Commentary: The six biological inventions in Haldane's Daedalus
Department of Epidemiology and Public Health, The Queen's University of Belfast, Belfast, UK. E-mail: a.evans{at}qub.ac.uk
Accepted 20 March 2008
The great English Geneticist, John Burdon Sanderson Haldane (1892–1964), described six ‘biological inventions’ in his famous essay Daedalus. Three of these, and possibly a fourth, were straightforward domestications, the other two were more convoluted.
Haldane originally read ‘Deadalus, or Science in the Future’,1 as a paper to the ‘Heretics’ in Cambridge on February 4, 1923, and it was published later that year. ‘Daedalus’ is full of Utopian vision, following in the footsteps of many other works, such as Samuel Butler's ‘Erewhon’2 to which Haldane refers, and William Morris’ ‘News from Nowhere’.3
In ‘Daedalus’, before proceeding to ‘prophecy’, Haldane (Figure 1) wanted ‘... to examine very briefly the half dozen or so biological inventions which have already been made’.1 He then listed four ‘... which were made before the dawn of history’—the first of these was the domestication of animals. Haldane would have been well aware of some of the consequences of animal domestication for Public Health because his great-uncle, Sir John Burdon-Sanderson, was a major figure in Public Health in 19th century England. He was also a physician, physiologist and pathologist, and performed the post mortem on the Emperor Napoleon III in 1873.4 He was also the source of Haldane's christian names.
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Undoubtedly animal domestication confers many advantages, including providing a ready source of food, (either from meat or dairy products), transport and motive power, sources of clothing, bedding, upholstery, shelter, sport, vanity and companionship. There are in addition a number of attendant disadvantages.
In ‘Guns, Germs and Steel’, Jared Diamond reviews5 the reasons why domestication was successful in some species but not in others (Box 1), gives the dates of domestication of certain species and lists some of the diseases that we have come to share with them. In fact, as early as 1798, Edward Jenner, in the introduction to his ‘Enquiry into ... The Cow- Pox’ observed:6
The deviation of man from the state in which he was originally placed by nature seems to have proved to him a prolific source of disease. From the love of splendour, from the indulgence of luxury, and from his fondness for amusement he has familiarised himself with a great number of animals, which may not originally have been intended for his associates. The wolf, disarmed of ferocity, is now pillowed in the lady's lap [Jenner is quoting his Mentor, John Hunter,7 that the dog is the wolf in a degenerate state6]. The cat, the little tiger of our island, whose natural home is the forest, is equally domesticated and caressed. The cow, the hog, the sheep, and the horse, are all, for a variety of purposes, brought under his care and dominion.Evidently, Jenner was well aware of the link between animal and human disease, in this case cow pox and smallpox, two centuries ago. The earliest domesticate was ‘man's best friend’, the dog from the wolf, around 12 000 years ago,5 or as long as 100 000 years ago,8 which may explain why humans share 65 infectious diseases with dogs, and almost 300 with all domesticated animals.9 It is not even clear if cattle were the original hosts for smallpox,9 with counter-claims made for rodents as the source.10
A central theme of Diamond's in ‘Guns, Germs and Steel’5 is that domestication ultimately yielded agents of conquest. The effect of unfamiliar infectious disease in a susceptible population is vividly illustrated by the outbreak of measles (from rinderpest in cattle or distemper in dogs10), which devastated the Faeroe Islands in 1846. The Islands had been measles-free since 1781 and on its re-introduction, by a Faroese cabinet-maker who had visited some measles patients on a visit to Copenhagen, the attack rate was close to 80% with a very significant mortality.11
It must be pointed out, however, that in his introduction to ‘Infectious Disease and Host-Pathogen Evolution’, Dronamraju states12 that it was Haldane ‘... who first drew attention to the significant role infectious diseases have played in shaping our own evolution’.
In ‘Disease and Evolution’ in 1949, Haldane states that13 ‘... the struggle against disease, particularly infectious disease, has been a very important evolutionary agent ...’ and he eloquently observes that ‘Europeans have used their genetic resistance to such viruses as that of measles ... as a weapon against primitive people as effective as fire-arms’.
He suggested further that, in certain circumstances, parasitism will be a factor promoting polymorphism and may even tend to encourage speciation.12
According to Max Perutz,14 it was in this same article that Haldane first suggested that ‘... individuals with various red cell disorders might be protected from fatal infections with malaria’. Haldane had been first to estimate15 a human mutation rate and predicted that:14
... the high incidence of thalassaemia in malarial countries might have arisen by balanced polymorphism, heterozygotes for the disease having a selective advantage relative to homozygotes of either kind.Curiously, this is not mentioned in the paper that Perutz cites but, as Dronamraju explains,12 it was contained in a footnote in Italian. Even more curiously, Haldane subsequently credited the concept to his former student, Lionel Penrose,16 in a paper entitled, ‘Biological Possibilities for the Human Species in the Next Ten Thousand Years’.17 Haldane was probably being over-generous as he had sown the seeds of the concept as early as 1932.15
Pearce-Duvet lists10 the other major diseases to emanate from livestock as Pertussis, Tuberculosis, Taenid Tapeworms and Falciparum malaria. To bring this up to date, Bovine Spongioform Encephalopathy in cattle, which has been linked18 to Creutzfeldt-Jakob Disease in humans, must be added. There remains the imminent threat of an Influenza Type A pandemic.19 All through history these pandemics have tended to arise in the Far East where a large human population lives in close proximity with domesticated birds. Pandemics can be disseminated through the wild bird population by migrating birds19 before adapting to humans. Bird migration is another evolutionary legacy that has evolved over tens of millions of years. Jenner wrote about bird migration20 and he also helped unravel the secrets of the bizarre reproductive habits of the cuckoo.21
Apart from infectious disease, there are a number of other problems inherent in our relationship with domesticated animals. These include, particularly in the past, trauma inflicted by the use of animals in transport and warfare, but by far and away the most important effects today are nutritional. These have been amply reviewed elsewhere, but briefly, foods derived from animal sources are ‘energy-dense’, contributing to the global obesity epidemic and Type II Diabetes; moreover, the consumption of red and processed meat is associated with the risk of certain cancers.22 Diets high in saturated fats, overwhelmingly of animal origin, are associated with cardiovascular diseases23 and food preservatives, particularly the use of salt in the past, has had deleterious effects. Also, animals kept in captivity may have less healthy lipid profiles than their wild counterparts, e.g. salmon, unless their food is fortified with n-3 fatty acids24 (although they are definitely more contaminated), and often more likely to harbour infection, e.g. factory-farmed chickens, and to contain antibiotics and other contaminants.
The consumption of cow's milk, which requires adult-persistent lactase,8 and other dairy products provide a rich source of saturated fat, which the dairy industry has promoted for its ‘natural goodness’. After a quadruple by-pass graft, the attitude of the great cardiologist, Frank Pantridge, hardened to dairy products. In his autobiography, ‘An Unquiet Life’, he wrote:25
Common sense, however, would suggest that it might be unwise to consume large quantities of milk or milk products. Man is the only mammal which consistently consumes the milk of another species.For once Frank was wrong, the Bishnoi, are a community of desert people in Rajasthan in NW India. They known as India's original ecologists and their womenfolk have been known to suckle orphaned antelopes (Figure 2).26
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Haldane in Daedalus had this to say on the topic of milking:1
Consider so simple and time-honoured a process as the milking of a cow. The milk which should have an intimate and almost sacramental bond between mother and child is elicited by the deft fingers of a milk-maid, and drunk, cooked or even allowed to rot into cheese. We have only to imagine ourselves as drinking any of its other secretions, in order to realise the radical indecency of our relation to the cow.In a footnote, he gives details of the use of these ‘other secretions’ in Indian religious ceremonies, before taking a sideswipe at both Hinduism and Catholicism.
In 1949, Haldane wrote ‘... the possibility that the heterozygote is fitter than the normal must be seriously considered ...’27 Haldane was addressing himself to the issue of Hybrid Vigour (Box 2), which is advantageous in animal breeding. He explored the theory behind this in a paper written with Bartlett in 193528 where it is stated: ‘For a large number of purposes, both theoretical and practical, it is desirable to produce a population as heterozygous as possible’. The concept is brilliantly illustrated by the mule, whose failure to propagate itself, despite a healthy sexual appetite, conforms perfectly to ‘Haldane's Rule’.
What about the ethics of our relationship with domesticated animals, with whom we share a common ancestry and the vast majority of our genes? Christianity, at least, is comfortably disposed to it as the Bible furnishes man with complete dispensation:29
Then God said: "Let us make man in our image, after our likeness. Let them have dominion over the fish of the sea, the birds of the air, and the cattle, and over all the wild animals and all the creatures that crawl on the ground.The absurdity of our treatment of our fellow animals, with whom we have so much in common, has not been lost on everyone. It was refreshingly satirized in 1853 by the Irish physician and poet, James Henry, in ‘Man’s Universal Hymn’, which begins:30
‘The Lord's my God and still shall be,‘Man’s Universal Hymn’ coruscates along in this vein for another 157 lines.For a kind God he is to me,
And gives me carte-blanche to rob
His other animals, and to fob
For my own use their property,
So good and kind he is to me’.
Haldane's second ‘biological invention’ was the domestication of plants. According to Diamond, this began around 10 500 years ago in The Fertile Crescent of SW Asia with wheat, peas and olives.5 The health consequences were of course not nearly as marked as with the domestication of animals. Rather, it was the insidious changes in dietary intake, which occurred as humans swapped a hunter-gatherer existence for an agriculturally based one. In ‘Leviathan’, in a chapter entitled, ‘Of the Natural Condition of Mankind as Concerning their Felicity and Misery’, the 17th century English Philosopher, Thomas Hobbes, famously described the hunter-gathers’ existence as, ‘... solitary, poor, nasty, brutish, and short’.31 (The English novelist, Lawrence Durrell, described life in post-war England as ‘... solitary, poor, nasty, British, and short’.32) Hobbes came from a background in Alchemy and wanted the inscription on his gravestone to read,33 ‘This is the true Philosopher's stone’, but sadly his wish was not granted.
The hunter-gatherers’ diet, in common, to an extent, with the Mediterranean diet, would have been highly mixed,34 and probably, thanks to the large range of nutrients, trace elements and anti-oxidants, coupled with high levels of physical activity, made for a healthy lifestyle, despite Hobbes’ analysis. In Europe by the Late Mesolithic period (4000 BC) wild plant food husbandry was already supplanting the opportunistic use of plants for food.35
Not all plant domestications were conducive to good health: both tobacco and coca were originally domesticated in the Andes.36 Another problem with propagating plants is getting them to breed true—and this is one of the main aims of domestication. In 19th century America, traders selling maize seed that was the result of a hybrid cross between two true-breeding inbred varieties had a characteristic, not disclosed, with special commercial value. Hybrid corn was not true breeding-breed, so it did not produce reliable seed.37 Farmers were forced to buy fresh seed from the trader the following year, as is the case with terminator seed today.
Grafting was one strategy to overcome this problem, as is commonly employed with fruit trees. Another was vegetative propagation, and today, two of the world's great staple foods, potatoes and bananas, are produced in this way. Because both plants are cloned, they lack biological diversity, and so are susceptible to disease. The fungi that attack them are genetically evolving to bypass the plants’ defences. It is a perfect example of Darwin's Natural Selection in action.
In the case of potatoes the result is blight caused by the fungus Phytophthora infestans that from 1845–49 resulted in the Irish Potato Famine.38 Burgundy mixture was the first anti-fungal agent to be used: it was developed after the serendipitous observation in 1882 that a mixture of copper sulphate and hydrated lime, sprayed on French vineyards to stop the pilfering of grapes, prevented mildew.39
Haldane cited13 the cultivated banana clone, Gros Michel, as a beautiful example of the danger of homogeneity. It is now extinct having long ago lost its one-sided battle with Fusarium cubense. The current commercial clone, Cavendish is now under attack on several fronts, especially from the Black Sigatoka Fungus, Mycosphaerella fijiensis, and looks set to go the same way as Gros Michel.40 The only defence is to step up levels of insecticides which bring their own drawbacks.
The combination of animal and plant domestication led to the establishment of agriculture. What is clear is that with the advent of domestication, the range of plants and animals consumed began to fall. Diamond estimates that of 148 species weighing 45 kg or more, only 14 were domesticated. Of over 200 000 species of higher plants, only about 100 were usefully domesticated8 and the range of grass seeds eaten today is tiny in comparison to the past.
In 1962 Neel observed41 ‘... during the first 99% or more of man's life on earth, while he existed as a hunter and gatherer, it was either feast or famine’. Neel's ‘Thrifty’ genotype hypothesis stated that those who could store energy more easily in a ‘feast’ were better able to withstand famine. The diet that humans have consumed since the introduction of agriculture, has been characterized by a foodstuffs richer in simple carbohydrates, saturated fats and latterly in more calories and salt, and are lower in fibre, complex carbohydrates, calcium and unsaturated fat. According to Neel,41 the ‘Thrifty’ genotype, thanks to this dietary ‘progress’, has rendered mankind susceptible to Diabetes Mellitus. Moreover, in contrast to the hunter-gatherer state, the glucose-mobilizing effect of adrenalin is no longer followed by physical activity.
Diamond gives a good account of the tensions that must have arisen between the hunter-gatherers and the new agriculturalists.8 It seems to have been much like the American Wild West when the middle classes arrived to displace the cowboys. In ‘The Conquest of Happiness’, the English philosopher, Bertrand Russell commented42 that ‘... with the coming of agriculture life began to grow dull, except, of course, for the aristocrats, who remained, and still remain, in the hunting stage’. The settled agriculturalists naturally won out because they had the enormous advantage of abundant food production, which could support a burgeoning population.
Agriculture, as Vallee has pointed out, very soon brought its own problems:43
The creation of agriculture led to food surpluses, which in turn led to ever larger groups of people living in close quarters, in villages or cities. These municipalities faced a problem that still vexes, namely, how to provide inhabitants with enough clean, pure water to sustain their constant need for physiological hydration. The solution, until the 19th century, was non-existent. The water supply of any group of people rapidly became polluted with their waste products and thereby dangerous, even fatal, to drink. How many of our progenitors died attempting to quench their thirst with water can never be known.Vallee makes a strong case that one of the answers to this problem was alcoholic drink, which provided a safe source of drinking water.43
This brings us to Haldane's third ‘biological invention’, which was ‘... the domestication of fungi for the production of alcohol’. Diamond totally ignores this in ‘Guns, Germs and Steel’,5 although by his own definition domestication involves:8
... a species bred in captivity and thereby modified from its wild ancestors in ways making it more useful to humans who control its reproduction and (in the case of animals [and domesticated Saccharomyces cerevisiae]) its food supply.Haldane was perfectly correct and other authorities would agree with him,44,45 with the same applying to yeast used in baking.46 Diamond does subsequently recognize8 the contribution of the ‘... consumption of large quantities of nutritionally important beer’, over the last several thousand years but does not acknowledge this to be the result of domestication. In his quote on milking (see above) Haldane refers to letting milk rot into cheese. This is a complex process involving lactobacilli, streptococci, and sometimes fungi (for veining and lacunae). Aroma, that crucial element of taste, is important—apparently the French recommend their Camembert in the words of their poet Léon-Paul Fargue: ‘Les pieds de Dieu’.47
It seems that beer has been brewed since the early Mesolithic period and wine was first produced in Armenia about 6000 BC,43 but human exposure to alcohol must predate this considerably. All types of life would have found an ability to metabolize alcohol an advantage when fruit ‘spoiled’, and fermented in the wild. Aristotle refers to ‘... a gnat produced by larvae engendered in the slime of vinegar’.48 This must have been Drosophila (dew lover), which has so heroically contributed to our knowledge of genetic mechanisms. Sturtevant regretted that its earlier name, Oinopota (wine drinker) was not kept.48
In any case, Vallee's observations41 concerning wine and beer as a safe source of drinking water were not new. Drummond, in ‘The Englishman’s Food’, had this to say about ‘small beer’: ‘It had the advantage of being a good deal safer and probably more palatable than most of the drinking water available’.49 In this respect, the absence of cases of disease in the Brewery during the Golden Square Cholera outbreak in 1854 should have been most informative.50 Lastly, Nathaniel Hulme advised the botanist, Joseph Banks, to take turpentine and molasses with him on the Endeavour's voyage to the South Pacific in 1768:51
... in order to brew Beer with, for your daily drink, when your Water becomes bad ... In this case you will want Yeast and the manner for preserving this at sea [reference to an Appendix]... So small a quantity of Molasses as two gallons, or two Gallons and a half are said to be sufficient for making a Hogshead of tolerably good beer ... I find by Experience that the smell of stinking water will be entirely destroyed by the process of fermentation.
Alcoholic beverages had another important advantage, they represented a way of preserving calories over long periods, thus beer is still known as ‘Liquid Bread’ in central Europe. It was also an intoxicant, which has fuelled warfare throughout history, and popular, in some cases too popular with those who imbibe it. In his Nobel Prize Acceptance Speech in 1950, Bertrand Russell observed:42
When white men first effect contact with some unspoilt race of savages, they offer them all kinds of benefits, from the light of the gospel to pumpkin pie ... What they really value ... is intoxicating liquor, which enables them for the first time in their lives, to have the illusion, for a few brief moments, that it is better to be alive than dead.
One thing that persists, for whatever reason, is that alcohol in moderation and drunk fairly constantly, seems to confer cardioprotection.52
Another valuable observation made by Valee was that:43
... genetics played an important role in making Asia avoid alcohol: approximately half of all Asian people lack an enzyme53 necessary for complete alcohol metabolism, making the experience of drinking quite unpleasant.
This unpleasant experience, related to gene morphism,53 is immediate as opposed to the ‘morning after’, which Westerners endure. For at least 2000 years tea has been the staple beverage in the East, with the water rendered safe by boiling.43
Haldane's fourth ‘biological invention’ comes as a surprise as it concerned human sexuality, and one which he believed:1
... was of more ultimate and far-reaching importance ..., since it altered the path of sexual selection, focussed the attention of man as a lover upon woman's face and breasts, and changed our idea of beauty from the steatapygous Hottentot to the modern European, from the Venus of Brassempouy to the Venus de Milo. There are certain races which have yet to make this last invention.
The first three inventions have been domestications but Haldane has been careful to define what he means by an ‘invention’ at the start of the paragraph:1
By a biological invention I mean the establishment of a new relationship between man and other animals or plants, or between different human beings, provided that such relationship is one which comes primarily under the domain of physics, psychology or ethics.
This is all very well but seems reminiscent of Procrustes ‘the Stretcher’, a character in Greek Mythology who fitted his victims in an iron bed, either by stretching them or cutting pieces off their legs, before robbing them.54 Theseus slew Procrustes by adapting him to fit the iron bed.
Beginning in 1892, a series of carved artefacts were recovered from a cave, the grotte du Pape, at Brassempouy in SW France.55 The most famous is La dame à la capuche comprising a woman's hooded head. Several small figures and other items were unearthed along with a mammoth ivory figurine fragment known as La Poire (the pear). It consisted of ample, well-rounded buttocks and a prominent mons veneris, which became the only Brassempouy figure to be labelled a Venus: La Vénus de Brassempouy. According to White,55 the name was almost certainly a reference to the Hottentot Venus, Saartjie Baartman (see subsequently). It was the commemoration of the centenary of her death, and the African frenzy of la belle époque, which caused the term to be applied to the most robust of Palaeolithic figurines. This one figurine was steatopygous, most of the others were callipygian.
The figurines of Brassempouy date from around 25 000 BC. This date would fit in with them having been fashioned by the people who succeeded the Neanderthals about 30 000 years ago,56 who are named after another cave in SW France at Cro-Magnon, in which some of their bones were first discovered in 1868.57 The Cro-Magnon people's genes are still extant in modern Europe. As Steve Jones memorably observed:58 ‘Most people would change seats if Cro-Magnon ... sat next to them on the tube, but would change trains if Neanderthal did the same thing.’ One possible source of inspiration for La Poire is that some of the Cro-Magnon people originated from the very south of Africa and included some of Bushman origin. Another possibility that merits consideration is that, because it is now realized that Neanderthals and Cro-Magnon coexisted, arguably without interbreeding (see Box 2), for thousands of years,56 the Venus was based on a Neanderthal female. Neanderthals were short and stocky (The Hottentot Venus was less than four feet, seven inches in height59) and the date for the extinction of Neanderthals is constantly being brought forward.60
The Venus de Milo, on the other hand, was found on the Island of Melos and dates from the first century BC. She represents idealized beauty, to be viewed from the side with her missing arms outstretched towards Cupid.61 (there is an apochryphal story that a Californian newspaper magnate had the Venus de Milo delivered to his mansion, but when she arrived her arms were broken, so he sued the railroad company, and won).
The Hottentot Venus was in contrast a very real person (Figure 3): she has been the subject one book in French,62 and two recent books in English: the first a novel,63 the second a splendid biography by Rachel Holmes.59 Saartjie (anglicized to Sarah) Baartman was born in 1789, into a South African herding tribe known as the Eastern Cape Khoisan. Thanks to colonial war she was orphaned and widowed before being forced aboard a ship in Cape Town, along with a decomposing giraffe skin, and taken to England in 1810. Her abductors were, Hendrik Cesars, a brother of her Custodian, and William Alexander Dunlop, a military surgeon, whose name suggests he was Scottish (his Return of Service record was destroyed at the National Archive in Kew by a World War II bomb). Saartjie was exhibited by the two men in London as ‘The Hottentot Venus’.59
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Her initial celebrity was on account of her steatopygous rear (she was christened62 ‘big bum’ in London), but she became an accomplished and charming performer and, despite being vilely exploited, she danced, sang and played the guitar. The men attempted to sell Saartjie, but being unsuccessful decided to manage her themselves. She was exhibited in Piccadilly and feted and dined by the great, including the Marquis of Queensbury. To cut a long story short, Dunlop died in 1812,64 and eventually Saartjie ended up, after various illnesses, in Paris where she died in the cold winter of 1815.59 As if things had not been bad enough for Saartjie in life, what was to follow in death was grisly in the extreme.
Plaster casts were taken of Saartjie's corpse, which was then dissected by Georges Cuvier: first her brain was removed and preserved in a jar; then her ‘apron’ (mons veneris and genitalia) was modelled in wax, removed and bottled; finally her skeleton was disarticulated. After Cuvier's death in 1832, and before 1860, Saartjie's body parts went on display at the National Museum of Natural History and subsequently at the Musée de l’Homme until the 1970s. After the end of Apartheid in 1994, Nelson Mandela started negotiations62 for reclaiming the ‘cultural property’ pertaining to Saartjie. Her remains were given a state funeral at Hankey, Eastern Cape, in the Republic of South Africa, on August 9, 2002.65 She had been overseas for nearly two centuries.
Saartjie became a cause célèbre in Europe at the start of the 19th century. She was a victim of racial and sexual stereotyping but she also became a leader of fashion: Holmes ascribes the fashion of the bustle later in the 19th century to her—coyly named the ‘tournure dress improver’ in England, and ‘le faux-cul’ (false bum) in France. As the Morning Herald put it:59
‘Though Venus, of old,In the light of this, Haldane's fourth ‘biological invention’ appears all the more bizarre. Steatopygia may not have been attractive to him but is probably irresistible to Khosian males, which after all is the important thing. Steatopygia probably signals a potential mother who is likely to be successful at child rearing under arid conditions. Sexual characteristics will be successful if they achieve propagation and that is natural. There are, however, examples in nature where sexual characteristics are imitated to ensure propagation (Box 3).By records, we’re told,
Excited the praise of mankind;
Our Hottentot, still,
Let her die when she will.
Will not leave her equal behind’.
Another factor that should be considered is that in ‘Daedalus’, Haldane, still a relatively young man, set out to be controversial. His description of Einstein as ‘... the greatest Jew since Jesus’,1 was not intended to leave religious feathers unruffled, and even the alliteration is mildly shocking. According to Dronamraju, Haldane:66
... immensely enjoyed the sensation [Daedalus] caused and took great pleasure in expounding his views loudly at every opportunity, quickly clearing tea shops of the more susceptible church-goers.Also it was not that long after World War I and rebellion against Edwardian repression was in the air.
‘Daedalus’ was the first airing of Haldane's views on Eugenics and came just after his move to Cambridge. Dronamraju plausibly suggests66 that Haldane's social milieux had something to do with this. For some years, Haldane and his sister Naomi (later Lady Mitchison, the writer) had been drawn into wider intellectual circles where such ideas had gained currency. Through the Huxleys (Aldous, another Utopian novelist,67 who cribbed Haldane's ‘biological inventions’, and his brother Julian) they:66
... were drawn closer to the intellectual centres of Bloomsbury, particularly the left leaning group which found a generous hostess in Lady Ottolline Morrell.The circle included Bertrand Russell, who in 1924 had responded to ‘Daedalus’ with ‘Icarus, or the Future of Science’,68 which adopted a gloomier outlook about the topic. Russell, Julian Huxley and DH Lawrence had affairs with Ottolline, and Lawrence based ‘Lady Chatterley’s Lover’ on the experience. Other guests of Ottolline included Augustus John, WB Yeats, David Cecil, TS Eliot, John Maynard Keynes, Wyndham Lewis and many more.66
Ottolline has been the subject of biography, most recently by Miranda Seymour.69 Ottolline's wardrobe, comprising 600 garments, now adorns the Museum of Costume in Bath.70
Although only on the periphery of these circles, Haldane must have encountered a heady mix of Suffragettism and liberated sexual mores. One wonders how much his ‘fourth invention’ was influenced by these perspectives, or was he wryly poking fun at them?
Haldanes last two ‘biological inventions’ refer to his ‘own day’.1 The first of these is ‘bactercide’ and he is probably thinking of Ehrlich's Salvarsan, although Penicillin was to come along shortly afterwards and can be viewed as another fungal domestication. The last invention was ‘artificial control of conception’. This is an important plank of Eugenics but, also, Marie Stopes, the great advocate of birth control,66 was involved in a libel case at the time.
Commenting on the first four inventions, Haldane noted1 that they all had had a profound emotional and ethical effect: ‘... there was not one which had not formed the basis of a religion’.1 He was not so sure about contraception forming the basis for a religion, but knew that it was impossible to keep religion from discussing it. He concluded that every biological invention is a perversion:1
There is hardly one which, on first being brought to the notice of an observer from any nation which had not heard of their existence, would not appear to him as indecent and unnatural.Certainly, there are many aspects of animal domestication that may strike some people as unpalatable, and the notion that the modern European woman is a biological invention is oddly repugnant. Never-the less, this review has covered only nine sentences of ‘Daedalus’. These, along with the rest of the text, still bear testimony to Haldane's unique mind.
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 Top Appendix AcknowledgementsReferences |
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| Box 1 Zebra domestication Diamond recognizes at least six groups of reasons for a failure to domesticate animals: finicky diet, growth rate, problems of captive breeding (too shy), tendency to panic, social structure and nasty disposition. Diamond states,1 ‘Alas, zebras become impossibly dangerous as they grow older. Zebras have the unpleasant habit of biting a person and not letting go’ (rather like politics where I live). ‘They thereby injure even more American zoo-keepers each year than do tigers!’ A comparison of rates would have been appropriate here, but, although a superb evolutionist, Diamond does not burden himself with the complexities of the Floating Denominator. He continues, ‘Zebras are also virtually impossible to lasso with a rope—even for cowboys who win rodeo championships by lassoing horses—because of their unfailing ability to watch the rope noose fly toward them and to duck their head out of the way. Hence it has rarely (if ever) been possible to saddle or ride a zebra, and South Africans’ enthusiasm for their domestication waned’. He does state, however, that Lord Walter Rothschild drove through the streets of London in a carriage pulled by zebras (Figure 1.1).1 Walter's father, Nathaniel, was the leader of Zionism in Britain and in 1917, 2 years after Nathaniel's death, the Balfour Declaration, which was to set up the state of Israel was sent to Walter.2 Walter was highly eccentric, at times inarticulate, but a passionate devotee of natural history, founding his own zoological museum at Tring, north of London. According to his niece Miriam Rothschild2 (The Queen of Bee Research), ‘It was a grand year ... in November 1894 the live zebras arrived at Tring and were, amid great excitement, put into loose boxes in the carriage stables and Walter began the business of breaking them in’. In fact, Tegetmeier and Sutherland record that,3 Walter ‘... placed three in the hands of a very careful breaker’ (Walter's access to unlimited funds was a bone of contention with his fellow naturalists). Miriam Rothschild continues,2 ‘... this proved no mean task for the zebras objected strongly to harness and bridles and he had to devise a method of letting down their collars from the ceiling ... Walter first accustomed them singly to a small trap, but he was eventually able to drive them down Piccadilly as a four-in-hand—three zebras and a small pony was the usual combination—and into the forecourt of Buckingham Palace’. The Natural History Museum (London) photograph's legend states that the carriage was ‘drawn by four zebra’ but, sure enough, the far front equid was a pony. This was akin to the American ‘Bell Mare’, which was a mare with a bell round her neck who was willingly followed by the mule-train, wherever she led.4 Apparently, the children were told ‘... that the zebras’ camouflage was so good that halfway down the street they seemed to vanish, leaving Walter bowling along Piccadilly in a horseless carriage ... Nattie (Nathaniel) was not pleased about the expedition to the Palace which, in his opinion, invited disaster. Walter himself admitted it was rather risky—his heart was in his mouth when Princess Alexandra tried to pat the leading zebra’.2 Just as if to prove Diamond right, ‘Later one of his (Walter's) zebras seriously injured a groom. His father gave the man a pension for life. The man eventually died from his injuries’.2 Despite the experience of Lord Walter and others, and the gloomy character reference given by Diamond, according to Tegetmeier and Sutherland, Burchell's zebras were being used in the 1890s as coach teams in the Transvaal.3 There is a picture to testify to this in their book, also a charming photograph of Burchell's zebra harnessed to a four-in-hand and a Cape cart, and a picture of a demure young lady riding a zebra, side-saddle (Figure 1.2). Apparently, Burchell's zebra is ‘... easily broken to harness, and readily becomes a domesticated animal’. And, ‘It is only recently that the Cape Colonists have arrived at the conclusion that Burchell's zebra is a desirable beast of draft and of burden’.3 They were also ‘... free of that scourge of South Africa commonly called ‘horse sickness’ (a lethal form of distemper4) and tsetse fly’. Tegetmeier and Sutherland concluded ‘... it has been found that the idea as to the impossibility of taming and breaking the zebra is a perfect myth’.3 In 1893 Burchell's zebras were on sale in the Cape at prices varying from £10 upwards and several had been imported into Great Britain, presumably this was Lord Walter's source. A Captain Lugard wrote, ‘Some years ago (1888) I advocated experiments in taming the zebra and I especially suggested that an attempt should be made to obtain zebra-mules by horse or donkey mares. Such mules, I believe, would be found to be excessively hardy, and impervious to the fly and to climatic diseases’.3 In fact, ‘Lord Walter in his combination of a zoo and museum, successfully mated a pony and a zebra ... but unfortunately for science and future zoologists it was not considered practical’.2 Lord Walter was not the first to employ an eccentric mode of transport in London. In the late 18th century, John Hunter used to drive into central London from Earls Court on a cart pulled by three Zebus (Asian Buffaloes)5 and, ‘Early in this century (19th), a pair of hybrids, bred between horse and Burchell's zebra, were driven about London in the service of the Zoological Society’.3 References 1Diamond J. Guns, Germs and Steel: A Short History of Everybody for the Last 13 000 Years. London: Johnathan Cape, 1997. 2Rothschild M. Dear Lord Rothschild: Birds, Butterflies and History. London: Hutchinson, 1983. 3Tegetmeier WB, Sutherland CL. Horses, Asses, Zebras, Mules, and Mule Breeding. London: Horace Cox, 1895. 4Clutton-Brock J. Horsepower: A History of the Horse and the Donkey in Human Societies. London: Natural History Museum Publications, 1992. 5Moore W. The Knife Man: The Extraordinary Life and Times of John Hunter, Father of Modern Surgery. London: Bantam Press, 2005.
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| Box 2 Hybrid vigour Aristotle knew about mules so mule-breeding has a long history. He supposed that other animals were species hybrids, e.g. the giraffe was the result of a cross between the camel and the leopard. According to him such cross-breeding took place around water holes in Libya.1 Mules, despite suffering from a bad press, are a superb example of Hybrid Vigour: they are hardier than horses, can carry more and can get by on poorer fodder.2 Tegetmeier and Sutherland record3 that, ‘... if properly handled, the mules will come down on the knees at a pull as many times as you ask them’. Hybrid vigour, as Haldane highlighted,4 results from the increase in heterozygosity caused by the interbreeding of two genetically different individuals. Because of their hybrid vigour the mule was the favourite pack animal for long distance transport for at least 3000 years until the invention of the steam engine, so their economic contribution to modern civilization cannot be overestimated. As well as greater vigour, mules had an effective working life almost double that of the horse.3 Moreover, mules have a sense of fun and mischief, and if well treated, are intensely loyal, following their army handlers around and nuzzling them like huge dogs.5 Their handlers would become so fond of their charges that they would forego leave. They were very popular in America and in France, where a particularly large variety was bred in the old province of Poitou,3 but mules were traditionally viewed with less enthusiasm in Britain. Quite simply, the mule was yesterday's tractor and of great military significance well into the last century. They were introduced by the British Army to good effect in World War I, and in World war II 5563 mules and horses were air-lifted into Burma, but in order that they would not betray their positions to the enemy, each one had its vocal cords cut.6 Sadly, the British Army's ‘Mule School’ was closed in 1975. The various species of horses, despite their marked similarities of form differ greatly in their numbers of chromosomes.5 Fusion or fission of chromosomes may be a major mechanism of speciation in mammals, and these differences may, therefore, carry great evolutionary significance. Horses have 64 chromosomes, in common with the Grevy zebra. The Burchell or ‘plains’ zebra has 44 and the mountain zebra 32. The sterility of the hybrids does not mean that they lack either internal or external reproductive organs, or sexual drive,6 and, to make it practicable, it is necessary to geld the male mule in the same way as the stallion horse.5 The male hybrid is unable to produce spermatozoa because the two kinds of chromosomes of its parents are unequally matched for the chromosome number of the equids.5 The female hybrid, although she may come into oestrus and occasionally may accept the male, will only rarely produce viable offspring. This had been noted by John Hunter in 1787, which he attributed to, ‘... a degree of monstrosity in the organs of the mule’, despite ‘... the copulation of Mules being very frequent’.7 Infertility is certainly usual in male mules but female mules have produced young when mated with a stallion.8 In 1922, Haldane published9 an interesting generalization, which became known as ‘Haldane's rule’, It states that: ‘When in the F1 offspring of two different animal races one sex is absent, rare, or sterile, that sex is always the heterozygous one’. The problem in mules is that during Anaphase there is chaotic cell division due to the fact that the mule has not received the same number of chromosomes from each parent.8 This is compounded in male mules by the fact that although the genetic material contained on the Y-chromosome may be nearly identical between closely related species, there is large variation in the packaging of the DNA between species. The same mechanisms could explain why Neanderthal mtDNA sequences fall outside the variation of modern humans,10 despite having 99.5% genetic homology,11 demonstrating that no cross breeding occurred, or if it did, that the offspring were sterile. The mule is a progeny of a male donkey (jack or jackass) and a female horse. A ‘hinny’ or ‘jennet’ is the progeny of a male horse and a female donkey (jenny). These were much bred in Ireland, where in 1893 there were 200 000 donkeys.3 Hinnies were easier to breed on a large scale because one horse could be used for a large donkey population, a strategy that was adopted in 1892 by the Congested districts Board (a system to relieve destitution) in Ireland.3 Donkeys sometimes have linear markings, typically down the back and across the withers, fabled in Catholic Ireland to be the Sign of the Cross because the donkey had borne Christ. Animals can be reluctant to mate with a member of another species. As usual man has been most ingenious in hoodwinking (a word borrowed from falconry) his fellow animals. Crosses between asses and Burchell's zebra were tried in South Africa, and these occasionally occur in the wild.2 Hamilton Smith was much intrigued by the interbreeding of different species of equids and wrote that: ‘Already, in the time of Bouffon (1707–88) the idea of producing mules from the striped species of Equidae had occurred.12 Lord Clive in experiments to effect this purpose, had found it necessary to deceive a female zebra by painting a male ass with hippotigrine stripes’. This is the most convincing piece of evidence I know of that cosmetics really work. Other examples of the many subterfuges employed in mule breeding are described in Mr C L Sutherland's Memorandum on Mule Breeding prepared for the use of the Government of India.3 The 19th Century Irish-American Politician, Ignatius Donnelly, famously described the Democratic Party as like a mule: ‘It has neither pride of ancestry nor hope of posterity’.2 This is a good description of the male mule, at least. References 1Sturtevant AH. A History of Genetics. New York: Cold Spring Harbor Press, 2001. 2Clutton-Brock J. Horse-power: A History of the Horse and the Donkey in Human Societies. London: Natural History Museum Publications, 1992. 3Tegetmeier WB, Sutherland CL. Horses, Asses, Zebras, Mules, and Mule Breeding. London: Horace Cox, 1895, pp. 63–166. 4Haldane, JBS. The rate of mutation of human genes. Proc Eighth Int Cong Genet, 1949;Hereditas 35:267–73. 5Gould SJ. What, if anything, is a zebra? Nat Hist 1981;90:6–12. 6Cooper J. Animals in War. London: Corgi Books, 2000. 7Hunter J. Observations tending to shew that the wolf, jackel, and dog, are all the same species. Philos Trans R Soc 1787;LXXVII (Part II):253–66. 8Lloyd-Jones O. Mules that breed. J Hered 1916;7:494–502. 9Haldane JBS. Sex ratio and unisexual sterility in hybrid animals. J Genet 1922;XII:101–09. 10Krings M, Stone A, Schmitz RW, Krainitzki H, Stoneking M, Pääbo S. Neanderthal DNA sequences and the origin of modern humans. Cell 1997;90:19–30. 11Noonan JP, Coop G, Kudaravalli S et al. Sequencing and analysis of Neanderthal genomic DNA. Science 2006;314:1113–18. 12Hamilton Smith C. Horses, in the naturalist's; library. In: Jardine W (ed.). Mammalia. XX. Edinburgh: W H Lizars, 1845.
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| Box 3 Sexual deception Mimicry is rife in nature but one of the most outrageous subterfuges is perpetrated by the bee orchid (Figure 3.1). In 2006, a subheading in a British newspaper read: ‘Spectacular Orchids Double Due to Global Warming’.1 It continued, ‘The bee orchid, one of the most spectacular of Britain's plants, tricks bees into "pseudo-copulation" to spread its pollen’. The mimicry is so good that even the bees’ hairs are represented (Figure 3.2). Some flowers mimic a potential female mate visually but the key stimuli are often chemical and tactile. This rejoices in the name of Pouyannian mimicry,2 after the person who first described it in 1917.3 The orchid also secretes chemicals from glands called osmophores, which are indistinguishable from the bee's natural pheromones. This relationship between the bee orchid and bees must have evolved over millions of years. Mutualism implies that both parties are getting something out of the relationship: the bee, attracted by sight and chemicals, derives nectar and pollen from the orchid and is seduced into ‘pseudo-copulation’, whereby the orchid gets pollinated and has its pollen disseminated. The Belfast artist and botanist, Raymond Piper, set out many years ago to paint all the orchids that are native to Ireland,4 of which there are around 35. Raymond sadly died in 2007,5 but reproduced above, with his kind permission, is his exquisite water colour of Ophrys apifera. References 1McCarthy M. Spectacular orchids double due to global warming. The (London) Independent. April 24, 2006, 14–15. 2Pasteur G. A classificatory system of mimicry systems. Ann Rev Ecol Syst 1982;13:169–99. 3Pouyanne M. La fécondation des Ophrys par les insectes. Bull Soc Hist Nat Afr Nord 1917;8:1–2. 4Longley M. The orchid man. The Irish Times Magazine, January 11, 2003, 10–12. 5Clements P. Obituary: Raymond Piper. The Guardian, September 17, 2007, 38.
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TopAppendixAcknowledgementsReferences |
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Thanks are due to Professor Dan Bradley for his advice on the Y-chromosome, to Professor Krishna Dronamraju for his kind advice and generosity in furnishing Figure 1, and to the late Raymond Piper for supplying Figures 3.1 and 3.2. Special gratitude is appropriate to Joe Clint for his tenacity with the References, and to Deva Evans for her expert assistance with the Figures.
Conflict of interest: None declared.
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