolution debate
1860 Oxford evolution debate
The 1860 Oxford evolution debate took place at the Oxford University Museum on 30 June 1860, seven months after the publication of Charles Darwin's On the Origin of Species. Several prominent British scientists and philosophers participated, including Thomas Henry Huxley, Bishop Samuel...

, during a meeting of the British Association for the Advancement of Science in Oxford on Thursday 28 June 1860. After Charles Daubeny
Charles Daubeny
Charles Giles Bridle Daubeny was an English chemist, botanist and geologist.Daubeny was born at Stratton near Cirencester in Gloucestershire, the son of the Rev. James Daubeny. He studied at Magdalen College, Oxford under Dr. John Kidd...

's paper "On the Final Causes of the Sexuality of Plants with Particular Reference to Mr. Darwin's Work", the chairman asked Huxley for comments, but he declined as he thought the public venue inappropriate. Owen then spoke of facts which would enable the public to "come to some conclusions ... of the truth of Mr. Darwin's theory", reportedly arguing that "the brain of the gorilla was more different from that of man than from that of the lowest primate particularly because only man had a posterior lobe, a posterior horn, and a hippocampus minor." In response, Huxley flatly but politely "denied altogether that the difference between the brain of the gorilla and man was so great" in a "direct and unqualified contradiction" of Owen, citing previous studies as well as promising to provide detailed support for his position.

Anguish over the death of his son of scarlet fever
Scarlet fever
Scarlet fever is a disease caused by exotoxin released by Streptococcus pyogenes. Once a major cause of death, it is now effectively treated with antibiotics...

 in September 1860 pushed Huxley to the brink, from which Kingsley rescued him by a series of letters. Huxley put his fury over the death into composing a paper which violently assaulted Owen's ideas and professional reputation. It was published in January 1861 in the first issue of Huxley's relaunched Natural History Review
Natural History Review
The Natural History Review was a short-lived, quarterly journal devoted to natural history. It was published in Dublin and London between 1854 and 1865....

magazine, and presented citations, quotations and letters from leading anatomists to attack Owen's three claims, aiming to prove him "guilty of wilful and deliberate falsehood" by citing Owen himself, and (with less clear cut justification) the anatomists whose illustrations Owen had used in the 1857 paper. While readily agreeing that the human brain differed from that of apes in size, proportions and complexity of convolutions, Huxley played the significance of these features down, and argued that to a lesser extent these also differed between the "highest" and "lowest" human races. Darwin congratulated Huxley on this "smasher" against the "canting humbug" Owen. From February to May Huxley delivered a very popular series of sixpenny
Penny (British pre-decimal coin)
The penny of the Kingdom of Great Britain and later of the United Kingdom, was in circulation from the early 18th century until February 1971, Decimal Day....

 lectures for working men at the School of Mines
Imperial College London
Imperial College London is a public research university located in London, United Kingdom, specialising in science, engineering, business and medicine...

 where he taught, on "The Relation of Man to the Rest of the Animal Kingdom". He told his wife that "My working men stick by me wonderfully, the house being fuller than ever last night. By next Friday evening they will all be convinced that they are monkeys."
Gorilla
Gorilla
Gorillas are the largest extant species of primates. They are ground-dwelling, predominantly herbivorous apes that inhabit the forests of central Africa. Gorillas are divided into two species and either four or five subspecies...

s became the topic of the day with the return of the explorer Paul du Chaillu
Paul du Chaillu
Paul Belloni du Chaillu was a French-American traveler and anthropologist. He became famous in the 1860s as the first modern outsider to confirm the existence of gorillas and the Pygmy people of central Africa. He later researched the prehistory of Scandinavia.-Early life:His date and place of...

. Owen arranged for him to speak and display his collections on stage at a spectacular Royal Geographical Society
Royal Geographical Society
The Royal Geographical Society is a British learned society founded in 1830 for the advancement of geographical sciences...

 meeting on 25 February, and followed this by giving a lecture at the Royal Institution
Royal Institution
The Royal Institution of Great Britain is an organization devoted to scientific education and research, based in London.-Overview:...

 on 19 March on the brains of The Gorilla and the Negro, asserting that the dispute was one of interpretation rather than fact, and hedging his previous claim by stating that humans alone had a hippocampus minor "as defined in human anatomy". This lecture was published in the Atnenœum
Athenaeum (magazine)
The Athenaeum was a literary magazine published in London from 1828 to 1921. It had a reputation for publishing the very best writers of the age....

on 23 March with unlabelled and inaccurate illustrations, and Huxley's response in the next issue a week later, Man and the Apes, ridiculed Owen's use of these illustrations and failure to mention the findings of anatomists that the three structures were present in animals. In the following week's issue Owen's letter blamed "the Artist" for the illustrations, but claimed that the argument was correct and referred the reader to his 1858 paper. In the Atnenœum of 13 April Huxley responded to this repetition of the claim by writing that "Life is too short to occupy oneself with the slaying of the slain more than once."

Each Saturday, Darwin read the latest ripostes in the Atnenœum. Owen tried to smear Huxley by portraying him as an "advocate of man's origins from a transmuted ape", and one of his contributions was titled "Ape-Origin of Man as Tested by the Brain". This backfired, as Huxley had already delighted Darwin by speculating on "pithecoid man" (ape-like man), and was glad of the invitation to publicly turn the anatomy of brain structure into a question of human ancestry. Darwin egged him on from Down, writing "Oh Lord what a thorn you must be in the poor dear man's side". Huxley told Darwin's friend Joseph Dalton Hooker
Joseph Dalton Hooker
Sir Joseph Dalton Hooker OM, GCSI, CB, MD, FRS was one of the greatest British botanists and explorers of the 19th century. Hooker was a founder of geographical botany, and Charles Darwin's closest friend...

, "Owen occupied an entirely untenable position ... The fact is he made a prodigious blunder in commencing the attack, and now his only chance is to be silent and let people forget the exposure. I do not believe that in the whole history of science there is a case of any man of reputation getting himself into such a contemptible position. He will be the laughing-stock of all the continental anatomists."

Public interest and satire

This very public slanging match attracted wide attention, and humorists were quick to take up the opportunity for satire. Punch
Punch (magazine)
Punch, or the London Charivari was a British weekly magazine of humour and satire established in 1841 by Henry Mayhew and engraver Ebenezer Landells. Historically, it was most influential in the 1840s and 50s, when it helped to coin the term "cartoon" in its modern sense as a humorous illustration...

featured the issue several times that year, notably on 18 May 1861 when a cartoon under the heading Monkeyana showed a standing gorilla with a sign parodying Josiah Wedgwood's anti-slavery slogan "Am I Not A Man And A Brother?". This was accompanied by a satirical poem by "Gorilla" at the zoo asking to be told if he was "A man in ape's shape, An anthropoid ape, Or monkey deprived of his tail?", and noting:
It then recounts Huxley's ripostes, and:
The poem was actually by the eminent palaeontologist Sir Philip Egerton who, as a trustee of the Royal College of Surgeons
Royal College of Surgeons of England
The Royal College of Surgeons of England is an independent professional body and registered charity committed to promoting and advancing the highest standards of surgical care for patients, regulating surgery, including dentistry, in England and Wales...

 and the British Museum
British Museum
The British Museum is a museum of human history and culture in London. Its collections, which number more than seven million objects, are amongst the largest and most comprehensive in the world and originate from all continents, illustrating and documenting the story of human culture from its...

, acted as Owen's patron. When a delighted Huxley found out who the author of the piece was, he thought it "speaks volumes for Owen's perfect success in damning himself."

In the second issue of Huxley's Natural History Review, an article by George Rolleston
George Rolleston
George Rolleston MA MD FRCP FRS was an English physician and zoologist. He was the first Linacre Professor of Anatomy and Physiology to be appointed at the University of Oxford, a post he held from 1860 until his death in 1881...

 on the orangutan
Orangutan
Orangutans are the only exclusively Asian genus of extant great ape. The largest living arboreal animals, they have proportionally longer arms than the other, more terrestrial, great apes. They are among the most intelligent primates and use a variety of sophisticated tools, also making sleeping...

 brain showed the features that Owen claimed apes lacked, and when Owen responded in a letter to the Annals and Magazine of Natural History
Journal of Natural History
The Journal of Natural History is a scientific journal published by Taylor & Francis focusing on entomology and zoology. The journal was established in 1841 under the name Annals and Magazine of Natural History and obtained its current title in 1967...

that the issue was a matter of definition rather than fact, Huxley made a public dissection of a spider monkey
Spider monkey
Spider monkeys of the genus Ateles are New World monkeys in the subfamily Atelinae, family Atelidae. Like other atelines, they are found in tropical forests of Central and South America, from southern Mexico to Brazil...

 that had died at the zoo, to support his case. In the following issue John Marshall provided detailed measurements making the same point about the chimpanzee
Chimpanzee
Chimpanzee, sometimes colloquially chimp, is the common name for the two extant species of ape in the genus Pan. The Congo River forms the boundary between the native habitat of the two species:...

, as well as explaining how a chimpanzee's brain could be distorted by not being properly preserved and removed from the skull, so that it would look like the one in Owen's illustration.

The Great Hippocampus Question

The debate continued in 1862. A detailed paper by William Henry Flower
William Henry Flower
Sir William Henry Flower KCB FRCS FRS was an English comparative anatomist and surgeon. Flower became a leading authority on mammals, and especially on the primate brain...

 in the prestigious journal, the Philosophical Transactions of the Royal Society
Philosophical Transactions of the Royal Society
The Philosophical Transactions of the Royal Society is a scientific journal published by the Royal Society of London. It was established in 1665, making it the first journal in the world exclusively devoted to science, and it has remained in continuous publication ever since, making it the world's...

, reviewed the earlier literature and presented his own studies based on having dissected sixteen species of primates, including prosimian
Prosimian
Prosimians are a grouping of mammals defined as being primates, but not monkeys or apes. They include, among others, lemurs, bushbabies, and tarsiers. They are considered to have characteristics that are more primitive than those of monkeys and apes. Prosimians are the only primates native to...

s, monkeys and an orangutan
Orangutan
Orangutans are the only exclusively Asian genus of extant great ape. The largest living arboreal animals, they have proportionally longer arms than the other, more terrestrial, great apes. They are among the most intelligent primates and use a variety of sophisticated tools, also making sleeping...

. Having stated at the outset that he had no opinion on transmutation or the origin of humans, he refuted Owen's three claims, and went further, stating that in relation to the mass of the brain, the hippocampus minor was proportionately largest in the marmoset
Marmoset
Marmosets are the 22 New World monkey species of the genera Callithrix, Cebuella, Callibella, and Mico. All four genera are part of the biological family Callitrichidae. The term marmoset is also used in reference to the Goeldi's Monkey, Callimico goeldii, which is closely related.Most marmosets...

, and proportionately smallest in mankind. The paper used terms recently coined by Huxley, and Flower was one of his close colleagues. Huxley presented more evidence against Owen in his Natural History Review
Natural History Review
The Natural History Review was a short-lived, quarterly journal devoted to natural history. It was published in Dublin and London between 1854 and 1865....

. The Dutch anatomists Jacobus Schroeder van der Kolk
Jacobus Schroeder van der Kolk
Jacobus Ludovicus Conradus Schroeder van der Kolk was a Dutch anatomist and physiologist, and an influential researcher into the causes of epilepsy and mental illness....

 and Willem Vrolik
Willem Vrolik
Willem Vrolik was a Dutch anatomist and pathologist who was a native of Amsterdam. He was a pioneer in the field of vertebrate teratology....

 found that Owen had repeatedly used their 1849 illustration of a chimpanzee's brain to support his arguments, and to prevent the public from being misled they dissected the brain of an orangutan that had died in the Amsterdam zoo, reporting at a meeting of the Royal Netherlands Academy of Arts and Sciences
Royal Netherlands Academy of Arts and Sciences
The Royal Netherlands Academy of Arts and Sciences is an organisation dedicated to the advancement of science and literature in the Netherlands...

 that the three features Owen claimed were unique to humans were present in this ape. They admitted that their earlier illustration was incorrect due to the way they had removed the brain for inspection, and suggested that Owen had become "lost" and "fell into a trap" in debating against Darwin. Huxley reprinted the report, in French, in his Review. His confrontations with Owen went on.

At the 1862 British Association meeting in Cambridge that year, Owen presented two papers opposing Darwin: one claimed that the adaptations of the Aye-aye
Aye-aye
The aye-aye is a lemur, a strepsirrhine primate native to Madagascar that combines rodent-like teeth and a special thin middle finger to fill the same ecological niche as a woodpecker...

 disproved evolution, and the second paper reiterated Owen's claims about human brains being unique, as well as discussing the question of whether apes have toes or thumbs. Huxley said Owen appeared to be "lying & shuffling", and Huxley's allies presented successive attacks on Owen. This was the first British Association annual meeting attended by Charles Kingsley
Charles Kingsley
Charles Kingsley was an English priest of the Church of England, university professor, historian and novelist, particularly associated with the West Country and northeast Hampshire.-Life and character:...

, and during the meeting he produced a privately printed satirical skit on the argument, "a little squib for circulation among his friends" written in the style of the then popular stage character Lord Dundreary
Lord Dundreary
Lord Dundreary is a character of the 1858 British play Our American Cousin by Tom Taylor. He is the personification of a good-natured, brainless aristocrat. The role was created on stage by Edward Askew Sothern. The most famous scene involved Dundreary reading a letter from his even sillier...

, a good natured but brainless aristocrat known for huge bushy sideburns and for mangling proverbs or sayings in "Dundrearyism
Dundrearyism
A Dundrearyism is an aphorism, proverb, colloquial phrase, saying or riddle humorously combined with another in such a way to render it nonsensical. An example is "Birds of a feather gather no moss."...

s". The skit was titled Speech of Lord Dundreary in Section D, on Friday Last, On the Great Hippocampus Question.
The British Medical Journal
BMJ
BMJ is a partially open-access peer-reviewed medical journal. Originally called the British Medical Journal, the title was officially shortened to BMJ in 1988. The journal is published by the BMJ Group, a wholly owned subsidiary of the British Medical Association...

asked, "Is it not high time that the annual passage of barbed words between Professor Owen and Professor Huxley, on the cerebral distinction between men and monkeys, should cease? ... Continued on its present footing, it becomes a hindrance and an injury to science, a joke for the populace, and a scandal to the scientific world." The London Quarterly Review
Quarterly Review
The Quarterly Review was a literary and political periodical founded in March 1809 by the well known London publishing house John Murray. It ceased publication in 1967.-Early years:...

took up the joke, describing the confrontation of Owen with Huxley and his supporters Rolleston
George Rolleston
George Rolleston MA MD FRCP FRS was an English physician and zoologist. He was the first Linacre Professor of Anatomy and Physiology to be appointed at the University of Oxford, a post he held from 1860 until his death in 1881...

 and Flower dramatically: "Animation increased, 'decorous reticence' was at an end, and all parties enjoyed the scene except the disputants. Surely apes were never before so honoured, as to be the theme of the warmest discussion in one of the two principal university towns in England. Strange sight was this, that three or four most accomplished anatomists were contending against each other like so many gorillas, and either reducing man to a monkey, or elevating the monkey to the man!" In October the Medical Times and Gazette reported Owen's presentation with full detail of the responses by Huxley, Rolleston and Flower, as well as Owen's rebuttal. The dispute continued in the next two issues of the magazine.

The great hippopotamus test

At about the same time as he was attending the Cambridge British Association meeting in 1862, instalments of Charles Kingsley's story for children The Water-Babies, A Fairy Tale for a Land Baby
The Water-Babies, A Fairy Tale for a Land Baby
The Water-Babies, A Fairy Tale for a Land Baby is a children's novel by the Reverend Charles Kingsley. Written in 1862–1863 as a serial for Macmillan's Magazine, it was first published in its entirety in 1863...

were being published in Macmillan's Magazine
Macmillan's Magazine
Macmillan's Magazine was a monthly British magazine from 1859 to 1907 published by Alexander Macmillan.The magazine was a literary periodical that published fiction and non-fiction works from primarily British authors. Thomas Hughes had convinced Macmillan to found the magazine. The first editor...

as a serial. Kingsley incorporated material modified from his skit about Dundreary's speech On the Great Hippocampus Question, as well as other references to the protagonists, the British Association, and notable scientists of the day. When the protagonist Tom is turned into a water-baby by the fairies, the question is raised that if there were water-babies, surely someone would have caught one and "put it into spirits, or into the Illustrated News, or perhaps cut it into two halves, poor dear little thing, and sent one to Professor Owen, and one to Professor Huxley, to see what they would each say about it." As for the suggestion that a water-baby is contrary to nature;
Keeping up an even-handed treatment, Kingsley introduced as a character in the story Professor Ptthmllnsprts (Put-them-all-in-spirits) as an amalgam of Owen and Huxley, satirising each in turn. Like the very possessive Owen, the Professor was "very good to all the world as long as it was good to him. Only one fault he had, which cock-robins have likewise, as you may see if you look out of the nursery window—that, when any one else found a curious worm, he would hop round them, and peck them, and set up his tail, and bristle up his feathers, just as a cock-robin would; and declare that he found the worm first; and that it was his worm; and, if not, that then it was not a worm at all." Like Huxley, "the professor had not the least notion of allowing that things were true, merely because people thought them beautiful. ... The professor, indeed, went further, and held that no man was forced to believe anything to be true, but what he could see, hear, taste, or handle." A paragraph on "the great hippopotamus test" opens with the Professor, like Huxley, declaring "that apes had hippopotamus majors in their brains just as men have", but then like Owen presenting the argument that "If you have a hippopotamus major in your brain, you are no ape".
Then, presented with the awkward question, "But why are there not water-babies?", the Professor in Huxley's characteristic voice answered quite sharply: "Because there ain’t."

The Water-Babies was published in book form in 1863, and in the same year an even more satirical short play was published anonymously by George Pycroft. In A Report of a Sad Case Recently Tried before the Lord Mayor, Owen versus Huxley... the Great Bone Case, the vulgarity of the behaviour of Owen and Huxley is parodied as them being taken to court for brawling
in the streets and disturbing the peace. In court, they shout terms such as "posterior cornu" and "hippocampus minor". In giving evidence, Huxley states "Well, as I was saying, Owen and me is in the same trade; and we both cuts up monkeys, and I finds something in the brains of them. Hallo! says I, here's a hippocampus. No, there ain't says Owen. Look here says I. I can't see it he says and he sets to werriting and haggling about it, and goes and tells everybody, as what I finds ain't there, and what he finds is".

Man's Place in Nature

Huxley expanded his lectures for working men into a book titled Evidence as to Man's Place in Nature, published in 1863. His intention was expressed in a letter to Charles Lyell
Charles Lyell
Sir Charles Lyell, 1st Baronet, Kt FRS was a British lawyer and the foremost geologist of his day. He is best known as the author of Principles of Geology, which popularised James Hutton's concepts of uniformitarianism – the idea that the earth was shaped by slow-moving forces still in operation...

 which referred to the Monkeyana poem of 1861: "I do not think you will find room to complain of any want of distinctness in my definition of Owen's position touching the Hippocampus question. I mean to give the whole history of the business in a note, so that the paraphrase of Sir Ph. Egerton's line 'To which Huxley replies that Owen he lies', shall be unmistakable." Darwin exclaimed, "Hurrah the monkey book has come". A central part of the book provides a step by step explanation suitable for newcomers to anatomy of how the brains of apes and humans are fundamentally similar, with particular reference to both having a posterior lobe, a posterior horn, and a hippocampus minor. The chapter concludes that this close similarity between apes and mankind proves that the original definition by Linnaeus of the biological Order
Order (biology)
In scientific classification used in biology, the order is# a taxonomic rank used in the classification of organisms. Other well-known ranks are life, domain, kingdom, phylum, class, family, genus, and species, with order fitting in between class and family...

 of Primate
Primate
A primate is a mammal of the order Primates , which contains prosimians and simians. Primates arose from ancestors that lived in the trees of tropical forests; many primate characteristics represent adaptations to life in this challenging three-dimensional environment...

s was correct to include both, and mentions that an explanation of humans originating from apes is provided by Darwin's theory. The book also includes six pages of small print giving "a succinct History of the Controversy respecting the Cerebral Structure of Man and the Apes" describing how Owen had "suppressed" and denied what Huxley had now shown to be the truth regarding the hippocampus minor, posterior horn, and posterior lobe, describing this as reflecting on Owen's "personal veracity". Reviewers regarded the book as a polemic against Owen, and a majority of them sided with Huxley.
Sir Charles Lyell's authoritative Geological Evidences of the Antiquity of Man was also published in 1863, and included a detailed review of the hippocampus question which gave solid and unambiguous support to Huxley's arguments. In an attempt to refute Lyell's judgement, Owen again defended his classification scheme, introducing a new claim that the hippocampus minor was virtually absent in an "idiot". Then in 1866 Owen's book On the Anatomy of Vertebrates presented accurate brain illustrations. In a long footnote, Owen cited himself and the earlier literature to admit at last that in apes "all the homologous parts of the human cerebral organ exist". However, he still believed that this did not invalidate his classification of man in a separate subclass. He now claimed that the structures concerned – the posterior lobe, the posterior horn, and the hippocampus minor – were in apes only "under modified form and low grades of development". He accused Huxley and his allies of making "puerile", "ridiculous" and "disgraceful" attacks on his scheme of classification.

The publicity surrounding the affair tarnished Owen's reputation. While Owen's aim of finding an objective way of defining the uniqueness of humanity and distinguishing their brain anatomy in a qualitative way, not just a quantitative way, may be found laudable, his obstinacy in refusing to admit his errors in trying to find that difference led to his fall from the pinnacle of British science. Huxley gained influence, and his X Club
X Club
The X Club was a dining club of nine men who supported the theories of natural selection and academic liberalism in late 19th-century England. Thomas Henry Huxley was the initiator: he called the first meeting for November 3, 1864...

 of like minded scientists used the journal Nature
Nature (journal)
Nature, first published on 4 November 1869, is ranked the world's most cited interdisciplinary scientific journal by the Science Edition of the 2010 Journal Citation Reports...

to promote evolution and naturalism, shaping much of late Victorian science. Even many of his supporters, including Charles Lyell
Charles Lyell
Sir Charles Lyell, 1st Baronet, Kt FRS was a British lawyer and the foremost geologist of his day. He is best known as the author of Principles of Geology, which popularised James Hutton's concepts of uniformitarianism – the idea that the earth was shaped by slow-moving forces still in operation...

 and Alfred Russel Wallace
Alfred Russel Wallace
Alfred Russel Wallace, OM, FRS was a British naturalist, explorer, geographer, anthropologist and biologist...

, thought that though humans shared a common ancestor with apes, the higher mental faculties could not have evolved through a purely material process. Darwin published his own explanation in 1871 in the Descent of Man.

Modern relevance

In a talk about biological systematics
Systematics
Biological systematics is the study of the diversification of terrestrial life, both past and present, and the relationships among living things through time. Relationships are visualized as evolutionary trees...

 (classification) and cladistics
Cladistics
Cladistics is a method of classifying species of organisms into groups called clades, which consist of an ancestor organism and all its descendants . For example, birds, dinosaurs, crocodiles, and all descendants of their most recent common ancestor form a clade...

 given at the American Museum of Natural History
American Museum of Natural History
The American Museum of Natural History , located on the Upper West Side of Manhattan in New York City, United States, is one of the largest and most celebrated museums in the world...

 in 1981, the paleontologist
Paleontology
Paleontology "old, ancient", ὄν, ὀντ- "being, creature", and λόγος "speech, thought") is the study of prehistoric life. It includes the study of fossils to determine organisms' evolution and interactions with each other and their environments...

 Colin Patterson discussed an argument put in a paper by Ernst Mayr
Ernst Mayr
Ernst Walter Mayr was one of the 20th century's leading evolutionary biologists. He was also a renowned taxonomist, tropical explorer, ornithologist, historian of science, and naturalist...

 that humans could be distinguished from apes by the presence of Broca's area
Broca's area
Broca's area is a region of the hominid brain with functions linked to speech production.The production of language has been linked to the Broca’s area since Pierre Paul Broca reported impairments in two patients. They had lost the ability to speak after injury to the posterior inferior frontal...

in the brain. Patterson commented that this reminded him of "The Great Hippocampus Question" as recorded in fiction by Kingsley, and as in fact being a controversy between Huxley and Owen that "eventually as usual, Huxley won."
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