Wilmot James is fascinated by science: by the people who do research, by their results and by the implications of their efforts

for creating a just society. In this book, he discusses ideas, people and history and does it with elegance, skill and a deep human sympathy.   – David Baltimore, California Institute of Technology Wilmot James has produced a book which will appeal to readers in South Africa and farther afield. The topics dealt with include the genetics of skin colour; human genetic diseases; the interaction between the humanities and science and the lives of two South African intellectuals, Eddie Roux and Eugène Marais. James writes with flair and style, and emerges as a fine populariser of science – a science unashamedly rooted in, and of particular relevance to, southern Africa. – Trefor Jenkins, University of the Witwatersrand   From genes to geology, medicine to music, bacteria to beauty, Wilmot James sheds light on a cornucopia of ideas. Skilfully weaving the personal and the global, the scientific and the political, his thinking is rooted in South Africa, but with a worldwide vision. At the core is the triumph of science as enlightenment and liberation, a potent force for the public good. You will relish Nature’s Gifts. – Helena Cronin, London School of Economics.

Nature’s Gifts
Why we are the way we are

WILMOT JAMES

Published in South Africa by: Wits University Press 1 Jan Smuts Avenue Johannesburg 2001 http://witspress.wits.ac.za Copyright © Wilmot James 2010 First published 2010 ISBN 978-1-86814-515-7 All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, without the written permission of the publisher, except in accordance with the provisions of the Copyright Act, Act 98 of 1978. Edited by Pat Tucker Cover design by Hybridesign Book design and layout by Hybrid Creative Printed and bound by Ultra Litho (Pty) Ltd.

I dedicate this book to the X chromosome so abundant in my life: my wife Delecia Forbes and daughters Gabriele and Isabella, and of course, my mother, Shelma James.

Contents
Acknowledgements viii 1. Of what use are genomes? 1 2. Reading Genes 21 3. Skin Colour 41 4. Blood 61 5. Senses and Sensibilities 81 6. Ways of Learning 101 7. Edward Roux – Communist and Botanist 121 8. Science in the Life of Eugène Marais 141 Notes 162 Bibliography 177 Index 188

Acknowledgements
Nature’s Gifts is based on a series of lectures I gave at the Human Sciences Research Council (South Africa), Duke University, California Institute of Technology, University of California at Berkeley, University of Wisconsin at Madison, Massachusetts Institute of Technology, University of Cape Town, University of the Witwatersrand, University of Pretoria, University of London, University of Texas at Brownsville, the Open University at Milton Keynes and Monash University of Australia. It also draws on the more than 100 columns I have written for the Cape Times on science-related issues in the past four years. I am very grateful to David Baltimore and the Faculty of the California Institute of Technology’s Humanities and Social Sciences Division for the invitation to spend the 2003-2004 academic year there as the Moore Visiting Professor, during which time I started the initial research for this book. Thanks to Steve Sturdy of the University of Edinburgh’s ESRC Genomics Policy & Research Forum for a visiting fellowship undertaken in 2007, during which time I refined some of the work on skin colour pigmentation. While there it was my privilege to meet Jonathan Rees, the first person to describe the melanocortin receptor 1 or MCR1 gene. Raj Ramesar, Professor and Head of the Division of Human Genetics at the University of Cape Town, persuaded the university to appoint me as an Honorary Professor in his division in 2005; a novel move to have a sociologist based in genetics. It is my privilege to have Raj as a colleague, someone who shares with me great enthusiasm for what Edward O Wilson once called ‘consilience’, which is the navigation in search of answers through the disciplines of science, social science and the humanities. Raj and I introduced the Darwin Seminars at the University of Cape Town and we have had four years of developing Cape Town’s only public forum for presenting innovations in evolutionary biology, put together so ably by Beryl Eichenberger.

At the University of Pretoria, where I am an Honorary Professor, Janis Grobbelaar provided a platform for taking the meaning of genes to sociologists. David Chidester, Trefor Jenkins, Himla Soodyall, Janis Golding, Nhlanhla Dlamini, Udo Shucklenk, Heather Sherwin and Roger Trythall, fellow directors at various times at the Africa Genome Education Institute, were my sounding boards and advisors. Neville Alexander, David Baltimore, Koos Bekker, Jo Ann Chataway, Ampie Coetzee, Cheryl Douglas, Janis Grobbelaar, Michael Kahn, Trefor Jenkins, Jeffrey Lever, Nicoli Nattrass, Steve Olsen, Raj Ramesar and Mamphela Ramphele commented on aspects of the chapters.The publisher’s two anonymous reviewers made extensive comments, prompting serious surgery to many of the chapters, and the copy editor, Pat Tucker, recommended major rewriting and a necessary gutting of some verbose parts of the manuscript. It made for a much better book. Carryn Smit helped with the laborious business of the notes. Himla Soodyall provided the informative maps. Veronica Klipp is a writer’s dream of a publisher – supportive and caring all along the way. I am, of course, responsible for any errors. The Charles Stewart Mott Foundation provided support for the completion of the manuscript and Naspers’s Ton Vosloo and Koos Bekker committed funds for the conference on skin colour variation and biological science leadership in Africa upon which parts of this book are based. The ideas I explore in Nature’s Gifts come from a life shared with my wife, Delecia Forbes, and daughters Gabriele and Isabella. Biologists will not be surprised by what is contained here. The curious lay public and those involved in the humanities and social sciences might be. Wilmot James Cape Town
ix

EUROPE

H, U, X G M B F B M B
AUSTRALIA SOUTH AMERICA NORTH AMERICA

Z Y B X

A, C, D

A*, D*

UK

T, U, V, W

ASIA

I, J, K

N M

A, C, D

L2

L3, M

AFRICA

L1

X? A, C, D

Human Migration Map Using Mitochondrial DNA

3 Skin Colour

RAcE cLASSificATiON

The release in 2010 of the documentary film Skin, the story of Sandra Laing, the woman with ‘coloured’ features born to a ‘white’ Afrikaner couple in the 1960s, creates an opportunity to examine more closely the interaction between genes and the environment when it comes to pigmentation. These reflections occur in the context of a post-apartheid South Africa. I spent some time in the Cape Archives consulting material on ‘group areas’, apartheid’s system of residential segregation. On reading through some initial materials it became immediately apparent that ‘group areas’ could not be understood in isolation from sex and marriage across the ‘colour line’ and that I was looking at an historical picture, at the centre of which stood a government effort led by T E Dönges, apartheid’s first Minister of the Interior, to apply a programme of population engineering that built on and refined existing measures to segregate the country’s population groups. What they tried to create was a national ‘breeding programme’ for human beings. In pursuit of this project, Dönges had to create laws to classify the South African population. He did so by means of the Population Registration Act of 1950, which divided the population into four main groups along lines of appearance and social recognition: Europeans (meaning whites), Asian, ‘coloureds’, and ‘natives’ (meaning blacks).

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Of course, the designation ‘European’ for the descendants of immigrants who came largely from the Netherlands, the United Kingdom, Germany and France was a wistful reclaiming of an identity lost over time; the use of Asian for the descendants of indentured workers who came from certain parts of India, a small part of the Asian sub-continent, was an admission of ignorance or indifference to areas of origin; ‘coloured’ was a fictional assembly of individuals from a diverse set of backgrounds living in one place at a particular time and ‘native’ was later replaced by ‘Bantu’, disposing an already troubled and misleading term to the language of offence. The method of dividing the population in terms of this nomenclature was based on appearance and social recognition: a person was white because he or she was ‘obviously’ white unless proven otherwise by the lack of social recognition by friends or neighbours. The challenge to Dönges and his cronies was what to do with those who did not fit in anywhere in particular, or with those who wanted to alter their classification once it had been set down, or with the thousands of light-skinned coloured individuals who very quickly moved into white neighbourhoods and made white friends before the law was passed. The task was left to the director of the apartheid government’s census, Mr Jan Raats. He had to prepare a detailed system of racial classification in time for the 1951 census, the urgency for which was not simply the need for a population count but for government departments to know to whom they ought to pay state pensions and at what rate.1 Raats came up with the following definitions: • Asiatic means a person whose parents are or were members of a race or tribe whose national or ethnical home is Asia, and shall include a person partly of Asiatic origin living as an Asiatic family, but shall not include any Jew, Syrian or Cape Malay. • Bantu means a person both of whose parents are or were members of an aboriginal tribe of Africa, and shall include
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a person of mixed race living as a member of the Bantu community, tribe, kraal or location, but shall not include any Bushman, Griqua, Hottentot or Koranna. • Coloured means any person who is not a white person, Asiatic, Bantu or Cape Malay as defined, and shall include any Bushmen, Griqua, Hottentot or Koranna. • A white person means a person both of whose parents are or were members of a race whose national or ethnical home is Europe, and shall include any Jew, Syrian or other person who is in appearance obviously a white person unless and until the contrary is proven.2 The Population Registration Act required the governorgeneral to provide definitions for sub-divisions of coloured and ‘native’ people.3 These sub-categories would, in time, lead to the identification of Cape Malays for exclusive residence in Schotsche Kloof, situated on prime property on Signal Hill. For the ‘native’ people the consequences were less than beneficial. In fact, they would prove to be particularly severe – in this subcategorisation lay the demographic basis for the ten ‘homelands’, to which they would, in time, be relegated. What to do, though, with the classification of marginal cases? Most of these people – and there were many: close to 100 000 individuals – fell between the definition of whites and that of coloureds. Raats complained that in his attempt to develop a rasse-skeidslyn (a line of racial division), the marginal cases proved to be a ‘geweldige en uiters moelike taak (a formidable and immensely difficult task)’. Raats cited the instance of Anthony Jooste, a coloured teacher in Krugersdorp, whose mother’s death certificate indicated that she was coloured, but whose father’s race was unknown. Jooste’s marriage certificate described both him and his wife as coloured, but her death certificate said she was white. Their three children were classified as white and attended white schools. ‘They were therefore accepted as white despite the fact that their descent was coloured,’ Raats noted.4 To deal with instances such as this,
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he wanted to know the political terms by which the classification device could be used to drive populations in given directions. Raats and his colleagues decided that a rigid racial line should be drawn around whites as the dominant group and that ‘blood-mixing’ with coloured people should not be allowed to further verydel (corrupt) the already impure white race. However, coloured people who looked like whites posed a problem. Many of them, in anticipation of the passage of the racial laws, rapidly disappeared into white life and met the legal criteria of both appearance and social recognition. Fearing that if members of this group were not allowed into the ranks of the whites they might, given their high fertility rate, become a competing white group, Raats stated that it would be better to classify as white all marginal cases where the individuals involved looked white. This argument was accepted by Dönges and, in the 1950s, was used as the rule of thumb, swelling the ranks of whites overnight. Indeed, if one factors in those who slipped into the white community before apartheid along with those who were made white by the stroke of Dönges’s pen, about 10 per cent of the white population in the early 1950s consisted of ‘coloured’ individuals. Dönges and his state officials planned it that way. They also agreed that Cape Malays, descendants of Javanese slaves brought to the Cape by the Jakarta-based Dutch East India Company in the 17th and 18th centuries, should be protected as a sub-group of coloured people, partly at their request, and partly because Dr I D du Plessis, the Commissioner of Coloured Affairs, was their benefactor.5 Du Plessis had invested a great deal of time developing his anthropological interests in the Cape Malays and writing about them and went out of his way to put what he thought was their case.6 Dönges had come to an understanding with Du Plessis that Cape Malays would get special and favoured treatment, which is why the wonderfully located Schotsche Kloof remained relatively unscathed by group areas removals. As for coloured people who were neither Malay nor of sufficiently fair complexion, they were to be nurtured and
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protected so that a strong sense of ‘coloured’ national pride could emerge. In time they would get their own university, the University of the Western Cape (UWC) and, hopefully, their own geographical homeland. The proposed city of Proteaville, built around the nucleus of what is today referred to as Bellville South, was considered to be a proto-capital of the coloured homeland, which is why the council chambers of the dummybody Coloured Representative Council were built there and why the proposed location of UWC was moved from Athlone, where it was initially intended to be, to Bellville. Raats came into his own when discussing the Indian population. He found their cultural distinctiveness, apparent aloofness and endogamous marriage practices threatening. Indians, he observed, kept their ‘race pure’, but the men were promiscuous with coloured and African women, taking no responsibility for their mixed offspring, who were then absorbed into either the coloured or African communities. He recommended that the state deliberately break down their cohesiveness by classifying the mixed offspring as Indian and not as coloured or African, as was typically the case. Raats suggested a process of what he called verkaffering (‘to make kaffir’): Many bastards are born of parents one of whom is Indian, and this bastard should go to the Indian community, be treated as one and live in the same neighbourhood, even though he might speak Zulu, Sotho or any other language. These bastards will be the medium of disintegration by which the solidarity of the Indian will be eroded, and will turn the Indian into a coloured group with a view of life more compatible with the conditions of our country.7 This grotesquely perverse man admired the extent to which dysfunctional coloured communities served white interests. Raats’s colleagues from the Departments of Justice, Native Affairs and the Interior found this argument to be far-fetched, and failed to respond to it.8 In reviewing his recommendations,
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they suggested that he be given greater investigative powers into the descent lines of marginal cases (descent was not normally a criterion for race in the legislation); that the various departments of state become more consistent in the use of racial classification and that the 30 000 or so fair-skinned coloured people be absorbed into the white group.9 Here, then, emerged a picture of the racial future state officials premeditated – a society divided along racial lines, with distinct cultural and colour characteristics. To this end, racial classification defined the races towards which the officials wanted to move society; sex and intermarriage laws invoked criminal penalties against those individuals who broke the emergent rules of colour and race; and group areas put an end to ‘racial mixing’, as the state officials saw it, by circumscribing the propinquity of sexually available populations and minimising points of social and interpersonal contact. The guiding principle of social apartheid was endogamous reproduction, which, if it continued for some decades its designers believed, would result in the emergence of racially distinct populations. It was ‘racial hygiene’ and ‘population engineering’ practised on a national scale.
SANdRA LAiNg ANd gENETic iNHERiTANcE

Many lives were wrecked. The story of Sandra Laing, as told in Skin, was only one of them but was, perhaps, the most searing and emblematic. Laing, born to an Afrikaans-speaking white couple from the Free State, had a darker skin than her parents and siblings and her facial features resembled those of members of the coloured community. She, like many other white South Africans of European descent, had genes from African and Asian populations. The problem for her was that it showed and she stood out like a sore thumb. As a result, her racial classification was changed and she was driven from her school. Ultimately, she eloped with a black man and lived a hard life in black townships. Although her story is still branded on the conscience of many, hers was not a unique
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tragedy. I knew of a young coloured man, my age at the time, who fell in love with a white woman, but they were not allowed to see one another. He committed suicide by throwing himself at an oncoming train in the Cape suburb of Heathfield. How did genetic inheritance work in the historical genealogy of the larger Laing family? Were they part of the ‘passing for white’ group or among those reclassified by Dönges, or did matters go much further back? We know that many whites, including Afrikaners, shared descent lines with other, darkerskinned South Africans. Genetic material from an ancestor or ancestors that coded for a darker skin, curlier hair and facial features such as raised cheekbones came down the Laing genealogy, some of it by-passing Sandra’s parents, but switched on in her and her brother. What is the current state of knowledge about genetic inheritance that could shed some light on the process? This is what we know today: differences in human skin colour are a consequence of the distribution and biochemical properties of a melanin group of brown, black, yellow and red pigments found among animals. Melanin is a mixture of lightabsorbing biopolymers, with properties similar to those that give colour to, for example, synthetically produced nylons or paints. Pheomelanin contains yellow and red pigments and eumelanin brown and black. The nature of melanin ‘frustrates precise chemical description’, writes University of Edinburgh dermatologist Jonathan Rees.10 Melanin biochemistry is complex and remains poorly understood. Early in the first trimester of foetal development, cells called melanocytes migrate from the neural crest, so named because it is perched on top of the neural tube, to its destination in the skin. In her book Skin: A Natural History, Nina G Jablonski writes of melanocytes as ‘immigrant cells’, which are those ‘that migrate into the skin from elsewhere in the body during early development and that retain some ability to move out of the skin during life’.11 Cell migration patterns have increasingly been captured by modern digital imaging technology and, given the
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speed of innovation in this sector, we will increasingly be able to see in pictorial form processes of cell division and movement. These immature cells appear to know when to migrate, where to migrate to, how to recognise their destinies and, of course, when to stop. To help understand these processes, biologists have developed the concept of the ‘fate map’, which, as the word suggests, is a diagram of what becomes of the individual cell during embryogenesis, a word referring to the maturation of the embryo among animals (and other organisms). Fate maps are, of course, under full genetic control. The destination of melanocytes is that part of the epidermis just below the surface of the skin as it borders with the deeper skin layer called the dermis. In mature form the melanocytes possess dendrites, tiny octopus-like extensions wedged into the skin layer, and there are thousands of these distributed throughout our bodies. It is within these mature cells that organelles called melanosomes produce melanin by way of a pathway that involves a process called tyrosine metabolism. Once the melanosomes reach maturity they move and give colour to the keratinocytes, another cell type we have by the million and which gives our skin its colour. We could, therefore, say that the melanosomes are the factories of melanin made for the benefit of populating the keratinocytes. It is significant in terms of biological function that the keratinocytes use ultraviolet B radiation to convert cholesterol into basic vitamin D, which the liver and then the kidneys progressively convert into the active form of vitamin D – for here resides the first clue to the functional biology of skin pigmentation. To talk about cells ‘knowing’ what to do, when to do it and when to stop doing it, suggests that they are capable of communicating with one another and that they are given instructions by our genes. ‘It is widely understood,’ even among non-specialists, writes Rees, ‘that human skin color and hair color are largely under genetic control.’12 Genes are, of course, our instruction manuals. We know of some (but not all) the genes responsible for encoding the cellular
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machinery that makes melanin. Finding the others is the job of the scientists. It is of interest to the biomedical community, and particularly the dermatologists among them, to understand better genetically inherited disorders of the skin like albinism and cancer. It is also of interest to the cosmetics and sun-tanning industries in order for them to make better products, gain more market share in what is a multi-billion-rand industry, and perhaps to develop innovations that can biochemically stimulate a lighter or darker skin. There are some products on the market that absorb ultraviolet rays and prevent the sun from darkening our skins, and others that directly inhibit the production of melanin itself. Genetic manipulation remains, for now, an abstract possibility. We know that there are many genes responsible for skin colour variation, but we do not know how many. Gregory Barsh notes, for instance, that 100 genes are known to affect mouse pigmentation.13 Our best guess is that we have anywhere between 5 and 15 candidate loci, with a number of genes at each. A growing number of skin-colour genes have been discovered and described, one of them the melanocortin receptor MC1R gene found on chromosome 16, which is associated with regulating sun sensitivity (first described by Rees, Tony Thody and Ian Jackson as the gene for ‘red hair’, or, less flatteringly, the ‘carrot top’).14 Another is the gene that goes by the letter P and is found on chromosome 15, where mutations or changes in the DNA sequence have been linked to various forms of albinism; recently Keith Cheng and his collaborators found a gene, SLC24A5, which is the human counterpart of a gene found among zebra fish. This ‘apparently accounts for a significant part of the difference between African and European skin tones’, Michael Balter wrote in the magazine Science, as ‘one variant of the gene seems to have undergone strong natural selection for lighter skin in Europeans’.15 Cheng presented his findings at the fourth annual meeting of the Africa Genome Education Institute, noting that ‘lighter variations of pigmentation in man and golden zebra fish are
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caused by similar alterations in melanosome morphogenesis’.16 Balter observes that although Cheng’s work does not solve the question of why fair skin might have been favoured among Europeans ‘it is consistent with a long-standing but unproven hypothesis that light skin allows more absorption of sunshine and so produces more Vitamin D, a trait that would be favored at less sunny European latitudes’.17 More recently, a study of 2 986 Icelanders and 1 214 Dutch individuals by Patrick Sulem and his colleagues found five genetic variants to be associated with eye, hair and skin colour differences among Europeans, the population that has the greatest phenotypic variety of the sources of colour in the world.18 From the point of view of functional biology melanin is a very effective sun-block agent, protecting against the harmful effects of a certain band of ultraviolet radiation. Rees observed that ‘[M]elanosomes … tend to produce caps over the nuclei, shielding the nuclear material from ultra-violet radiation.’19 Advances in understanding the biochemistry of melanin confirm today what people observed long ago: that individuals living in equatorial areas tend to have darker skins because of much higher levels of melanin than those who live closer to the poles. The Swedish botanist Carolus Linnaeus recorded in the 18th century the correspondence between geography, sun exposure and skin colour.20 Skin colour is a form of quantitative variation, something we all have, but in varying amounts, like skeletal architecture and bodily frame, for example. Such forms of variation are usually very sensitive to the environment: nutrition in the case of height and weight, ultraviolet radiation in the case of skin colour. This is unlike qualitative forms of variation, a quality you either have or do not have, like blood group type, for example, which are environmentally insensitive. The result of what Richard Lewontin calls ‘joint actions’ and ‘norms of reaction’ of life, the evolution of skin colour variation involves a causal interplay between biological actions, environmental factors and human decisions, though in ways that are not always obvious.21
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Presenting the environmental aspects of the ‘joint actions’ of skin colour evolution, Nina Jablonski and George Chaplin write that ‘[R]ecent epidemiological and physiological evidence suggests to us that the worldwide pattern of human skin color is the product of natural selection acting to regulate the effects of the sun’s ultraviolet (UV ) radiation on key nutrients crucial to reproductive success.’22 Scientists initially believed that pigmented skins developed to protect against skin cancer-causing ultraviolet radiation, but, as cancer tends to develop after reproductive age it is not the risk of developing melanomas that drives the evolutionary process. Therefore, ‘[A]n alternative theory suggests that dark skin might have evolved primarily to protect against the breakdown of folate (folic acid), a nutrient essential for fertility and for fetal development.’23 Jablonski and Chaplin were not the first to make the link between skin colour variation, the sun’s ultraviolet rays and vitamin D processing. In their book, Man’s Evolution, published in 1965, C Loring Brace and M F Ashley Montagu devoted a full 20 pages to the shaping of human variation that included an analysis of skin colour, laying the anthropological foundations for today’s molecular explanations.24 Jablonski and Chaplin built their findings on two earlier scientific studies: one showing that folate (part of the B vitamin complex) in the human body breaks down rapidly in experimental conditions during exposure to intense ultraviolet (UV ) radiation, especially among lighter-skinned people. Up to half the folate in blood plasma, tested under experimental conditions, can be lost in under an hour. The second study to which they refer found ‘that too low folate levels can cause debilitating neural tube defects (NTDs) in foetuses (the reason why women should take folic acid before and during pregnancy)’.25 Spina bifida is one of those neural tube defects. Jablonski and Chaplin go on to talk about the role of folate in the synthesis of DNA in cell division. Anything associated with rapid cell proliferation such as spermatogenesis requires folate,
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they say. Their hypothesis is that dark skin evolved to ‘protect the body’s folate stores from destruction’.26 Their conclusion is a combination of anthropology, human biology and high-technology satellite data. Building on the earlier mapping exercises of the Italian geographer Renato Basutti, Jablonski and Chaplin took the global ultraviolet measurements established for the first time by NASA’s Total Ozone Mapping Spectrometer between 1978 and 1993 and compared these with published materials on the global distribution of human skin colour variation. They were able to model the distribution of UV radiation on the earth and relate the satellite data to the amount of ultraviolet B radiation (UVB) necessary to produce vitamin D. They found that the earth’s surface could be divided into three vitamin D zones: one comprising the tropics, one the sub-tropics and temperate regions, and the last the circumpolar regions north and south of about 45 degrees latitude. In the first, the dosage of UVB throughout the year is high enough for humans to have ample opportunity to synthesise vitamin D all year. In the second, at least one month during the year has insufficient UVB radiation, and in the third area there is not enough UVB on average during the entire year to prompt vitamin D synthesis. This distribution could explain why indigenous peoples in the tropics generally have dark skin, whereas people in the subtropics and temperate regions are lighter-skinned but have the ability to tan, and those who live in regions near the poles tend to be very light skinned and burn easily.27 It is true that Native Americans have a distribution of skin colour that varies much less than that of others who populate the continents across the spectrum of ultraviolet radiation zones as described by Jablonski and Chaplin. Perhaps this is due to their relatively recent arrival as migrants across what is now known as the Bering Strait an estimated 13 500-20 000 years ago, when
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the American archaeological record suddenly springs to life.28 Other anomalous examples are the skin colour of the Inuit populations of Alaska and Northern Canada, who have darker skins than we would, perhaps, expect. Relatively late settlement (about 5 000 years ago) and a vitamin D-rich, fish-based diet, possibly explain this phenomenon. Lewontin believed that we need to find more systematic evidence in support of the relationship between vitamin D and which peoples survived and which did not; who produced offspring and who did not, that is to say, our survival and fertility rates. Harshly put, perhaps, but without systematic data on what he calls ‘survivorship and reproduction’, ‘adaptive reconstruction of the causes of human differences remains essentially an amusing pastime, testing our ingenuity as imaginative story-tellers’.29 So, what story can we tell with incomplete data about the Laing family? There are African versions of the genes that code for melanin synthesis and these presumably became a part of the Laing genealogy. To find a precise answer would involve taking DNA from a multi-generational sample (called a pedigree) from the Laings and their related families. We do know that, in general, genes regulating skin colour have ‘throwback’ features in the inheritance process, and can go back to grandparents, great-grandparents and further back to ancestors of whom we often have no memory or photographic record. I was recently surprised to see a photograph of my wife’s grandmother, who is as dark as she is, although her mother is much lighter in skin tone. Genes expressing a certain skin tone often bypass one or even two generations in the shuffling process that is part of bisexual heredity.
cOLOuR

The most recent hominid-chimpanzee ancestor probably lived about 8- to 10-million years ago. Today’s chimps have a pink skin covered with hair. Perhaps what became the Homo line also started off with a pink skin. We had the biological ability to change colour inter-generationally through natural and sex
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selection (unlike the more sophisticated chameleon and some amphibians, where skin is camouflage and changes by habitat). We have to remember that we vary far more and in much greater detail than the human eye can apprehend. It is entirely possible that all the Homo lines possessed the adaptive capacity to evolve skin colour pigments that varied by geophysical location and ecological niches on the planet, at least to the extent that the concentration of bodily hair required it. Skin is soft tissue and does not fossilise, except under highly exceptional circumstances. It seems as though the hominid line progressively lost bodily hair as a result of the evolution of heat-dissipating sweat glands. We do not know what colour their skins actually were. DNA has been extracted from both our last surviving co-inhabitants, Homo neanderthalensis (which died out between 24 000 and 30 000 years ago) and Homo floresiensis (which went extinct about 13 000 years ago), but it is uncertain whether one may determine skin colour from the samples. In the past 100 000 years, modern Homo sapiens started to migrate out of our African homeland, probably first turning eastwards and expanding along the coasts of Arabia and southern Asia towards China and Australia. Nina Jablonski argues that the original African populations of modern human beings had a dark skin because of the tropical environment in which they evolved.30 Those who settled in western Asia or Arabia may have been the source of the moderns who started to ‘infiltrate Neanderthal strongholds in Europe, or they may have derived directly from a second wave from North Africa’.31 Within Africa, some of our ancestors reached the Cape about 180 000 years ago. The colour of their skin was probably dark brown, much like that of the remaining San populations living in Botswana, Namibia and South Africa today. Africa was largely a continent of brown-skinned peoples until the great black migration from the Niger-Congo area that started about 1 500 years ago.32 If our ancestors moved on average five miles a year, they were able to populate the Indian and South East Asian coastlines and, with primitive maritime technology, reach Australia’s Arnhem
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Land about 55 000 to 60 000 years ago. Their skin colour probably became as dark as that of today’s living Australian Aboriginal population.33 Middle Eurasian populations developed lightercoloured skins and northern Europeans even more lightly pigmented ones. Keith Cheng and his collaborators’ gene SLC24A5 is associated with the pale skin and gold hair found among northern European populations. Jonathan Rees’s MCR1 gene is associated with the red hair and pale freckled skin seen today in unusual incidence among the Scottish and the Irish. The movements into the Americas probably started about 20 000 years ago, when a landmass called Beringia connected presentday Siberia and Alaska. There were other waves of migrations which resulted in a permanent pattern of settlement of a people who developed dark to light-brown skins with a reddish tint, the Far East Asian populations from whom they descended having an already established light skin with a yellowish tint. Biochemically, both are a result of a more dominant pheomelanin activity in melanin production.34 Some of our more distinctive outward features, determined as they are by the tiniest fraction of DNA, suggest that groups of human beings reproduced in relative isolation from one another for some time; long enough for variations in certain features, like skin colour, to become more homogeneous. Gene-flow across populations would have been minimal in those times. Scientists today believe that the isolation was not so great that it allowed for races or sub-species to emerge. Also, there is no evidence that the world’s different populations were a consequence of crossbreeding with other Homo lines. Whenever the period of relative isolation began, it ended about 10 000 years ago, when the first innovations in agriculture were introduced and populations started to grow. Modern agriculture and medicine unleashed the explosion in human populations in the 16th to 17th centuries, initiating a period of growth and, as a result of today’s travel technologies, a modern
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period of unprecedented human mobility which erodes national patterns of sexual selection and, in the age of globalisation, also those between north and south. Richard Lewontin wrote, Countries of the Southern Hemisphere, are contributing many more genes to the species as a whole than are northern industrial countries. The consequence will be that the human species will come, more and more, to have gene frequencies that at present characterize Africans, South Americans, and South Asians. As far as anyone knows, this trend will have no interesting consequence, except that it will make the species as a whole look less variable but darker in skin color.35
cOLOuR By SigHT

The final part of the story is about the way we see or apprehend colour and its role in what Charles Darwin, in The Descent of Man, called sex selection.36 ‘We think of our eyes as wise seers,’ Diane Ackerman wrote in her Natural History of the Senses, ‘but all the eye does is gather light.’37 The brain interprets the lightwave patterns as colour. How much variation in skin colour do our eyes see? A lot. But not as much, I am sure, as those of an eagle or owl or cheetah, though there are 32 words in Brazilian Portuguese for variations in skin colour that the human eye apprehends in Brazil’s colour-blind democracy! In fact, we see much more homogeneity than there actually is. It is possible that, in reality, the variation between human beings spread across the globe in pre-modern times was much finer and more nuanced than we may think. We were thinly spread and had little contact with individuals who were strikingly different to our eyes. Still, most of our ancestors probably thought their skin colour was the only one. Travel narratives indicate that towards the end of the 19th century there were many peoples who had not yet encountered human beings with a strikingly different colour from their own. When contact occurred it must have been a surprise, perhaps even a shock.
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SkiN cOLOuR

In most species that reproduce bisexually the males and females have different reproductive strategies due to a difference in relative investment in producing offspring. The well-known Bateman principle relevant to this topic is that women have a variety of social and psychological strategies for sniffing out the right mates over, typically, a longer or shorter courtship period and, of course, they may be unsuccessful. Some studies suggest that, for men, the skin-colour tones of women can be read as indicating their reproductive health status and that the common assumption that human beings will use any marker that is socially available as a basis for discrimination is mistaken. Skin colour triggers a memory of reproductive success in particular environmental niches and, culturally, is associated with mate selection and the aesthetic repertoires of desire and beauty. To put it simply, a fair-skinned woman became desirable to men under circumstances – in the ice ages for example – where such a skin allowed for a life long enough for the women to bear many children. A fair-skinned woman would have no such advantage in the tropics, but a dark-skinned woman would, which is why records indicate that light-skinned women were seen as unattractive in tropical regions. Light-brown-skinned women can navigate many more worlds, which is perhaps why they are seen as the most desirable of all. Certainly, the sun-tanning and skin-lightening cosmetics industry depends mightily on the evolutionary history of skin colour aesthetics and our sense of what is beautiful and what is not. Karl Grammar and his co-authors provide a useful review of the field, writing that beautiful and irresistible features have evolved numerous times in plants and animals due to sexual selection, and such preferences and beauty standards provide evidence for the claim that human beauty and obsession with bodily beauty are mirrored in analogous traits and tendencies throughout the plant and animal kingdom.38

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They continue: Human beauty standards reflect our evolutionary distant and recent past and emphasise the role of health assessment in mate choice as reflected by analyses of the attractiveness of visual characters of the face and the body, but also of vocal and olfactory signals. Skin colour and tone are, therefore, cues to past periods of reproductive health and fitness. We are also a violent species and the rape of women, particularly during and in the aftermath of war, has contributed to modifying the skin tones of populations. In slave societies, too, like those of the southern United States and South Africa, the male promiscuity of the dominant group found its outlet, and violently so, among the darker-skinned women of the slave communities.
THE PROBLEM WiTH REdHEAdS

Less than one per cent of the human population has red hair, but Scotland boasts a 13 per cent incidence. Red hair is regulated by the recessive MCR1 gene which is carried by 40 per cent of the Scots and 35 per cent of the Irish. Redheads are also to be found in Wales, the USA, in Northern and Western European countries, and in Russia. There are redheads among the Berber and Kabylie populations of Northern Algeria and Morocco and in Northern India and Pakistan. Red hair is also common among the Pushtuns of Iran. As hair and skin colour work in tandem, Rees established that MCR1 is associated with a lightly-pigmented skin, sometimes freckled, and with red hair. The gene produces more pheomelanin chemicals, of the yellow and red variety. Scientists are interested in the connections between red hair, lightly-pigmented skin, pigment-bereft albinism and cancers. The health issues are serious because redheads burn very easily as their skins cannot tan. There is also evidence that individuals
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with red hair respond differently to anaesthetics. It seems that redheads experience certain categories of pain, particularly thermally induced pain, more powerfully. Their particular version of the MCR1 gene (called an allele) releases more of the hormone that stimulates histonal cells as well as stimulating a brain receptor linked to pain sensitivity. MCR1, dated as an old gene – 100 000 years – is found in many species (wolves, for example), located in the same genome place. Studies suggest that it was kept in the human gene pool because it is a by-product of a lighter skin that gave its owners an advantage in environments where sunlight was scarce. Scientific studies have shown that having a lighter skin prevents rickets in cold climates and allows for better heat retention. Redheads find it difficult to live in tropical climates. They should stay away from the sun and are vulnerable to skin cancer. There is evidence that red hair is kept out of African gene pools precisely because it not only confers no advantage but is positively harmful. It is linked, too, to some secondary diseases. Kwashiorkor can, for example, turn dark hair red or blond and one of the many varieties of albinism found among members of the population of Africa and New Guinea results in red hair and skin. Like so much else, human characteristics can be both a blessing and a curse. Red hair has been associated, mythologically, with hotheadedness and, in mediaeval times, with evil spirits. Today it is often a point of insult (being called a ‘carrot top’ is no compliment) and discrimination. Discrimination against redheads is listed in the United Kingdom as an example of the infringement of civil liberties and a jurisprudence has developed based on some case histories. In this day and age it seems perverse. Still, if the red-hair story were as perverse as apartheid there would emerge an organised and society-wide system of allowing redheads only to marry redheads and have them all live together in neighbourhoods set aside by law especially for them. If a redhead was caught having sex with an individual with throwback non-redhead genes, public embarrassment and
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criminal penalties would follow. Over the course of time a nation of redheads would emerge. Sound crazy? That was apartheid in South Africa; an effort to create an organised society-wide system of population inbreeding, and it succeeded … up to a point.

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Human Tree Of Descent

H U7 U6 U8 K U9 U3 U2 U4 U1 3 T X W U5

V B F Y A E Q D G M2 M1 C Z

R

I

N M

L3k

L3a L3b L3c L3d L3j L6 L1c4 L1c6 L1c3 L3f L3i

L3x L3e

L3 L2

L3h

L4

L2d L2c L2b L5 L2a L0k L0a1 L0a2 L0a3 L0a4 L0b

L2e

L1c2 L1c1 L1b

L1 L0

L0d1 L0d3 L0d2
AFRICA ASIA AND THE AMERICAS EURASIA EUROPE

L0f

The letters and numbers represent particular lines of descent grouped into categories called haplotypes.

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