William Hamilton's worries about the future of the human genome

Was
Darwin
Wrong?

What others say

What's the problem?

Hamilton's worries

Discussion

Conclusion*

Further Reading

William Hamilton's worries about the future of the human genome

A review by Gert Korthof 25 Aug 2011 (updated 10 Feb 2012)

This review was triggered by the BBC documentary with the mysterious title All watched over by machines of loving grace (1). The documentary appeared to be a story about Bill Hamilton, George Price and altruism (and much, much more). One of the themes of the documentary was altruism and selfish genes, but I will focus here on what was the biggest surprise for me: Hamilton had eugenetic ideas.

Bill Hamilton at the Kyoto Prize ceremony 1993 (list of winners).

(images from the documentary captured with Open Movie Player and edited with GIMP Image Editor)

Philosopher Michael Ruse interviewed in the BBC documentary (2)

According to the documentary Hamilton wrote about eugenics in Narrow Roads of Gene Land (2). In the documentary philosopher Michael Ruse confirmed that Hamilton had eugenic ideas:

"Hamilton believed that some people were genetically inferior to others. We should take a stand against the slide into degeneration."
"... it would let the genetically inferior to survive and so would weaken society".

If Michael Ruse, a philosopher of biology, who devoted his academic career to Darwin and evolution, tells us these things about the famous evolutionary biologist, there must be some truth in it. However, we do not get any details in the documentary. So, I wanted to find out what exactly Hamilton wrote about eugenics, and what his position was. This is not easy because the 3 volumes of Narrow Roads of Gene Land add up to no less than 1904 pages! Fortunately, the index points us to a number of interesting locations in the text. The most extensive discussion can be found in Volume 2 chapter 12 with the mysterious title 'The hospitals are coming' which is a 60 page introduction to the article 'Sex and Disease' (25 pages).

Volume 1 Evolution of Social Behaviour

Volume 2 Evolution of sex

Volume 3 Last Words

Hamilton's worries
Hamilton worries that modern medicine (16) eliminates natural selection and our genome will steadily accumulate more and more deleterious mutations. He fears that our scientific and technological abilities to diagnose and repair these mutations will be insufficient, now and in the near future. The accumulation of mutations has accelerated since humans have partly eliminated natural selection by medical practice. Medical intervention amounts to phenotypic curing (spectacles, phenylalanine-free diets for PKU) of every defect in the germ line (see: What's the problem? below). He does not believe that future technologies will improve to the point that we will be able to correct all deleterious mutations and can keep our genome healthy. Hamilton is more pessimistic than scientists like Steve Jones and John Maynard Smith who think we can repair the known deleterious mutations by engineering the germ line or soma. There are hundreds known single gene disorders, but many more unknown genes that affect disease. Hamilton estimates there are 100.00 genes in the human genome. Most mutations are bad. The majority of genes will accumulate bad mutations. This happens at a rate of perhaps one mutation per genome per individual per generation ('Kondrashov threshold', p. 464), but the rate could be an order of magnitude above or below the Kondrashov threshold, so between 0.1 and 10.

"If humans turn out to be near the Kondrashov limit —that is, if on average every gamete has one bad mutation created during the lifetime of its producer— it is obviously not going to be nearly enough to test a baby for the subset of the few hundreds or so of well-characterized genetic defects (...) There are certainly tens of thousands more possible mutations that the baby could have that have not been characterized well enough ..." (p. 465). (my emphasis)

Correcting those is an impossible task just like Maxwell's Demon (66). This is a depressing future. We are unable to prevent the degradation of the human genome.

The human embryo is like a fish
The above arguments are about empirical data. Controversies can be solved by data. However, there is a second argument in Hamilton's writings. It involves moral values, although it is mixed with empirical data. Consider this: natural selection kills at every age: egg, sperm, fertilized egg, fetus, embryo, baby, child, adult. However, humans can choose that moment as early as possible (when the individual is still in the womb) avoiding infanticide. Hamilton:

"for me, tiny embryos that seem to be in fish-like stages of human development, or earlier, are genuinely fish-like or even more primitive. Indeed, I believe additionally that they experience less of pain, fear, and danger than fish experience. Because of this I would genuinely be happier 'terminating' such early human embryos in a Petri dish than I would be 'terminating' an adult fish in an aquarium." (p. 460).
"By kidding ourselves about some weird kindness to embryos, to neonates, and the like now, we are actually being very unkind to numerous far more sentient persons of the future." (p. 476)
"If one is going to kill a baby, clearly it is only tolerable for it to be killed painlessly" (p. 481 Vol 2 Narrow Roads).

Empirical data about pain sensation in human embryo can be found at Neonatal perception. Hamilton's argument about pain in the human embryo —whether we agree or not (26)—stands on its own feet, and is not connected with the argument about the burden of deleterious mutations. Whether one is against abortion of human fetuses and infanticide, this does not change the fact that humans accumulate mutations when natural selection is eliminated. So far Hamilton (Vol 2 p.449 - 497).

What is the problem?

What is the problem Hamilton is worried about? In short: natural selection is weakened because medical science saves lives that otherwise would die, and helps reproduce those that otherwise could not, and subsequently transmit the medical problem to the next generation. And so on. Examples:

Caesarean section is mentioned by Hamilton. It is against natural selection because the baby would have died without medical intervention. When the problem is caused by genetic factors and the baby is a girl then she has an increased risk for Caesarean section also. The problem is transmitted to the next generation. Hamilton is in favour of disincentive schemes by the state (Vol 2 p. 504). Hamilton did not research the subject. Here are some recent data: (8). In fact, any assisted birth in a hospital, contrary to home birth or natural birth, are examples of 'assisted reproduction'. However, Hamilton forgets that Caesarean section could also have a positive outcomes after many generations by eliminating constraints on head size of the baby at birth. It could enable an evolutionary increase in skull and possibly brain size. At the moment head size is constrained by natural birth.
Assisted Reproductive Technology (ART) (31) such as: in vitro fertilization, artificial insemination (30), eggcell or sperm donation, Intra Cytoplasmic Sperm Injection (ICSI) (65), surrogacy are against natural selection because without medical intervention these couples could not reproduce. The infertility problem is transmitted to the next generation. However, ivf in combination with preimplantation genetic diagnosis (PGD) could have eugenic effects.
mother's milk: when mothers are unable or unwilling to breastfeed their baby they can use a commercial breast milk substitute (formula) and their baby will not die. There are also substances that promote lactation in humans. In all cases natural selection is prevented. If the inability to breastfeed is (partly) hereditary, the problem will be transmitted to the next generation. And so on.
Medical treatments: regenerative medicine, organ transplantation, blood transfusion, pharmaceutical drugs including essential medicines (52), antibiotics (46), vaccination, medical devices (e.g. hearing aid (37), and non-medical inventions such as clothing, cooking, soap, houses with air co and central heating, etc. all affect survival and indirectly reproduction.
Examples of hereditary diseases where new treatment makes reproduction possible:
- PKU is a hereditary disease mentioned by Hamilton. There is no cure for PKU, but newborns who are diagnosed early and maintain a diet low in phenylalanine (Phe) can have a normal life span with normal mental development. Women with PKU can have normal pregnancies when they maintain the low-Phe diet. With new technologies, some are men becoming fathers. In those cases the mutation is transmitted to the next generation. Because of this success PKU has become increasingly common.
- Hemophilia: prior to the 1960s when effective treatment became available, average life expectancy was only 11 years (no reproduction possible). By the 1980s the life span was 50–60 years. According to the WHO: "Children with haemophilia now face few limitations. They certainly attend normal schools, most jobs are open to them, and full participation in society through employment, marriage and having children is now the norm. It is anticipated, however, that the number of people with haemophilia in developed countries will increase steadily over the next few decades".
- Cystic Fibrosis (CF or mucoviscidosis) is a recessive genetic disease which affects about 30,000 people in the United States. Hundreds of mutations cause defects in the CFTR protein. In 1959, the median age of survival of children with cystic fibrosis in the U.S. was six months. Through improved treatment, it is now (2011) 37.4 years. For example: a new drug VX-770 improves lung function and results in fewer infections in patients 6 years and older with the rare G551D mutation (82). Female infertility may be overcome by assisted reproduction technology, particularly embryo transfer techniques. Male infertility caused by absence of the vas deferens may be overcome with testicular sperm extraction (TEST), collecting sperm cells directly from the testicles. If the collected sample contains too few sperm cells to likely have a spontaneous fertilization, intracytoplasmic sperm injection can be performed. In 2008, 240 American women with CF were pregnant.

If all these medical interventions are added up, they substantially reduce natural selection, so mutations will accumulate in the human genome (which has already a high spontaneous mutation rate).

Can medicine help natural selection?
Medical science could 'help' natural selection in specific cases by sterilization, chemical or surgical castration, prenatal diagnosis, abortion. The paradox: 'helping' natural selection is interfering with nature. Hamilton is against eliminating natural selection, but is for abortion and infanticide which is interfering with nature. One could argue that civilization is one big interference with nature.

Has natural selection been eliminated?
No! Miscarriage or spontaneous abortion, habitual abortion, stillbirth, premature birth (45), incurable infertility are examples of natural selection in humans (36). Further examples: childhood deaths, infectious diseases (HIV, influenza, measles, EHEC), and incurable genetic diseases (i.e. Tay-Sachs disease). When looking globally, infectious diseases are the number one killer of humans and therefore the main selective pressure exerted on our species (74). The H2N2 flu virus killed 70,000 people in 1957 in the United States alone (75). Single Nucleotide Polymorphisms (SNPs) in a number of genes governing immunity affects a person's susceptibility to infectious diseases, and how sick he or she becomes from those infections (76). This creates an opportunity for natural selection.

Not natural selection are:

road accidents, war, death by starvation, air pollution, radioactivity, natural disasters (tsunami, earthquake, lightning, fires) do kill but are not natural selection because organisms cannot adapt to these events.
No effect on natural selection: all late-onset disorders such as Alzheimer's, Huntington's, ALS and heart disease because they have children before the disease is manifest.
All medical treatments after reproductive age which do not affect the number of children, do not have an effect on natural selection.
Mixed, small or unknown effects on natural selection have all interventions that enhance sexual attractiviness and sex: breast enlargement, bra, lipstick, high heel shoes, plastic surgery, erectile dysfunction drugs, personal lubricant and so on. In so far these treatments result in enhanced reproductive success, they fall into the category 'assisted reproduction', and weaken natural selection.

The long term effect is that humans cannot reproduce without medical assistance.

Opportunities for natural selection:
when every conception produces a baby and when every couple has the same number of children, then there is no natural selection. Demographic developments in the EU show that the average number of children per woman is 1.5 children. The opportunity for natural selection is smaller compared with large families and when differences between the number of children per woman is also smaller, then natural selection is also less. According to Michael Lynch as a result of an orders-of-magnitude reduction in Effective population size in all multicellular eukaryotes compared to unicellular organisms, there is a decline in the efficiency of natural selection (34).

William D. Hamilton (1936 - 2000)
© James King-Holmes/Science Photo Library.
printed in Vol 2 Narrow Roads

What others say

Let me first quote Alan Grafen who wrote a biography of Hamilton which is included in the Narrow Roads as chapter 20:

"One non-scientific and even political theme recurs throughout the autobiographical essays, and it is one that troubles many readers. Hamilton develops a eugenic argument, which was deeply felt and persistently argued. (...) It shocks many readers when Hamilton advocates infanticide; suggests the denial or in more emollient mode the strict regulation of fertility treatment; and worries about the long-term effects of saving the lives of mother and neonate with a Caesarean section." (...) It has to be said that the scientific basis of Hamilton's eugenic views is not established, though he provided interesting possible hypotheses."
"A start on an intellectual engagement with Hamilton's views has been made in the considered remarks of Haig (2003)"
(Narrow Roads Vol 3, chapter 20, p.447,448)

David Haig:

"I think that many of Hamilton's ideas on eugenics are naïve and misguided, but I support his plea that this should be a question on which one could have a reasoned, nonacrimonious exchange of views. The extent to which the human genome is deteriorating is an important question that should be squarely faced, rather than side-stepped."
David Haig (2003) (6)

John Maynard Smith:

"Don't you think that in the timescale in which medical treatment is going to lead to an increase in the frequency of deleterious genes, we are going to see techniques for actually changing the genes themselves (that is, eugenics)? If we can actually transform deleterious genes into beneficial genes in the germ line..." (Narrow Roads Vol 2 p.456).

Discussion

Old and new eugenics
Old eugenics is rejected by nearly every thinking person, including Hamilton^{<citation required>}. The main characteristics of old eugenics are: racism, involuntary sterilization, discrimination against certain groups of people, state interference in personal affairs, infanticide, and lack of genetic knowledge. New eugenics or 'neo-eugenics' (Armand Leroi) is based on voluntary reproductive decisions of parents informed by knowledge of the molecular and genetic basis of diseases. In the new eugenics there is no place for the bad practices of old eugenics. Also, the association of the new eugenics with 'closing the hospitals' (Hamilton's indiscriminate distrust of the medical profession and pharmaceutical industry) must also be avoided. In the new eugencis there should be no discrimination of mentally and physically handicapped people.

Dilemma: individual health versus the health of the human genome
Health is a worthy goal. If the goal of Hamilton's eugenics is genetic health, and if genetic health is a means of achieving phenotypic health, then genetic health is an important -albeit indirect- goal. A healthy genome is a good investment in the health of future generations. Would anybody consciously transmit a degenerate genome to future generations? It would be unethical if we could do something about it and didn't do so. We should care about the quality of our genome, just as we care about the resources of our earth such as food, clean air, clean water, and energy (60). All medical intervention so far is treating the body (41). Indeed that is myopic. Genetic deterioration seems imperceptible because it is abstract and occurs in small steps in many generations.
However, and this is important, if the goal of eugenics is genetic health, then it is not necessary to imitate natural selection by killing handicapped neonates or preventing patients with genetic diseases to have children. Nature is very cruel and wasteful. We should not copy that method, but find more humane ways (compare: planes are not copies of birds!). Hamilton apparently was obsessed by nature's way of eliminating less fit individuals (letting them die). His own death was probably caused by his refusal to have adequate drugs and medical treatment (3). He clearly sees the disadvantages of medical practice, but completely overlooks the advantages. The advantages are so great that a dilemma arises between the health of the individual and the long-term genetic health of the human species. It seems that Hamilton did not see that dilemma.

The dilemma creates the paradox of civilization
When the quality of the human genome becomes a goal in itself, it would justify letting people die from infectious diseases and cancer, because it would improve the human genome by letting natural selection do its job (43). For example: if we eradicate smallpox then there is no longer selection of smallpox on the human genome. That would be bad for the human genome. Should we therefore keep smallpox? That would be madness. Similarly, withholding adequate care during pregnancy and delivery, withholding care for the newborn baby and letting premature born babies die, would also be justified when the quality of the human genome would have the highest priority. In fact, the prevention and treatment of all infectious and genetic diseases, if it enabled people with harmful mutations to survive and reproduce, would be bad for the human genome. But the end of medical treatment would be the end of civilization (9). Therefore, the quality of human genome should be a means, not a goal. According to psychiatrist Laurent Mottron:

"variations in gene sequence or expression of the human genome may have adaptive or maladaptive consequences, but they cannot be reduced to an error of nature that should be corrected. The hallmark of an enlightened society is its inclusion of homosexuality, ethnic differences and disabilities. Governments have spent time and money to accommodate people with visual and hearing impairments, helping them to navigate public places and find employment" (68).

Is the dilemma always real?
There is not always a dilemma. For example, thalassaemia patients may have reduced fertility or even infertility. Children who are diagnosed with Huntington's disease do not usually live to reach adulthood and don't reproduce. Males with Down syndrome are usually unable to father children, while females have lower fertility than females without the syndrome (25). Women with sickle cell anemia are at increased risk for fetal loss (69). Tay-Sachs disease usually results in death by the age of four. At least 97% of men with cystic fibrosis are infertile, but not sterile and can have children with assisted reproductive techniques. Further, natural selection is still in operation in case of miscarriage, spontaneous abortion, habitual abortion, stillbirth, and (untreated) infertility. So, in many cases, nature herself solves the eugenic problem. However, as soon as medical treatment enable patients to become parents, the dilemma becomes real (21). In my country mentally handicapped persons can have children, because enforced anti-conception is forbidden by law. There you have the dilemma.

Is the dilemma recognized by professionals?
There are examples of complete lack of awareness of the dysgenic effect of medical treatment. For example, it is good news for the patients with genetic diseases when treatment enables them to have children, but it is bad news for the human genome (7). Another example is displayed in an article about the treatment of the hereditary disease Cystic Fibrosis (CF) in Scientific American where the authors discuss all sorts of treatments of CF that increase life expectancy and the quality of life of their patients without even mentioning, let alone discussing, the effects of having children has on the human genome (10). Many more examples could be given (40, 53). Hamilton deserves credit for trying to start a discussion. Unfortunately, he largely failed, maybe because he did not publish it in peer-reviewed scientific journals. Anyway, there was a great silence (18). Independently of Hamilton scientists discussed eugenics (32, 38, 49, 50) and it is present in some evolution textbooks (33, 51). However, even if the effect of modern medicine on the genome were negligible, professionals agree that humans have an exceptionally high spontaneous mutation rate: an average newborn is calculated to have acquired 50 to 100 new mutations in his or her genome and up to seven non-synonymous mutations in exons (55).

Who cares about future genomes?
Who cares about future genomes? Patients with heritable disorders want treatments, and they want it now. Doctors try to make their patients happy. The pharmaceutical industry tries to produce profitable drugs and treatments. Mother's don't want abortions. The only professional groups who potentially do care about the future genome are evolutionary biologists (such as Ronald Fisher and Hamilton) and medical geneticists. Further: economists? (see below) Philosophers? (15) Lawyers? The future quality of the human genome is an abstract idea. It is not a person. It has no rights. It cannot speak for itself. However, the future child could have the right of a healthy genome. Especially, when the future child is a fetus in the womb of a mother. When prenatal whole-genome analysis is possible and the parents refuse to do the test, the rights of the future child could be violated. So, the 'future human genome' could be defined as the total of all future children.

The economic costs of a degenerating genome
The economic costs of disease include not only the maintenance of hospitals, medical departments at universities, but also development, testing and production of drugs by the pharmaceutical industry and the environmental costs of waste disposal of the pharmaceutical industry (23). The US National Institutes of Health (NIH) invests over $31.2 billion annually in medical research for the American people (24). In 2006, health care costs In the USA reached 16% of the nation's gross domestic product, on a path to reach 20% by 2016 (27). The USA spends the largest amount per person as well as perc. of GDP on healthcare of all countries in the world (78). Developing a drug is a costly gamble. Getting one to market takes, on average, more than ten years and a billion dollars. It is true that all this creates jobs, but money spent on medical care cannot be spent for example on education.
The costs of genetic disease could be an argument in favour of eugenics, as Hamilton seems to suggest (20). Calculation of cost-effectiveness of genetic testing of newborns is ethically acceptable for genetic diseases that can be (somatically!) treated (48). It is difficult to get data about costs of genetic disease, but some related data are:

The total economic impact of premature death and disability from cancer worldwide was $895 billion in 2008 (11).
Congenital mental retardation afflicts about 51,000 children annually in the USA; the Centers for Disease Control and Prevention (CDC) estimate that each afflicted child will cost the US economy $1 million over the course of his or her life–that is, a collective cost of $51 billion (54).
Preterm birth is a significant cost factor in healthcare.
Europe: in a typical year, about 165 million people — 38% (!) of the total population of 30 European countries — will have a fully developed mental illness (28). Brain disorders cost Europe almost €800 billion (US$1 trillion) a year – more than cancer, cardiovascular disease and diabetes put together (59). The propertion that can be attributed to genetic conditions is unspecified or unknown.
The lifetime medical cost for a person with spina bifida was estimated to be $460,923 and nonmedical costs $56,511 in 2009 in the USA (56)
VX-770-Kalydeco, a drug that helps to reduce the symptoms of the hereditary disease CF (see above), costs $294,000 a year for the twice-daily pill, to be taken for a lifetime (82). If all 1200 CF (G551D) patients would use VX-770-Kalydeco it would cost $352,800,000 per year or $3,5 billion per 10 year. But those patients are only 4% of the CF population in the US.
During the past decade, the US federal government has spent about US$1 billion researching the genetics of autism and only about $40 million on studies of possible environmental factors (67).
China is contributing $470 million to the Human Variome Project, an international effort to catalog gene variations affecting human health.
According to Statistics Netherlands the total costs spent on health and welfare in the Netherlands were 74.4 billion euro in 2007. In 2010 an estimated 87.6 billion was spent (79).

Hamilton speculated what would happen to people whose lives depend on drugs, when -as a consequence of a big disaster- no therapeutic drugs could be produced anymore. This is not completely fantasy. The costs of helping disabled people should not be a big problem for rich countries, but even in rich countries disabled people have hard times during economic recession or financial crisis because governmental help is reduced (as happens in my country) (22). Furthermore, enough doctors must be present in a country. Romania has a major crisis in the medical system due to the lack of personnel, as 1,700 doctors left the country in the year 2011 (70). See also: Global burden of disease (wikipedia).

Can the dilemma be solved? The new eugenics
The dilemma can be solved if we don't sacrifice the individual (the benefits of medical care) and still prevent the accumulation of harmful mutations. In my view, a rational approach to eugenics could be:

we should forbid mutagenic (genotoxic) substances in our environment. Example: radioactivity. See also: euthenics.
voluntary prevention of the birth of humans with known genetic diseases (genetic diseases run in families). This is already happening: prenatal diagnosis, preimplantation genetic diagnosis (PGD), newborn screening, genetic counseling (47). Currently, more than 1000 genetic tests are available from testing laboratories. Example: because of genetic tests the number of children born with Tay Sachs in the United States have been reduced by 90% (57). Today it is economically feasible to test 448 severe recessive childhood diseases (58), and the number is increasing (64). A requirement for testing is that the mutations are known to cause disease. Hamilton claimed that prenatal diagnosis and germline gene therapy have little effect on the human genome. But what else can we do to reduce mutations in the human genome?
Avoid having children at a later age (for mothers the age is 31, 71). Reason: there is an increase in the frequency of chromosomal abnormalities in newborn children as a function of maternal and, to a lesser extent, paternal age (72).
Repair the human genome. Gene therapy seems to be the method of the future (13),(14). However, sofar all medical treatment, including gene therapy, is directed at the body of the individual and does not repair the germline. However, only germline gene therapy is eugenic (35). On the other hand, treatment of the germline does not cure the individual.
Gene therapy. More than a thousand proof-of-principle clinical studies have been done around the world, and dozens have shown positive results, but as yet no form of gene therapy has been approved for routine use (81). According to the WHO gene therapy holds great promise for treating cystic fibrosis (CF). Others (10) see gene therapy for CF as a failure and invest in phenotypic therapy (!). In general gene therapies for monogenetic disorders other than blood disorders have had limited success so far (12). Gene therapy has shown lasting benefits for the immune systems of a number of children with a rare and fatal immune-deficiency disease (SCID, Wiskott-Aldrich syndrome) (29); patients with β-thalassemia, hemophilia, some eye diseases, and a devastating brain disorder called Metachromatic Leukodystrophy (61).
However, all these cases are necessarily somatic gene therapies because they help the patient, not the descendants of the patient. A solution that benefits both the incipient patient and its children would be genetic therapy of the fertilised egg (before the first division). This is sofar only an option if the mother has a mitochondrial disease (80). Repairing the genome by germline gene therapy is prohibited by law in many countries. Maybe in the far future synthetic biology could reengineer the human genome (42).
enhancing natural levels of DNA-repair (proofreading). In stead of targeting specific mutations, a more universal method could be to enhance the natural level of DNA-repair (19). All organisms have repair mechanisms, but the bacterium Deinococcus radiodurans, which resembles E. coli in physiological terms, yet has DNA repair mechanisms of far greater efficacy and represents the most radiation-resistant organism known. (for example: many 'redundant' copies of the repair-genes plus additional mechanisms that help the repair process). We could learn something from this bacterium and introduce it into the human genome. Mutations that are certainly important to prevent are mutations in genes that are involved in DNA replication or repair because they cause mutations in other genes. Furthermore, the germline should be protected from the mutagenic effects of Transposable Elements (LINE-1 and Alu elements) (63). Cellular processes that control retrotransposon activity should be improved (73). Also, prevention of the generation of reactive oxygen species would be useful.

Should genetic health have priority over phenotypic health?

The above world health map shows that health is determined by geographic, political and economic factors. Health has national borders. It is not likely that national differences are caused by genetic differences (4,5). As long as these factors cause huge inequalities in health and as long environmental and life-style factors (for example: one out of five Americans smoking; or smoking and alcohol during pregnancy, 62) damage health, would it make sense to give priority to genetic health? Furthermore, viewed from a short term perspective, improving genetic health is a very inefficient, slow and indirect way to improve phenotypic health. Apart from good genes, phenotypic health requires food, water, shelter, clothing and protection against pathogens in the environment. Without all that, we would not be able to profit from genetic health. But without genetic health, all those environmental factors are useless. We need both.

However, here is the dilemma again: according to Hamilton's eugenic argument, the genetic health of poor developing countries (Africa, Asia, Latin America) is in fact better than in the developed countries because natural selection is not hindered by modern medicine. So, we should not give medical help to the developing countries? Our medical help would be bad for the genomes of the poor people. Following Hamilton's eugenic argument to its logical conclusion, the goals of the World Health Organization are completely wrong. Following Hamilton's argument, medical care in poor countries should not be stimulated, and in the rich countries it should be stopped; all medical care should be eradicated and natural selection should do its job; that would be good for the human genome. But that would set us back a few hundred years in history, to a time when there was no effective medicine available. But modern medicine brought us health and increasing lifespans. To abandon that would be the end of civilization. There is the dilemma again. It won't go away.

Conclusion

Conclusion rewritten
11 Oct 2011

Main conclusions:

The main problem is the high spontaneous mutation rate in humans
and the ever decreasing levels of natural selection (caused mainly by medical science)
the inescapable result is that our mutational load is increasing
we cannot correct this by technological fixes

Hamilton could have given a far more scientific account of the problem. It's amazing that he allows for a large range (at least 50x) in the rate of mutation: somewhere between 0.1 and 5 fitness-lowering mutations per genome per generation. We need better estimates of the real mutation rate. According to the latest findings there are on average 31 deleterious mutations per individual (39). That is far more than Hamilton's estimate. His worries would become only deeper.
Hamilton pointed out the dysgenic effects of medicine and he also argued for infanticide. Still, Michael Ruse's short statements in the documentary are unfair. Hamilton did more than that. His worries about the future of the human genome were based on the genetical theory of evolution.
It is understandable that Hamilton was pessimistic about the future of the human genome. However, we can now be a little bit more optimistic because techniques to detect mutations in human embryos have tremendiously improved beyond everything Hamilton could have dreamed of. We know now our mutations in unprecendented detail (whole genome diagnosis). Improvement of these detection techniques is almost our only reason for optimism.

Calculating economic costs of hereditary disease seems disrespectful for the unfortunate people with hereditary diseases, but costs are the only thing that economists and politicians understand. In order to calculate global burden of hereditary disease we need data about the number of people with hereditary diseases. The WHO made a crude estimate (17). How does it compare to infectious disease and other environmental diseases? Furthermore, we need data about the extent of Hamilton's problem. What are the effects of medical treatment on the reproductive success of patients with hereditary diseases? Does medical treatment really cause an increase in genetic diseases? If so, how much? How does it compare to spontaneous mutations? What percentage of genetic disease in the population can be prevented on a voluntary basis and how much is in fact prevented? Is it realistic that we can reduce the amount of harmful mutations in the human population?(44).

Hamilton is fully aware of the eugenic problem, but not the dilemma it creates. I guess, this is because he only sees the disadvantages of modern medicine, not its triumphs. Without modern medicine we would have no civilization, no culture, no science, no evolutionary biologists (!), because an epidemic would destroy two-thirds of the world population and we would die at 30 years. So, he misses the dilemma. The WHO and nearly everybody else only see the advantages of modern medicine, but are blind to the damage modern medicine is doing to our genome (17). Nobody sees the full depth, the inevitability and the nastiness of the dilemma. We can close our eyes, but the dilemma won't go away.

Notes

Adam Curtis All watched over by machines of loving grace, part 3.
Narrow Roads of Gene Land. The collected papers of W. D. Hamilton. Volume 1-3. W. H. Freeman: Volume 1 Evolution of Social Behaviour (1996), Oxford University Press: Vol 2 Evolution of sex (2001), Vol 3 Last Words (2005) (edited by Mark Ridley).
"During the second expedition (...) Bill contracted malaria in the Congo, collapsed with haemorrhaging soon after returning to London, and without fully recovering consciousness a few weeks later, at the age of only 63." (page 445, Narrow Roads Vol 3) (biography by Alan Grafen).
On the other hand there is no map of the worldwide distribution of genetic diseases. The UC Atlas of Global Inequality mentions 'Non-communicable diseases' (heart disease and cancer) in the glossary, but there is no map.
It is unlikely that national differences in lifespan are genetical. The life expectancy at birth of the world is 67.2 years, from 39.6 (Swaziland =40% below world average!) to 82.6 years (Japan). See: wikipedia. Additionally, changes in lifespan are too fast to be genetical, see: wikipedia. Only a few genetic diseases (Thalassaemia, Sickle cell anemia, Tay sachs disease) have a specific geographic distribution (WHO).
David Haig (2003) 'The science that dare not speak its name', Quarterly Review of Biology 78: 327-335. Free pdf. The second part of this review is about eugenics. One of the very few reviews about Hamilton's eugenic views I know of.
WHO seems unaware of the dysgenic effects of helping CF patients to become fathers: "CF also can cause reproductive problems - more than 95 percent of men with CF are sterile. But, with new technologies, some are becoming fathers. Although many women with CF are able to conceive, limited lung function and other health factors may make it difficult to carry a child to term." (source). As long as having children is part of the quality of life, the dilemma is unsolved.
Tollånes MC, Rasmussen S, Irgens LM. Caesarean section among relatives, Int J Epidemiol. 2008 Dec;37(6):1341-8. Epub 2008 Jun 7. This publication appeared after Hamilton died. Caesarean section rates are increasing: A mother born by caesarean section had a 55% increased risk of having her first child by caesarean section A female-to-female familial predisposition to caesarean section was observed. It could be caused by biologic inheritance, primarily working through maternal alleles and/or environmental factors. The results imply that both mechanisms could be important. Conclusion: "it represents an economic burden to society". So Hamilton was right in that respect.
A pair of spectacles is a medical device that prevents natural selection. Just as power-assisted-steering in cars, central heating and air conditioning.
Steven Rowe, J.P. Clancy, Eric Sorscher (2011) A Breath Of Fresh Air. Scientific American August 2011. Further: Strachan and Read (2011) state "Until recently, virtually nobody with CF lived long enough to reproduce" (Human Molecular Genetics, p. 88).
THE GLOBAL ECONOMIC COST OF CANCER Please compare with costs of tsunami in Japan in 2011.
Tom Strachan, Andrew Read (2011) Human Molecular Genetics, p. 712.
Verma IM, Weitzman MD. (2005) Gene therapy: twenty-first century medicine. Annu Rev Biochem. 2005;74:711-38.
Timothy P. O'Connor & Ronald G. Crystal (2006) Genetic medicines: treatment strategies for hereditary disorders, Nature Reviews Genetics 7, 261-276 (April 2006)
Antina de Jong et all (2011) Advances in prenatal screening: the ethical dimension, Nature Reviews Genetics 12, 657-663 (September 2011). The authors discuss 'The autonomy rights of the future child', but this is about individuals and pregnant mothers, and is not generalized to the human genome of the future.
"I predict that in two generations the damage being done to the human by the ante- and postnatal life-saving efforts of modern medicine will be obvious to all and be a big taling point of science and politics." (Vol 2, xlvii, Preface).

It would be wrong to say that the WHO is unaware of genetic diseases. See table:

genetic condition	estimated number of cases
Down Syndrome	Each year approximately 3,000 to 5,000 children are born
Thalassaemia	Iran: about 8,000 pregnancies are at risk each year
Sickle cell anemia	millions
Haemophilia	5400 people in the UK with haemophilia A and about 1100 with haemophilia B
Cystic Fibrosis	European Union 1 in 2000-3000 new borns is found to be affected by CF
Tay sachs disease	from 1 : 27 to 1 : 250
Fragile X syndrome	1 in 3600 males and 1 in 4000 to 6000 females with full mutation worldwide
Huntington's disease	about 5 - 7 people per 100,000
Cancer	over 4.2 million deaths
Diabetes	about 180 million
Cardiovascular Disease	16.6 million deaths
Asthma	20.3 million people currently have asthma
data from: Genes and human disease (WHO). (8)
Angelman syndrome	1 in 15,000 live births (77) or about 66 per million live births

WHO Summary:
- 7 million children around the world are born annually with severe genetic disorders or birth defects.
- 90% of infants born with genetic disorders are found in developing countries, contributing significantly to global child mortality. (factsheet Human Genetics programme WHO). I did not find the economic burden of genetic disorders. Disabilities is reported but not genetic. Here is information from the WHO: Amniocentesis and chorionic villus sampling for prenatal diagnosis.

Apart from Alan Grafen and David Haig, nobody discusses the eugenic views of Hamilton. For example the encyclopadic Evolution: The First Four Billion Years edited by Michael Ruse, does contain an entry about Hamilton written by Marlene Zuk, but nothing about eugenics. The Wikipedia article mentions eugenics without explaining. The eugenics wikipedia article does not mention Hamilton (assessed aug 2011).
We can learn from patients with DNA repair-deficiency disorders.
"...it is obvious that each wheelchair or bed-bound victim must add to the burden of those fit and tax-paying individuals who ultimately have to provide for them to survive". p.475 Vol 2 Narrow Roads. (Not completely true, because there is also money from charity.)
Exceptions: there is not always a sharp boundary between genetic and non-genetic diseases. For example, the cause of asthma may include: genetic heredity, lifestyles, smoking, pollution, and viral infections. Finally, "some kinds of insanity can produce better leadership" as is argued in Nassir Ghaemi (2011) A First-Rate Madness: Uncovering the Links between Leadership and Mental Ilness. An example is Winston Churchill. This seems an exception to me, many leaders come to mind which are really mad, bad and dangerous. Artists like Vincent van Gogh suffered from mental illness.
In my country the government does economize on special education, for example for children with learning disabilities such as dyslexia. Dyslexia is partly genetic (source). Also reduction of money going to the support of mentally handicapped people. See also: Tulisa: My Mum and Me about parents with mental health problems.
Discharges from pharmaceutical factories contaminate rivers on three continents (Nature 18 Aug 2011).
Here is a very impressive, extensive overview of NIH-funding per type of disease per year (2007-2012). Please note that the categories: Gene Therapy, Gene Therapy Clinical Trials, Genetic Testing, Genetics together have a budget of 8,1 billion dollars (Estimated 2012). This is apart from the vague category 'Clinical Research' the biggest budget! One has to add many diseases listed with a genetic basis. Then there is 'economic damage' of disease, which is not included. For example: "Schizophrenia alone costs the United States tens of billions of dollars each year." (Nature 27 Oct 11). 27 Oct 2011
About 3% of Down syndrome cases are heritable (familial Down syndrome). Since most cases of Down are not hereditary, eugenics does not make sense. An increase in life expectancy by better medical care is not an eugenic problem because most Down syndrome patients are infertile.
Those who think Hamilton is exceptionally, consider that "between 2 and 3 million unwanted dogs are euthanized in animal shelters each year" and "in 1970, 23 million dogs and cats were euthanized in animal shlters in the United States" according to Hal Herzog (2010) 'Some we love, some we hate, some we eat'.
Thomas R. Insel (2008) Assessing the Economic Costs of Serious Mental Illness, American Journal of Psychiatry 165:663-665, June 2008 (editorial):
"Unlike other medical disorders, the costs of mental disorders are more "indirect" than "direct." The costs of care (e.g., medication, clinic visits, or hospitalization) are direct costs. Indirect costs are incurred through reduced labor supply, public income support payments, reduced educational attainment, and costs associated with other consequences such as incarceration or homelessness. Another kind of indirect cost results from the high rate of medical complications associated with serious mental illness, leading to high rates of emergency room care, high prevalence of pulmonary disease (persons with serious mental illness smoke 44% of all cigarettes in the United States), and early mortality (a loss of 13 to 32 years)", "The $317 billion estimated economic burden of serious mental illness excludes ...". Please note: the author talks about more efficient treatment, but it never occurs to him that the real problem is the degenerating human genome.
Brain burdens, Nature 477, 132 08 September 2011. "A good measure of disease burden is the disability –adjusted life year (DALY) – the person-years lost in a population owing to disability and shortened life. The authors establish brain disorders – both mental and neurological – as the greatest health burden on the population, comprising 23.4% of all DALYs among men and 30.1% for women". The bad news is that the Nature article does not discuss what the genetic contribution to mental disorders is, in other words: the eugenic problem. 8 Sep 2011
Long-term fix for SCID kids, Nature 477, 8-9 01 September 2011. 8 Sep 2011
The reason for artificial insemination is not always infertility, another reason is that more women choose to have babies on their own. "No one knows how many children are born in this country [USA] each year using sperm donors. Some estimates put the number at 30,000 to 60,000, perhaps more" (source). Young adults born by AI are worried about unknowingly having children with a AI half-brother or -sister, and they are worried about having children with genetic diseases, but eugenic worries are not yet among them. The eugenic worries are: (1) natural selection on the natural form of coitus is eliminated, and this could result in disappearance of for example orgasm; (2) infertility is transmitted to the next generation. Potentially, AI could have positive eugenic effects if donors are selected for good health 11 Sep 2011
European Science Foundation (2010) 'Male Reproductive Health. Its impacts in relation to general wellbeing and low European fertility rates', Sep 2010. "It has been estimated that more than 7% of all children born in 2007 in Denmark were conceived by use of ART, which includes in vitro fertilisation, intracytoplasmic sperm injection (ICSI), and intrauterine insemination". Remarkably, altough the ESF is worried: "Dependency on ART would dramatically influence society, since only limited resources are available for state-supported healthcare" (p. 3), they seem only worried about who is going to pay for ART, not about the dysgenic effects of ART itself. The ESF ascribes the increasing use of ART to preventable environmental causes of infertility (because the increase is too rapid to be genetic). On page 7 the report says: "It will be of great importance to ensure that the sperm used for ICSI or IVF procedures is as well selected in terms of DNA integrity as under natural conception" which seems to show eugenic awareness, but they fail to explain how to ensure genetic integrity of sperm of men with fertility problems. Furthermore, since their goal is to treat male reproductive disorders (p. 7), which is dysgenic, I doubt whether they are fully aware of the eugenic problems of ART.
Dudley Kirk (1966) 'Demographic factors affecting the opportunity for natural selection in the United States', Biodemography and Social Biology, Volume 13, Issue 3, 1966. "This amazing reduction in mortality, clearly due to environmental factors, has given rise to concern that relaxation of selective pressures may be enabling the survival of the unfit and deterioration of the genotype". Also published in: Eugenics quarterly. 13 (3): 270-273; 1966.
Scott Freeman and Jon Herron (2007) Evolutionary Analysis has a very good section on 'Compulsory sterilization' starting on page 208.
Ariel Fernández, Michael Lynch (2011) Non-adaptive origins of interactome complexity, Nature 474, 502-505 (23 June 2011)
Germline therapy is very risky because an error is propagated to all future generations. Therefore, it has not been done in humans. For a discussion see Box 21.6 'The ethics of germ-line gene therapy' in 12. In the mouse a human artificial chromosomes (HAC) has been successfully used to deliver the gene Duchenne muscular dystrophy (DMD) to muscle cells. Hamilton was very sceptical about HACs. Source: Francesco Saverio Tedesco (2011) Stem Cell–Mediated Transfer of a Human Artificial Chromosome Ameliorates Muscular Dystrophy, Sci Transl Med 17 August 2011: Vol. 3, Issue 96, p. 96ra78.
In general: if basic processes in the early embryo such as DNA replication and ATP synthesis are damaged by mutation the embryo dies, which is natural selection. Only embryos with low reactive oxygen species (ROS) leak make it through embryonic development. Nick Lane (2011) 14 Oct 2011
Genetic Heterogeneity of Deafness, Marriages among the Deaf (Gallaudet University) 18 Sep 2011
James F. Crow (1997) The high spontaneous mutation rate: Is it a health risk? PNAS August 5, 1997. This free article is based on a public lecture of population geneticist James Crow at the National Academy of Sciences, November 14, 1996. It is an important article. It is about eugenics without mentioning the word. He concludes:
"I do regard mutation accumulation as a problem. It is something like the population bomb, but it has a much longer fuse. We can expect molecular techniques to increase greatly the chance of early detection of mutations with large effects. But there is less reason for optimism about the ability to deal with the much more numerous mutations with very mild effects. But this is a problem with a long time scale; the characteristic time is some 50–100 generations, which cautions us against advocating any precipitate action. We can take time to learn more. Meanwhile, we have more immediate problems: global warming, loss of habitat, water depletion, food shortages, war, terrorism, and especially increase of the world population. If we don't somehow reduce the global birth rate to a sustainable level commensurate with economic viability, we won't have the luxury of worrying about the mutation problem."
Crow recognizes that for the past few centuries natural selection has been greatly reduced in wealthy nations by 'rapid environmental improvements'. As a consequence harmful mutations have been accumulating. Crow is especially worried about the numerous mutations with very mild effects. This results in health risks (headaches, stomach upsets, depressed periods). But humans can improve their environment, so mutations are masked. But this depends on wealth. What if we can't afford it economically anymore? These are essentially Hamilton's worries! Crow certainly is worried about the future of the human genome, and he would reduce the mutation rate to zero if he could. However, he does not specify the causes that reduce natural selection. So, the dilemma of Janus-faced modern medicine is completely out of his sight. Also, he thinks that mutations are a problem of the far future. The word 'spontaneous' in the title is a little misleading if the problem is caused by improved medical technology. I agree with his statement about the more immediate problems. 21 Sep 2011.
Alexey Kondrashov (2012) shows that Crow comfirms Hamilton's worries ('due to advances in medicine'!):
"Crow was very concerned about the negative impact on human health of mildly deleterious alleles that, due to advances in medicine, can accumulate almost unchecked by natural selection. Individually, these may do little – causing a slight increase in blood pressure, for instance. But cumulatively their effect could prove fatal. Unlike some other prominent geneticists, Crow was always careful to separate science and policy and never advocated simplistic approaches to dealing with this problem." Alexey Kondrashov (2012) 'James Crow (1916–2012)', Nature, 481, 444 26 January 2012.
David N. Cooper et al (2010) Genes, Mutations, and Human Inherited Disease at the Dawn of the Age of Personalized Genomics, Human Mutation, Vol. 31, No. 6, 631–655, 2010. 21 Sep 2011
Prader-Willi syndrome: boys have a very small penis; may be corrected with testosterone. Low levels of sex hormones may be corrected at puberty with hormone replacement (source). No discussion of the possibility that successfull treatment could enable patients to have children. 22 Sep 2011.
"Tumours removed, joints replaced, organs transplanted: every weekday, 85,000 non-emergency operations take place in the United States alone", review of Invasion of the Body: Revolutions in Surgery, Nature 22 Sep 2011. 22 Sep 2011
"Scientists have begun to overhaul a yeast's genome to make it more stable, engineerable and evolvable. Remarkably, the part-natural, part-synthetic yeast cells function and reproduce without obvious ill effects". Nature 477, 413–414 22 Sep 2011
A few HIV-infected individuals (dubbed 'elite neutralizers') do develop antibodies that potently nullify diverse HIV-1 strains. So, if this has a genetic basis, these individuals are genetically immune to HIV and can be expected to survive HIV and have children. Nature 22 Sep 2011.
Jon W. Gordon (1999) Genetic Enhancement in Humans, Science Vol. 283 no. 5410 pp. 2023-2024 26 March 1999. 22 Sep 2011. Gordon has a very pessimistic view about genetic enhancement: "Thus, any effort to enhance the human species experimentally would be swamped by the random attempts of Mother Nature.". When applied to evolution it would follow that no positive mutation could become fixed in the population and the theory of evolution would be undermined. 27 Sep 2011
In the Netherlands neonatologists do not treat premature babies born before the 26th week, in other countries neonatologists treat premature babies of 23 - 24 weeks. 25 Sep 2011.
The discovery of antibiotics is more than 70 years ago. This is about 2 - 3 generations. Therefore, the evolutionary effect on the human genome will be very small. However, the effect on the genomes of microbes is very strong (antibiotic resistance!). 26 Sep 2011
The field of genetic counseling is too complex to summarize here. A few notes: Genetic counseling can have eugenic effects or it could have exactly the opposite effect according to Walter Fuhrmann and Frierich Vogel (1983) Genetic Counseling Third Edition, p.162. 26 Sep 2011. Peter Harper and Angus Clarke (1997) Genetics, Society and Clinical Practice, distinguish two main paradigms of prenatal screening: 'genetic cleansing' ("aimed at preventing the birth of affected individuals in the most cost-effective manner" which could be said to be eugenic); and 'informed choice' ("aimed at maximizing client autonomy by facilitating informed choice") (p. 125). 27 Sep 2011
Cost-effectiveness: the costs of screening and early treatment of newborns are less than no screening and late treatment of the disease. Hamilton would have dismissed this as dysgenic, I suppose, because the individual is treated somatically. Prenatal screening could be eugenic. The literature on cost-effectiveness is vast. For criticism see: T. G. Ganiats (1996) Justifying prenatal screening and genetic amniocentesis programs by cost-effectiveness analyses: a re-evaluation, Med Decis Making 1996 Jan-Mar;16(1):45-50. 28 Sep 2011
Julian s. Huxley (1936) 'Eugenics and Society', reprinted in: Carl J. Bajema (1976) Eugenics Then and Now:
"But in civilized human communities of our present type, the elimination of defect by natural selection is largely (...) rendered inoperative by medicine, charity, and the social services. (...) The net result is that many deleterious mutations can and do survive." (p.263)
Hamilton would agree. 28 Sep 2011
H. J. Muller wrote many eugenic articles (largely ignored in wikipedia), see: Carl J. Bajema (1976) Eugenics Then and Now. Famous is his 1949 article: 'Our Load of Mutations' (not mentioned in wikipedia). 28 Sep 2011
Monroe Strickberger (2000) Evolution, third edition, p.617-628 and fourth edition (2008). (good discussion of eugenics) 28 Sep 2011
Essential medicines: drugs needed to satisfy all basic human needs. This is similar to 'essential amino acids'. However, essential medicines are worrying: it means we are all sick and cannot live without medicines. The number of drugs on the WHO list of essential medicines has nearly doubled, from 186 in 1977 to 320 in 2002. The increase is worrying too. 29 Sep 2011
Randolph Nesse, George Williams (1996) Why we get sick (see here) are clearly against eugenics, but are also blind to the degeneration of the human genome and the possible role of modern medicine in that process. Examples: treatment of PKU (p.106), "injections of extra oxytocin have stopped excessive bleeding and saved thousands of lives" (p.201) (my empahsis). It does not occur to them that treatment prevents natural selection. 1 Oct 2011
Armand Leroi (2006) The future of neo-eugenics, EMBO reports (2006) 7, 1184 - 1187. 2 Oct 2011 See also his book Mutants. On Genetic Variety and the Human Body (2003): "our health and happiness are being continually eroded by an unceasing supply of genetic error." (page 18). 18 Oct 2011
Lisenka E L M Vissers et al (2010) A de novo paradigm for mental retardation, Nature Genetics 42, 1109–1112 (2010) 4 Oct 2011
Spina Bifida. Economic Cost (CDC). See also Birth Defects page. (5 Oct 2011)
David B. Goldstein (2011) Growth of genome screening needs debate, Nature 476, 27–28 (04 August 2011) (5 Oct 2011)
Bell CJ et al Carrier testing for severe childhood recessive diseases by next-generation sequencing, Sci Transl Med. 2011 Jan 12 (5 Oct 2011)
Kerri Smith (2011) Trillion-dollar brain drain, Nature News 6 Oct 2011 (6 Oct 2011)
I found this also in John Harris (2007) Enhancing Evolution. The ethical case for Making Better People, p. 79. However, I don't like 'Making Better People' and 'enhancing', we are only in the phase of correcting errors in DNA. Otherwise, recommended reading. (8 Oct 2011)
Jocelyn Kaiser (2011) Gene Therapists Celebrate a Decade of Progress, Science 7 October 2011 (9 Oct 2011)
The World Health Organization (WHO) lists indoor air pollution (IAP) from primitive household cooking fires as the leading environmental cause of death in the world, as it contributes to nearly 2.0 million deaths annually –more deaths than are caused each year by malaria. (14 Oct 2011)
Sergio Lukic, Kevin Chen (2011) Human piRNAs Are Under Selection in Africans and Repress Transposable Elements, Mol Biol Evol (2011) 28 (11): 3061-3067 (24 Oct 2011)
Eleanor Raffan, Robert K. Semple (2011) Next generation sequencing–implications for clinical practice, Br Med Bull (2011) 99 (1): 53-71.
"Stephen Kingsmore at Children's Mercy Hospital in Kansas City, Missouri, argues that clinical sequencing should be limited in scope. He advocates sequencing just what he calls the Mendelianome, the genetic regions known to be involved in inherited diseases. His group is developing methods that use a panel of mutations associated with just over 600 recessive diseases for such screening." Brendan Maher (2011) Human genetics: Genomes on prescription, Nature 478, 22-24 (2011). (7 Nov 2011)
There is a Y-chromosome deletion causing almost complete infertility. This infertility is bound to be passed from father to son (!). However, it is treatable by surgically extracting sperm and micro-injecting it into an egg. This acts strongly against natural selection. Source: Alan Handyside (2010) Let parents decide, Nature 464, 978–979 (15 April 2010) (31 Oct 2011)
However, the resolution of techniques to detect genomic aberrations are becoming better. Array-comparative genomic hybridization begins to look like a genomic Maxwell's Demon. (31 Oct 2011)
Karen Weintraub (2011) The prevalence puzzle: Autism counts, Nature 479, 22–24 (2011) (3 Nov 2011)
Laurent Mottron (2011) Changing perceptions: The power of autism, Nature 479, 33–35 (3 Nov 2011)
Recurrent Early Pregnancy Loss, Genetic Causes - Genetic Abnormalities/Mendelian Disorders, Medscape Reference. (10 Nov 2011)
Medical system in crisis as thousands of doctors leave the country (04.10.11) (14 Nov 2011)
Egbert R. te Velde, Peter L.Pearson (2002) The variability of female reproductive ageing, Human Reproduction Update, Vol.8, No.2 pp. 141–154, 2002. "the frequency of chromosome abnormalities at conception can be expected to rise rapidly with increasing maternal age, such that the majority of embryos are chromosomally abnormal in women approaching 40 years old." (14 Nov 2011)
Thomas B. L. Kirkwood, Steven N. Austad (2000) Why do we age? Nature 408 9 November 2000 (14 Nov 2011)
"Examples of human genetic disorders caused by de novo L1, Alu and SVA insertions continue to accumulate, and 65 cases have been shown to cause heritable diseases, such as haemophilia, cystic fibrosis, Apert syndrome, neurofibromatosis, Duchenne muscular dystrophy, β-thalassaemia, hypercholesterolaemia and breast and colon cancers", from: Richard Cordaux, Mark A. Batze (2009) The impact of retrotransposons on human genome evolution, Nature Reviews Genetics 10, 691-703 (October 2009). (28 Nov 2011)
Michel Tibayrenc (editor) (2011) Genetics and Evolution of Infectious Diseases, Elsevier. (9 Dec 2011)
R. Dolin (2009) How to understand your risk and protect your health (Harvard Health Publications, 2009).
Christine Junge (2011) Morbidity: A personal response, Nature 480, S14–S15 (08 December 2011)
Greg Miller (2011) New Hope for a Devastating Neurological Disorder, Science 23 December 2011.
Wikipedia: "Sexual development is thought to be unaffected, as evidenced by a single reported case of a woman with Angelman syndrome conceiving a female child who also had Angelman syndrome", "General health is fairly good and life-span near average."
Gapminder world: life expectancy and total health spending (US$) per person per country.
rivm: More than a fifth of all costs were spent on mental disorders. This amounts to 15.9 billion euro. A breakdown to specific diseases reveals that mental retardation and dementia are the most expensive single conditions, with 5.6 and 3.5 billion euro. (28 dec 2011)
Ewen Callaway (2012) UK sets sights on gene therapy in eggs, Nature News 24 January 2012
Alison Abbott (2012) French institute prepares for gene-therapy push, Nature News 25 January 2012
Jocelyn Kaiser (2012) New Cystic Fibrosis Drug Offers Hope, at a Price, Science 10 Feb 2012. About the costs: "As more such costly drugs emerge, "there's going to have to be a reality check," Francis Collins says. He anticipates a 'complicated conversation' between companies, insurers, and 'society as a whole' to make the drugs an affordable part of health care."

Reviews

A review of Narrow Roads of Gene Land Volume 2.
Ullica Segerstrale (2013) Nature's Oracle: A Life of W. D. Hamilton, Oxford University Press, USA (April 1, 2013) (not yet published). I guess chapter 'It is a Sick World' deals with eugenics.

guestbook (moderated)	home: Towards the Third Evolutionary Synthesis	http://home.planet.nl/~gkorthof/korthof97.htm
Copyright ©G. Korthof	First published: 25 Aug 2011	Updated: 10 Feb F.R./N: 10 Feb 2012