The costs of outbreeding: a few more points

Although the house mouse (Mus musculus) is a single species, fertility is reduced in crosses between geographic populations (Wikipedia - Bolid74)



It's widely believed that mating between two species never produces fertile offspring. Untrue. Interspecific mating sometimes produces fertile offspring, and the genomes of closely related species often show gene flow from one to the other. It would be more accurate to say that such crossings are usually compromised in one way or another—incompatibilities arise during development, which lead to miscarriage, sterile offspring, or less viable offspring. 

There is a similar misunderstanding about mating within a species. It isn't true that mating within a species is uniformly possible. Even below the species level, genetic incompatibilities can arise between different geographic populations. Keep in mind the arbitrariness of all these terms: “geographic population,” “race,” “subspecies,” and “species.” These are points on a continuum of increasing reproductive isolation and genetic incompatibility. 

Various problems have been observed in crossings between geographic populations of the same species (Presgraves 2010; Turner et al. 2011). In the case of crossings between two subspecies of the house mouse (M. m. domesticus and M. m. musculus), the problems have no single cause. There is not only reduced fertility but also reduced immune function, as shown by higher loads of intestinal parasites in hybrids. Even the reduction in fertility has multiple causes: "The type and severity of fertility defects observed depends on the geographic origin of the strains and also varies among individuals within regions. This variability suggests that multiple genetic incompatibilities contribute to hybrid male sterility" (Turner et al. 2011). Multiple loci seem to be responsible, and incompatibilities that cause sterility likely act in combination with each other and with other incompatibilities (Turner et al. 2011).

In general, as genetic distance increases, so does the risk of faulty interactions between different gene loci:

The Dobzhansky-Muller model proposes that populations diverging independently in allopatry will accumulate differences through drift or selection, but will maintain coadaptation between divergent loci within each single population. If secondary contact and hybridization occurs between these populations, these divergent loci from the previously allopatric populations may interact deleteriously in hybrids leading to lowered hybrid fitness (Leppälä and Savolainen 2011)

Icelandic and Danish studies on outbreeding

The above model is consistent with two recent studies on humans, one in Iceland and the other in Denmark. The Icelandic study used a database of couples born between 1800 and 1965. In the authors' own words, "the advantage of using the Icelandic data set lies in this population being small and one of the most socioeconomically and culturally homogeneous societies in the world, with little variation in family size, use of contraceptives, and marriage practices" (Helgason et al. 2008). The authors found that fertility rose and then fell with increasing genetic distance between husband and wife. Specifically, the relationship was:

[...] an n-shaped curve from the relatively low reproductive success of couples related at the level of second cousins or closer, to the maximum for couples related at the level of third and fourth cousins, after which there is a steady decrease in reproductive success with diminishing kinship between spouses. A similar picture emerges when the number of grandchildren per couple is examined (Helgason et al. 2008)

The Icelandic study was criticized in a tweet by Jayman:

Doubt it. The Icelandic study is likely confounded in various ways so I don't think genetic relatedness is the thing directly affecting fertility.

He was retweeted by hbdchick, but neither of them identified the "various" confounds. Nor did I find this criticism in the 140 papers that cited the study on Google Scholar. So we're not in the realm of common knowledge.

The original paper did identify a possible confound. From 1800 to 1965, there was a decrease in mean fertility and an increase in marriages with more distantly related individuals. The relationship between fertility and kinship may thus be an historical confound:

These results are based on couples born during a period of almost 200 years, in the course of which there was a marked decline both in the mean fertility and in kinship between couples. (Helgason et al. 2008)

To eliminate the confound, the authors broke the data down by 25-year intervals:

Nonetheless, the same general relationship between kinship and reproductive outcome was observed within each 25-year subinterval (fig. S2). We evaluated the correlation between the standardized variables of kinship and reproductive outcome for all couples and for each time interval separately (Table 2), adjusting for the impact of geographical differences in the kinship and fertility of couples within Iceland (10). Each test revealed as significant association with kinship, with correlation coefficients of 0.063 (P = 1.5 × 10-129) for the number of children, 0.045 (P = 3.6 × 10-66) for the number of children who reproduced, and 0.042 (P = 7.6 × 10 -58) for the number of grandchildren. (Helgason et al. 2008)

There might be other confounds. That's why the Danish study is important. It went through a number of socioeconomic factors: "education, family income, urbanicity, mother's age at first birth, and six variables representing proximity to kin [maternal radius, i.e., distance between mother's and child's birthplace; paternal radius; and presence of each grandparent in the child's birth parish or neighboring parish" (Laboriau and Amorim 2008). Controlling for those factors did not change the findings.


Davenport's study on outbreeding in Jamaica

Jayman also criticized Davenport's study, apparently in reference to the finding that between 5 and 30% of the "brown" children did worse than predicted.

As for mixed race children, you have to keep in mind that people don't enter those relationships randomly - i.e., also confounded

Jayman is arguing that the parents of the brown children were atypical, apparently in the sense of being below-average. In its early days, Jamaica did have many poor whites, but most of them left because of competition from slave labor. European ancestry entered the Jamaican population largely from landowners and businessmen, particularly people of Scottish descent:

Jamaica has more people using the Campbell surnames than the population of Scotland itself, and it also has the highest percentage of Scottish surnames outside of Scotland. Scottish surnames account to about 60% of the surnames in the Jamaican phone books. The first Jamaican inhabitants from Scotland were exiled "rebels". Later, they would be followed by ambitious businessmen who spent time between their great country estates in Scotland and the island. As a result, many of the slave owning plantations on the island were owned by Scottish men, and thus a large number of mixed-race Jamaicans can claim Scottish ancestry. (Wikipedia 2020).

Today, "brown" Jamaicans are generally middle class, and they were even more so during the time of Davenport's study. It is indeed strange to intimate that his biracial subjects were the offspring of below-average whites and blacks.


Conclusion

Davenport's study probably corresponds to the highest degree of outbreeding possible. Even in that case, adverse mental effects were observable only in a minority of offspring. The most widespread effect is probably lower fertility and perhaps a higher risk of testicular cancer. Joffe (2009) suggests that an increase in outbreeding might explain a century-long decrease in semen quality and a corresponding increase in testicular cancer:

One implication of the proposed pathogenesis is that as D&D [duplications and deletions of genetic material during meiosis] accumulate, mating between individuals who are genetically unalike would be associated with lower reproductive success as pairing at the start of meiosis would be more likely to be impaired. On the face of it, this contradicts the earlier observation that hybrid vigour would tend to increase fertility in unrelated individuals. Yet these two ideas may be compatible: both extremes—genetic similarity (inbreeding) and genetic distance (D&D accumulation)—could decrease fertility, so that an intermediate degree of relatedness would be associated with the highest degree of fertility. This could explain the evidence from Iceland that the greatest reproductive success, measured as the number of grandchildren, was observed in couples who were third or fourth cousins (Helgason et al., 2008). Comparable findings have been reported from elsewhere, for example Denmark (Labouriau and Amorim, 2008).

I realize this may be a sensitive subject. If you feel offended, please read something else.


References

Davenport, C.B. and M. Steggerda. (1928). Race Crossing in Jamaica. Washington: Carnegie Institution, Publication no. 395.
http://www.velesova-sloboda.info/archiv/pdf/davenport-race-crossing-in-jamaica.pdf  

Helgason, A., S. Pálsson, D.F. Guðbjartsson, þ. Kristjánsson, K. Stefánsson. (2008). An association between the kinship and fertility of human couples. Science 319(5864): 813-816.
http://facelab.org/debruine/Teaching/EvPsych/files/Helgason_2008.pdf   

Joffe, M. (2009). What has happened to human fertility? Human Reproduction 25(2): 295-307.
http://citeseerx.ist.psu.edu/viewdoc/download?doi=10.1.1.1003.7270&rep=rep1&type=pdf 

Labouriau, R., and A. Amorim. (2008). Comment on "An Association Between the Kinship and Fertility of Human Couples." Science 322(5908): 1634
http://science.sciencemag.org/content/322/5908/1634.2.full   

Leppälä, J. and O. Savolainen. (2011). Nuclear-cytoplasmic interactions reduce male fertility in hybrids of Arabidopsis lyrata subspecies. Evolution 65(10): 2959-2972
https://onlinelibrary.wiley.com/doi/full/10.1111/j.1558-5646.2011.01361.x

Presgraves, D.C. (2010). The molecular evolutionary basis of species formation. Nature Reviews Genetics 11:175-180.
https://www.nature.com/articles/nrg2718  

Turner, L.M., D.J. Schwahn, and B. Harr. (2011). Reduced male fertility is common but highly variable in form and severity in a natural house mouse hybrid zone. Evolution 66(2): 443-458.
https://onlinelibrary.wiley.com/doi/full/10.1111/j.1558-5646.2011.01445.x  

Wikipedia (2020). Jamaica
https://en.wikipedia.org/wiki/Jamaica

The costs of outbreeding: what do we know?

Charles Davenport, circa 1929 (Wikicommons)



To produce healthy children, you should marry a third or fourth cousin. Farther out, the genetic costs of outbreeding begin to outweigh those of inbreeding. That was what a cohort study found in examining Icelanders born between 1800 and 1964. Fertility was lower if the woman's husband was either closer in or farther out (Helgason et al. 2008).

That finding is supported by a Danish study. Using data on Danes born in 1954, the authors measured the distance between the wife's home parish and the husband's home parish. Fertility peaked at a distance of around 75 km. The relationship between fertility and marital radius was not explained by education, family income, urbanicity, or mother's age at first birth. The authors concluded that their findings were consistent with those of the Icelandic study, the cause being the same: fertility rises with decreasing relatedness up to a peak and then starts to fall. Inbreeding depression gives way to outbreeding depression (Labouriau and Amorim 2008).

How far can you marry out before the costs of outbreeding become unacceptable? One problem lies with that word. What may be perfectly acceptable to one person may not be to another.

So let me review what is known and what remains to be known. You be the judge.

A half-century ago, Ernst Mayr wrote: "Hybridization between species leads almost invariably to unbalance through deleterious combinations of genes. [...] in Drosophila even the hybridization of races may lead to destruction of well-integrated gene combinations" (Mayr 1970, p. 399). He then went on to say that it is "still uncertain" whether these findings apply to our species. In fact, "all investigations of race mixtures in man have failed to produce any evidence of decreased fitness."

It would be better to say "indisputable evidence." In fact, there is evidence from two major studies of the 1920s.


Davenport’s research in Jamaica

I will begin with a study by Charles Davenport, a Harvard zoology professor. In the early years of the twentieth century, he became concerned about human heredity and the genetic consequences of outbreeding. In 1926, with his assistant Morris Steggerda, he went to Jamaica and examined 100 black, 100 white and 100 “brown” adults of equal social status, together with 1200 children between the ages of 10 and 16. They were given anthropometric, physiological, and psychological tests. The psychological results are worth quoting in full:

Disharmonies in the mental sphere are socially more significant, perhaps, than those in the physical sphere, and such disharmonies are apparently common in the adult Browns [...]. Such disharmony and confusion apparently appear in visualization and reproduction, as in putting together the parts of the manikin. The proportion of failures of the Browns is 9.6 per cent, as opposed to 3.1 per cent in Blacks and 2.1 per cent in Whites. In copying of geometric figures 5 per cent of Browns fail completely, as compared with 3 per cent of the Blacks and 0 per cent of the Whites. In the Army Alpha Test IV (opposites and similars) 41 per cent of Browns got only 3.7 or fewer correct, while only 23 per cent of Blacks did so poorly and none of the Whites. In Test V (pied sentences) 30 per cent of the Browns got fewer than 3 out of 24 questions right, while only 26 per cent of the Blacks and 13 per cent of the Whites did. In Test VII (of analogies) 45 per cent of the Browns got less than 10 per cent of the questions correct, while only 30 per cent of the Blacks and 21 per cent of the Whites did so badly. One gains the general impression that, though on the average the Browns did not do so badly, there was among them a greater number of persons than in either Blacks or Whites who were muddled and wuzzle-headed. The Blacks may have low intelligence, but they generally can use what they have in fairly effective fashion; but among the Browns there appear to be an extra 5 per cent who seem not to be able to utilize their native endowment. There are so many variables, however, and the numbers are so small, that the results merely propose an hypothesis and do not warrant a conclusion.

The question arises: are there any traits in which, on the average, the adult Browns are superior to the Whites? We might, theoretically, expect such, yet when we have tested all of the probably genetically distinct traits between Blacks and Whites, we find only one in which the mean of the adult Browns is clearly higher than that of either parental stock.

On the other hand, if we examine the means for children of 10 to 13 or 13 to 16 years there are a few in which the brown children grade higher than either the black or the white children. [...] The fact that youthful Browns sometimes score higher than youthful Blacks or Whites suggests the conclusion that brown children develop in some mental capacities precociously; and then fall behind in development.

The relative underperformance of biracial individuals looks significant, given the sample sizes. Less clear is its magnitude and, most importantly, its cause. If we exclude the hypothesis of incompatibilities during development, there are only two other explanations:

- The “brown” individuals had suffered some kind of deprivation that the black and white individuals hadn't.

- “Brown” individuals with mental problems had been oversampled.

Neither explanation seems likely. In the Jamaica of the 1920s, biracial individuals were if anything a privileged group. They dominated the middle class. Nor is it clear why those with mental problems would be oversampled. 


Mjøen’s research in Norway and Sweden

The second study was by Jon Alfred Hansen Mjøen, a Norwegian who in 1906 founded the Vinderen Biological Laboratory, a center for international research in genetics. He likewise was concerned about the possible negative effects of outbreeding and became a proponent of eugenics. In 1934, he supported a Norwegian law for forced sterilization. He died in 1939. 

With the support of his laboratory, Mjøen did fieldwork in northern Norway and Sweden, where he observed and measured about 600 Norwegians/Swedes, about 600 Sami, and more than 300 people of first-generation mixed ancestry. Most individuals in the last group were described as having "harmonious faces, general good looks, decent stature, height, and strength, as well as a good correlation between volume of lungs and body-size, and muscular strength and body-size." However, "we found more disharmonies, both physical and mental, than in the two parent races."

The disharmonies were: "Relatively large or small ears, disproportionate extremities, unusual length of body in the F1 generation, abnormal range of variation with regard to such characteristics as lung-volume and muscular strength [...], greater prevalence of diabetes [...], loss of balance, and diminished resistance to tuberculosis" (Mjøen 1931).

Mjøen argued that disharmonies are most likely to occur when a trait is determined by large numbers of genes acting together, such as when hormones act on genes for growth and development: "It is highly probable that the frequently observed exaggerated growth of the hybrid and his disproportionately large extremities are due to a glandular disturbance of genetic origin." He also attributed the higher rate of diabetes to errors in hormonal regulation.


After

There were no subsequent studies. It would be easy to say that research ended with the widespread revulsion against Nazi Germany in 1945. Actually, the end came earlier, in the 1930s with the triumph of Boasian anthropology and behaviorist psychology. Both stressed the plasticity of the human mind and shifted the focus of research from nature to nurture, i.e., to "society" and possibilities for social change. 

Nonetheless, even within academia, important people continued to voice support for hereditarian and eugenic positions. John Maynard Keynes served as the director of the Eugenics Society of London (1937-1944) and in 1946 was still calling eugenics "the most important and significant branch of sociology" (Brignell 2010). The science fiction series Star Trek ran an episode in 1967 on eugenics ('Space Seed'). The Eugenics Review would be renamed The Journal of Biosocial Science only in 1969. Eugenic laws remained on the books and were enforced as late as the 1970s in some jurisdictions. 

All of this may seem surprising because there has been an effort to push the systematic rejection of eugenics farther and farther back in time. The Wiki entry on Charles Davenport, for instance, states that "only his ardent admirers" took his research seriously. The truth is that he was widely respected until the 1930s. His 1911 work, Heredity in Relation to Eugenics, became a college textbook (Wikipedia 2020). Mjøen was likewise well regarded:

His fame was such that he was one of three expert advisors consulted by the US Congress in the appointment of the first President of the American Eugenics Society, he gave a lecture tour of eighteen American universities in 1927, and, in a period in which The New York Times voiced anxieties about the future of the 'Nordic Type' in America, the newspaper of record invoked Mjøen repeatedly (Reinert 2015)

Even on the left, the general attitude was cautious acceptance, as seen in this 1928 report from the Eugenic Society of Leningrad:

P. I. Ljublinsky remarked that there was not much to be said for the two extreme views of this problem—the doctrine of complete indifference to the matter and the system of strict measures. Therefore what must be recognised as most advantageous is something lying between the two extremes,—a certain system of compromise, of which he himself was an adherent. The meeting gave its support to this last standpoint. It was decided that certain legislative prohibitions in this field are undoubtedly of service. (Philiptschenko 1928)

On the specific issue of outbreeding, feelings were more mixed:

After the conclusion of the discussion the meeting decided that, although the question of race crossing is very important, it is difficult to regard is as definitely solved. But in contrast to the attitude adopted in the Norwegian programme it must be stated that, so long as we have no other data available, we cannot express an opinion against the crossing of different races and nations. (Philiptschenko 1928)

There would be no more data. The Soviet Union, like the world in general, looked up to progressive thought in the West, particularly in the United States. By the 1930s, the shift toward blank slate thinking had become overwhelming among American intellectuals.


Conclusion

If we go back to the recent studies on fertility and outbreeding, we see that fertility progressively declines when outbreeding takes place beyond one's fourth cousin. A growing proportion of embryos or fetuses fail to develop beyond a certain point, apparently because of some incompatibility.

Outbreeding depression seems to be a real phenomenon, but how serious is it? Davenport examined perhaps the maximum degree of outbreeding possible, and only a minority of individuals showed a degree of dysfunction that was higher than expected. The proportion varied according to the mental test, from a low of about 5% to a high of 30%. It is likely that subsequent generations would show progressively lower proportions, with natural selection removing the least functional individuals through reduced fertility, inability to find a mate, or higher risk of illness. 

Davenport was cautious in interpreting his results, noting the difficulty in estimating the size of the adverse outbreeding effect. His self-criticism was taken up by his critics, and the current prevalent view is that his study has been thoroughly discredited, if not by methodological problems, then by Hitler.


References

Brignell, V. (2010). The eugenics movement Britain wants to forget. New Statesman. December 9 http://www.newstatesman.com/society/2010/12/british-eugenics-disabled 

Davenport, C.B. and M. Steggerda: (1928). Race Crossing in Jamaica. Washington: Carnegie Institution, Publication no. 395.
http://www.velesova-sloboda.info/archiv/pdf/davenport-race-crossing-in-jamaica.pdf

Helgason, A., S. Pálsson, D.F. Guðbjartsson, þ. Kristjánsson, K. Stefánsson. (2008). An association between the kinship and fertility of human couples. Science 319(5864): 813-816.
http://facelab.org/debruine/Teaching/EvPsych/files/Helgason_2008.pdf

Labouriau, R., and A. Amorim. (2008). Comment on "An Association Between the Kinship and Fertility of Human Couples" Science 322(5908): 1634
http://science.sciencemag.org/content/322/5908/1634.2.full

Mayr, E. (1970). Populations, Species, and Evolution. Cambridge (Mass.): Belknap Press.

Mjøen, J.A. (1931). Race-crossing and glands. Some human hybrids and their parent stocks. The Eugenics Review 23(1): 31-40.
https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2984998/?page=1

Philiptschenko, J.A. (1928). The Norwegian eugenic programme discussed at meetings of the eugenic society of Leningrad. The Eugenics Review 19(4): 294-298.
https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2984720/

Reinert, S.A. (2015). The economy of fear:  H.P. Lovecraft on eugenics, economics and the Great Depression. Horror Studies 6(2): 255-281.
https://www.academia.edu/19676748/The_Economy_of_Fear_H.P._Lovecraft_on_Eugenics_Economics_and_the_Great_Depression

Wikipedia. (2020). Charles Davenport.
https://en.wikipedia.org/wiki/Charles_Davenport