Article

Estimating inbreeding in large, semi-isolated populations: Effects of varying generation lengths and of migration

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Abstract

The purpose of the study reported here was to investigate two important assumptions used in a recently reported new method of estimating inbreeding in large, relatively isolated populations over historic times. The method, based on modeling the genealogical "paradox," produces values of Pearl's coefficients, Z, a measure of inbreeding or genealogical coalescence, as a function of time. In this study, the effects on inbreeding of two important assumptions made in earlier studies, namely those of using a constant generation length and of ignoring migration, have been investigated for the population of Britain. First, by relating the median age of women at childbirth to the development level of various societies, the variation of the generation lengths for different periods in historic Britain were estimated. Values of Z for two types of varying generation lengths were then calculated and compared with the case of constant generation length. Second, the population curve for Britain used in earlier studies was modified to obtain the subpopulation at any time during the past two millennia that was descended from the pre-Roman British Celts. Values of Z for the case with migration were then calculated and compared with the case for no migration. It is shown that these two assumptions may be taken into account if and when required. Both the effect of a varying generation length and the effect of migration on Z were found to be 20-40%, when no known value of inbreeding was used, and 2-5%, when a known value of inbreeding was used.

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... An interest in consanguineous marriages and percentage inbreeding values, in addition to casual curiosity (Bramwell, 1939), arises from academic curiosity concerning marital mobility, demographics, population structure, etc. (Brennan, 1981;Coleman, 1980;Day 2010;Day & Smith 2013;Robinson, 1983;Smith, 2001), and medical curiosity concerning whether any genetic detrimental effects have occurred in the resulting offspring (Bell, 1940;Bundey et al., 1990;Darwin, 1875; Mitchell, 1866;Pearson, 1908a,b,c). Inbreeding values have also been used to model inbreeding over the past millennium based on the genealogical paradox (Pattison, 2001(Pattison, , 2003(Pattison, , 2004(Pattison, , 2007, which apparently corrects for the effective breeding population in the modelling method used. These few studies represent the little systematic information obtained to date on past levels of inbreeding in Britain (Day & Smith). ...
... Unfortunately these cousin relationships could not be used separately as 'once removed' data in this study as it was not indicated in which of her five time periods they occurred. A generation length of 28 years was used in this study because it has been shown to be more appropriate than 30 years (Pattison, 2007;Day). However, this was not really an issue, as discussed by Smith) because the allocated ranges of generational years are only rough estimates. ...
... Bramwell;Jacquard 1974;Shoumatoff 1985;and Wachter). There have been a number of different approaches to modelling the pedigree collapse involved in the solution to the 'paradox' (Chang 1999;Derrida et al. 1999Derrida et al. , 2000aPattison 2001Pattison , 2003Pattison , 2004Pattison , 2007Wachter) and each has found that there must have been an increasing trend in inbreeding as one moves back in time. ...
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There have been a number of previous estimates of human inbreeding for Britons of British descent in Britain; each generally for different social classes, geographical regions, and/or time periods. In this study an attempt was made to collect all relevant published studies and to combine the results of these disparate studies into an integrated whole for all of Britain. This was achieved by combining weighted means of the percentage of consanguineous marriages (f%) reported in these earlier studies: weighted according to the number of records each author examined, the proportion of social classes or geographical regions covered by the records, and the 'merit' of their individual research methodologies. The percentage occurrences of the various consanguineous marriages, from 1 st to 3 rd cousins, were partitioned into a number of time periods, which allowed the weighted mean percentage inbreeding coefficients (F%) to be obtained as a function of time over the period from 1820 to 1960. The resulting temporal scatter distribution of the weighted F% values closely followed a sigmoidal curve, with a non-linear correlation coefficient of η = 0.974, which fitted well to a generalized logistic function. After about 1900 the value of the weighted F% was essentially constant at about 0.038±0.004, whereas it decreased rapidly from about 0.256±0.011 between 1820 and 1900. The upper-bound value of weighted F%, before 1820, from the fitted logistic function is 0.276. Note that this corresponds to a value of the conventional mean inbreeding coefficient F = 0.00276. As the first known attempt to integrate the earlier disparate values of unweighted F% for Britons of British descent for all of Britain, the results of this analysis are promising and should be useful as reference values in other related studies.
... The common historical inclination of small and isolated populations was to marry those within walking distance (Pattison, 2001). In practical terms, close relatives are likely to live next to somebody, strengthening family ties, to be in the same social class, to share the same religion, culture, tradition, ethnic group, language as well as to be part of the same social circle (Pattison, 2007). This apparent 'genealogy paradox' is also named 'pedigree collapse' (Pattison, 2001). ...
... In fact, a closer look reveals that moderate inbreeding has constantly been the rule, not the exception in human history -some experts assume that up to 80% of all marriages in history have been between second cousins or closer (Pattison, 2007;Conniff, 2003). Some authors, such as Derrida et al. (1999), affirmed, according to their model, that when we look very far in the past, about 80% of the (adult) population appears in the genealogical tree of every individual. ...
... Some authors, such as Derrida et al. (1999), affirmed, according to their model, that when we look very far in the past, about 80% of the (adult) population appears in the genealogical tree of every individual. There are good manuscripts on the subject, relating human biological origins, evolutionary development, inbreeding and genetic diversity, even considering isolated or semi-isolated populations (Pattison, 2001;2007). ...
... Thomas et al.'s (2008) first and principal criticism was on the methodology used for the population reconstruction. I employed the back projection method, which is an established method used by historical demographers for modeling past populations (see Pattison 2007). The population estimates used to construct the historical population curve for Britain were uncertain, but were the best available at the time (Pattison 2003(Pattison , 2004. ...
... Compared to the later medieval period when bastardy was discouraged by the manorial fine of childwyte, it is possible that in Anglo-Saxon times there was a more indulgent attitude (Laslett 1980). Even so, when discouraged in the later medieval period, births outside of marriage were not uncommon, comprising up to 25% of births in some parishes in the pre-plague period, and crossing social boundaries (Pattison 2007). Also, adultery leading to illicit births is hidden within marriages (Laslett 1980). ...
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The genetic surveys of the population of Britain conducted by Weale et al. and Capelli et al. produced estimates of the Germani immigration into Britain during the early Anglo-Saxon period, c.430-c.730. These estimates are considerably higher than the estimates of archaeologists. A possible explanation suggests that an apartheid-like social system existed in the early Anglo-Saxon kingdoms resulting in the Germani breeding more quickly than the Britons. Thomas et al. attempted to model this suggestion and showed that it was a possible explanation if all Anglo-Saxon kingdoms had such a system for up to 400 years. I noted that their explanation ignored the probability that Germani have been arriving in Britain for at least the past three millennia, including Belgae and Roman soldiers, and not only during the early Anglo-Saxon period. I produced a population model for Britain taking into account this long term, low level migration that showed that the estimates could be reconciled without the need for introducing an apartheid-like system. In turn, Thomas et al. responded, criticizing my model and arguments, which they considered persuasively written but wanting in terms of methodology, data sources, underlying assumptions, and application. Here, I respond in detail to those criticisms and argue that it is still unnecessary to introduce an apartheid-like system in order to reconcile the different estimates of Germani arrivals. A point of confusion is that geneticists are interested in ancestry, while archaeologists are interested in ethnicity: it is the bones, not the burial rites, which are important in the present context.
... In these cases, it is unlikely that any resulting child would have taken the identity of the father, as stated byThomas et al. (2006). For instance, in the later Medieval period, births outside of marriage were not uncommon and crossed social boundaries (Pattison 2007). ...
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It has recently been argued that there was an apartheid-like social structure operating in Early Anglo-Saxon England. This was proposed in order to explain the relatively high degree of similarity between Germanic-speaking areas of northwest Europe and England. Opinions vary as to whether there was a substantial Germanic invasion or only a relatively small number arrived in Britain during this period. Contrary to the assumption of limited intermarriage made in the apartheid simulation, there is evidence that significant mixing of the British and Germanic peoples occurred, and that the early law codes, such as that of King Ine of Wessex, could have deliberately encouraged such mixing. More importantly, the simulation did not take into account any northwest European immigration that arrived both before and after the Early Anglo-Saxon period. In view of the uncertainty of the places of origin of the various Germanic peoples, and their numbers and dates of arrival, the present study adopts an alternative approach to estimate the percentage of indigenous Britons in the current British population. It was found unnecessary to introduce any special social structure among the diverse Anglo-Saxon people in order to account for the estimates of northwest European intrusion into the British population.
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