Genomic ancestry of North Africans supports back-to-Africa migrations

North African populations are distinct from sub-Saharan Africans based on cultural, linguistic, and phenotypic attributes; however, the time and the extent of genetic divergence between populations north and south of the Sahara remain poorly understood. Here, we interrogate the multilayered history of North Africa by characterizing the effect of hypothesized migrations from the Near East, Europe, and sub-Saharan Africa on current genetic diversity. We present dense, genome-wide SNP genotyping array data (730,000 sites) from seven North African populations, spanning from Egypt to Morocco, and one Spanish population. We identify a gradient of likely autochthonous Maghrebi ancestry that increases from east to west across northern Africa; this ancestry is likely derived from "back-to-Africa" gene flow more than 12,000 years ago (ya), prior to the Holocene. The indigenous North African ancestry is more frequent in populations with historical Berber ethnicity. In most North African populations we also see substantial shared ancestry with the Near East, and to a lesser extent sub-Saharan Africa and Europe. To estimate the time of migration from sub-Saharan populations into North Africa, we implement a maximum likelihood dating method based on the distribution of migrant tracts. In order to first identify migrant tracts, we assign local ancestry to haplotypes using a novel, principal component-based analysis of three ancestral populations. We estimate that a migration of western African origin into Morocco began about 40 generations ago (approximately 1,200 ya); a migration of individuals with Nilotic ancestry into Egypt occurred about 25 generations ago (approximately 750 ya). Our genomic data reveal an extraordinarily complex history of migrations, involving at least five ancestral populations, into North Africa.     

Insights into the demographic history of African Pygmies from complete mitochondrial genomes

Pygmy populations are among the few hunter-gatherers currently living in sub-Saharan Africa and are mainly represented by two groups, Eastern and Western, according to their current geographical distribution. They are scattered across the Central African belt and surrounded by Bantu-speaking farmers, with whom they have complex social and economic interactions. To investigate the demographic history of Pygmy groups, a population approach was applied to the analysis of 205 complete mitochondrial DNA (mtDNA) sequences from ten central African populations. No sharing of maternal lineages was observed between the two Pygmy groups, with haplogroup L1c being characteristic of the Western group but most of Eastern Pygmy lineages falling into subclades of L0a, L2a, and L5. Demographic inferences based on Bayesian coalescent simulations point to an early split among the maternal ancestors of Pygmies and those of Bantu-speaking farmers (～ 70,000 years ago [ya]). Evidence for population growth in the ancestors of Bantu-speaking farmers has been observed, starting ～ 65,000 ya, well before the diffusion of Bantu languages. Subsequently, the effective population size of the ancestors of Pygmies remained constant over time and ～ 27,000 ya, coincident with the Last Glacial Maximum, Eastern and Western Pygmies diverged, with evidence of subsequent migration only among the Western group and the Bantu-speaking farmers. Western Pygmies show signs of a recent bottleneck 4,000-650 ya, coincident with the diffusion of Bantu languages, whereas Eastern Pygmies seem to have experienced a more ancient decrease in population size (20,000-4,000 ya). In conclusion, the results of this first attempt at analyzing complete mtDNA sequences at the population level in sub-Saharan Africa not only support previous findings but also offer new insights into the demographic history of Pygmy populations, shedding new light on the ancient peopling of the African continent.     

Phylogeography, genetic structure and population divergence time of cheetahs in Africa and Asia: evidence for long-term geographic isolates

The cheetah (Acinonyx jubatus) has been described as a species with low levels of genetic variation. This has been suggested to be the consequence of a demographic bottleneck 10 000–12 000 years ago (ya) and also led to the assumption that only small genetic differences exist between the described subspecies. However, analysing mitochondrial DNA and microsatellites in cheetah samples from most of the historic range of the species we found relatively deep phylogeographic breaks between some of the investigated populations, and most of the methods assessed divergence time estimates predating the postulated bottleneck. Mitochondrial DNA monophyly and overall levels of genetic differentiation support the distinctiveness of Northern‐East African cheetahs (Acinonyx jubatus soemmeringii). Moreover, combining archaeozoological and contemporary samples, we show that Asiatic cheetahs (Acinonyx jubatus venaticus) are unambiguously separated from African subspecies. Divergence time estimates from mitochondrial and nuclear data place the split between Asiatic and Southern African cheetahs (Acinonyx jubatus jubatus) at 32 000–67 000 ya using an average mammalian microsatellite mutation rate and at 4700–44 000 ya employing human microsatellite mutation rates. Cheetahs are vulnerable to extinction globally and critically endangered in their Asiatic range, where the last 70–110 individuals survive only in Iran. We demonstrate that these extant Iranian cheetahs are an autochthonous monophyletic population and the last representatives of the Asiatic subspecies A. j. venaticus. We advocate that conservation strategies should consider the uncovered independent evolutionary histories of Asiatic and African cheetahs, as well as among some African subspecies. This would facilitate the dual conservation priorities of maintaining locally adapted ecotypes and genetic diversity.     

Phylogeny and evolutionary history of the silkworm

The silkworm, Bombyx mori, played an important role in the old Silk Road that connected ancient Asia and Europe. However, to date, there have been few studies of the origins and domestication of this species using molecular methods. In this study, DNA sequences of mitochondrial and nuclear loci were used to infer the phylogeny and evolutionary history of the domesticated silkworm and its relatives. All of the phylogenetic analyses indicated a close relationship between the domesticated silkworm and the Chinese wild silkworm. Domestication was estimated to have occurred about 4100 years ago (ya), and the radiation of the different geographic strains of B. mori about 2000 ya. The Chinese wild silkworm and the Japanese wild silkworm split about 23600 ya. These estimates are in good agreement with the fossil evidence and historical records. In addition, we show that the domesticated silkworm experienced a population expansion around 1000 ya. The divergence times and the population dynamics of silkworms presented in this study will be useful for studies of lepidopteran phylogenetics, in the genetic analysis of domestic animals, and for understanding the spread of human civilizations.     

Pan-African Genetic Structure in the African Buffalo (Syncerus caffer): Investigating Intraspecific Divergence

The African buffalo (Syncerus caffer) exhibits extreme morphological variability, which has led to controversies about the validity and taxonomic status of the various recognized subspecies. The present study aims to clarify these by inferring the pan-African spatial distribution of genetic diversity, using a comprehensive set of mitochondrial D-loop sequences from across the entire range of the species. All analyses converged on the existence of two distinct lineages, corresponding to a group encompassing West and Central African populations and a group encompassing East and Southern African populations. The former is currently assigned to two to three subspecies (S. c. nanus, S. c. brachyceros, S. c. aequinoctialis) and the latter to a separate subspecies (S. c. caffer). Forty-two per cent of the total amount of genetic diversity is explained by the between-lineage component, with one to seventeen female migrants per generation inferred as consistent with the isolation-with-migration model. The two lineages diverged between 145 000 to 449 000 years ago, with strong indications for a population expansion in both lineages, as revealed by coalescent-based analyses, summary statistics and a star-like topology of the haplotype network for the S. c. caffer lineage. A Bayesian analysis identified the most probable historical migration routes, with the Cape buffalo undertaking successive colonization events from Eastern toward Southern Africa. Furthermore, our analyses indicate that, in the West-Central African lineage, the forest ecophenotype may be a derived form of the savanna ecophenotype and not vice versa, as has previously been proposed. The African buffalo most likely expanded and diverged in the late to middle Pleistocene from an ancestral population located around the current-day Central African Republic, adapting morphologically to colonize new habitats, hence developing the variety of ecophenotypes observed today.     

New molecular data favour an anthropogenic introduction of the wood mouse (Apodemus sylvaticus) in North Africa

According to fossil data, the wood mouse arrived in North Africa 7500 ya, while it was present in Europe since Early Pleistocene. Previous molecular studies suggested that its introduction in North Africa probably occurred via the Strait of Gibraltar more than 0.4 Mya ago. In this study, we widely sampled wood mice to get a better understanding of the geographic and demographic history of this species in North Africa and possibly to help resolving the discrepancy between genetic and palaeontological data. Specifically, we wanted to answer the following questions: (1) When and how did the wood mouse arrive in North Africa? and (2) What is its demographic and geographic history in North Africa since its colonization? We collected in the field 438 new individuals and used both mtDNA and six microsatellite markers to answer these questions. Our results confirm that North African wood mice have a south‐western European origin and colonized the Maghreb through the Strait of Gibraltar probably during the Mesolithic or slightly after. They first colonized the Tingitana Peninsula and then expanded throughout North Africa. Our genetic data suggest that the ancestral population size comprised numerous individuals reinforcing the idea that wood mice did not colonize Morocco accidentally through rafting of a few individuals, but via recurrent/multiple anthropogenic translocations. No spatial structuring of the genetic variability was recorded in North Africa, from Morocco to Tunisia.     

Microsatellite variation and differentiation in African and non-African populations of Drosophila simulans

Drosophila simulans originated in sub-Saharan Africa or Madagascar and colonized the rest of the world after the last glaciation about 10 000 years ago. Consistent with this demographic history, sub-Saharan African populations have been shown to harbour higher levels of microsatellite and sequence variation than cosmopolitan populations. Nevertheless, only limited information is available on the population structure of D. simulans. Here, we analysed X-linked and autosomal microsatellite loci in four sub-Saharan African, one North African, one Israeli, and two European D. simulans populations. Bayesian clustering algorithms combined the North African, Israeli, and European populations into a single cosmopolitan group. The four sub-Saharan populations were split into two separate groups. Pairwise F(ST) analysis, however, indicated significant population differentiation between all eight populations surveyed. A significant signal for population reduction in cosmopolitan populations was found only for X-linked loci.     

Population changes of Antarctic fur seals at Nyrøysa, Bouvetøya

Antarctic fur seal Arctocephalus gazella pup production at Nyrøysa, Bouvetøya was estimated to be approximately 15,000 per annum during each of four summers from 1996 to 2002, indicating a total population of about 66,000. While the Bouvetøya population is the second largest for this species, pup production at this site still accounts for only 2.4% of the global total. This population experienced a mean annual rate of increase of 30.6% for the period 1989–1996, perhaps due, in part, to significant immigration, but has been stable since 1996. Historical accounts of significant numbers of animals being present towards the end of the period of sealing (C.1800–1930), indicate that the geographic isolation and inaccessibility of this site may have resulted in the Bouvetøya population being one of three populations that survived a series of periods of extreme exploitation of this species.     

Size and composition of the wild reindeer Rangifer tarandus platyrhynchus population in the Southeast Svalbard Nature Reserve

Alendal, E., de Bie, S. and van Wieren, S. E. 1979. Size and composition of the wild reindeer Rangifer tarandus platyrhynchus population in the Southeast Svalbard Nature Reserve. Holarct. Ecol. 2: 101-107. In the summer of 1977 we studied the reindeer population on the islands Barentsøya and Edgeøya in the eastern part of the Svalbard archipelago. A total of 1374 reindeer were observed: 326 animals in the western parts of Barentsøya and 1048 animals on Edgeøya. Considering those parts of Edgeøya which were not visited, the total number of reindeer on Edgeøya was estimated at 1300 animals. The total number of reindeer was lower than in previous years. The decline probably was due to severe winter conditions in 1975/1976 and 1976/1977 confirmed by the fact that many carcasses and few yearlings were observed. Nearly all reindeer occurred on the coastal plains and in the valleys. These areas have the relatively richest vegetation. The average recruitment of the total population (counted) was 15.9%. The adult sex ratio was in favour of females: 59% females versus 41% males. There were differences both in the recruitment and in the adult sex ratio between three distinct areas on Edgeøya and between two on Barentsøya. These differences may be due to dissimilarities in food quality and feeding conditions caused by climate, and by small exchange of reindeer between the areas. The high frequency of shed male antlers on Frankenhalvøya and Talaveraflya, north and south coast of Barentsøya respectively, indicates that these areas belong to the wintering grounds of reindeer on this island. Concentrations of shed female antlers on Barentsøya were less pronounced. The highest frequency was in the areas Sjodalen and Kvistdalen-Talaveraflya in the northwest and south respectively. Females may use these areas as late wintering grounds and possibly as calving areas. The average group size was 2.2 and the aggregation index 3.1. Seventytwo per cent of all groups, containing 48% of all reindeer, fell into group size 1 and 2. Males mostly were observed alone or together with one other animal. Females with calves most frequently occurred in groups of 2 and 4 animals.     

Non-indigenous introgression into the Norwegian red deer population

Rates of introgression from non-indigenous into native populations are increasing worldwide, often as a result of anthropogenic translocation events. In ungulates translocations have been common, especially among deer. European red deer consists of two distinct lineages, one western and one eastern. These probably originate from different glacial refuges, but it is unknown to what extent they hold different adaptations. Here we address dispersal and introgression into the Norwegian mainland population from an introduced island stock consisting of an admixture of both European lineages. The last decade this stock has grown considerably in number and dispersal could be expected to have increased. We therefore used samples separated by a 5 year interval from Otterøya, adjacent mainland areas and a more distant sub-population. Bayesian assignment analysis verified the genetic structure and identified dispersal between the Otterøya stock and the adjacent mainland coastal areas. Three individuals (two newly sampled) with second or third generation non-indigenous origin were found among the adjacent mainland samples (5 and 3 %, respectively). Two individuals with first and second generation mainland-origin were found on Otterøya (old samples). This suggests some non-indigenous introgression from Otterøya into the mainland Norwegian population.     

Population structure in African Drosophila melanogaster revealed by microsatellite analysis

Tropical sub-Saharan regions are considered to be the geographical origin of Drosophila melanogaster. Starting from there, the species colonized the rest of the world after the last glaciation about 10 000 years ago. Consistent with this demographic scenario, African populations have been shown to harbour higher levels of microsatellite and sequence variation than cosmopolitan populations. Nevertheless, limited information is available on the genetic structure of African populations. We used X chromosomal microsatellite variation to study the population structure of D. melanogaster populations using 13 sampling sites in North, West and East Africa. These populations were compared to six European and one North American population. Significant population structure was found among African D. melanogaster populations. Using a Bayesian method for inferring population structure we detected two distinct groups of populations among African D. melanogaster. Interestingly, the comparison to cosmopolitan D. melanogaster populations indicated that one of the divergent African groups is closely related to cosmopolitan flies. Low, but significant levels of differentiation were observed for sub-Saharan D. melanogaster populations from West and East Africa.     

Evolutionary history and biogeography of the drongos (Dicruridae), a tropical Old World clade of corvoid passerines

We address the phylogenetic relationships of the drongos (Dicruridae) at the species-level using sequences from two nuclear (myoglobin intron-2 and c-mos) and two mitochondrial (ND2 and cytochrome b) loci. The resulting phylogenetic tree shows that the most basal species is D. aeneus, followed in the tree by a trichotomy including (1) the Asian D. remifer, (2) a clade of all African and Indian Ocean islands species as well as two Asian species (D. macrocercus and D. leucophaeus) and (3) a clade that includes all other Asian species as well as two Australasian species (D. megarhynchus and D. bracteatus). Our phylogenetic hypotheses are compared to [Mayr, E., Vaurie, C., 1948. Evolution of the family Dicruridae (Birds). Evolution 2, 238-265.] hypothetical family "tree" based on traditional phenotypic analysis and biogeography. We point out a general discrepancy between the so-called "primitive" or "unspecialized" species and their position in the phylogenetic tree, although our results for other species are congruent with previous hypotheses. We conduct dating analyses using a relaxed-clock method, and propose a chronology of clades formation. A particular attention is given to the drongo radiation in Indian Ocean islands and to the extinction-invasion processes involved. The first large diversification of the family took place both in Asia and Africa at 11.9 and 13.3Myr, respectively, followed by a dispersal event from Africa to Asia at ca 10.6Myr; dispersal over Wallace line occurred later at ca 6Myr. At 5Myr, Principe and Indian Ocean Islands have been colonized from an African ancestor; the most recent colonization event concerned Anjouan by an immigrating population from Madagascar.     

Consequences for genetic diversity and population performance of introducing continental red deer into the northern distribution range

For several centuries, game management has involved translocations of non-native individuals of many species to reinforce local native populations. However, there are few quantitative studies of potentially negative effects on population viability as expected when taxa with different local adaptations hybridise. The European red deer has been subject to particularly many translocations. Around 1900, a total of 17 red deer of Hungarian (Cervus elaphus hippelaphus) and German (C. e. germanicus) origin were introduced onto the island of Otterøya in Norway where few native red deer (C. e. atlanticus) remained (n ~ 13). To assess interbreeding, the present stock on Otterøya and the indigenous Norwegian and Hungarian populations were characterised in 14 microsatellite loci and in the control region of mtDNA. An intermediate level of genetic variation in the Otterøya population and the presence of population specific alleles from both the indigenous Norwegian and the Hungarian population demonstrate that the introduced red deer interbred with the native. Even distributions of one indigenous and one non-indigenous mtDNA haplotype in the Otterøya population and two point estimates of admixture indicate similar genetic contributions from the two parental populations into the hybrid stock. Low numbers of migrants identified with Bayesian assignment tests demonstrate low recent gene flow from Otterøya into the Norwegian mainland population. The Otterøya hybrid stock has grown vastly in numbers during recent decades, suggesting a high population viability. We observed that the body mass of red deer on Otterøya was similar or greater than in adjacent indigenous Norwegian stocks, indicating that population performance has not been reduced in the hybrid stock and that gene flow probably has not had any negative effects.     

North African influences and potential bias in case-control association studies in the Spanish population

Background

Despite the limited genetic heterogeneity of Spanish populations, substantial evidences support that historical African influences have not affected them uniformly. Accounting for such population differences might be essential to reduce spurious results in association studies of genetic factors with disease. Using ancestry informative markers (AIMs), we aimed to measure the African influences in Spanish populations and to explore whether these might introduce statistical bias in population-based association studies.

Methodology/Principal Findings

We genotyped 93 AIMs in Spanish (from the Canary Islands and the Iberian Peninsula) and Northwest Africans, and conducted population and individual-based clustering analyses along with reference data from the HapMap, HGDP-CEPH, and other sources. We found significant differences for the Northwest African influence among Spanish populations from as low as ≈5% in Spanish from the Iberian Peninsula to as much as ≈17% in Canary Islanders, whereas the sub-Saharan African influence was negligible. Strikingly, the Northwest African ancestry showed a wide inter-individual variation in Canary Islanders ranging from 0% to 96%, reflecting the violent way the Islands were conquered and colonized by the Spanish in the XV century. As a consequence, a comparison of allele frequencies between Spanish samples from the Iberian Peninsula and the Canary Islands evidenced an excess of markers with significant differences. However, the inflation of p-values for the differences was adequately controlled by correcting for genetic ancestry estimates derived from a reduced number of AIMs.

Conclusions/Significance

Although the African influences estimated might be biased due to marker ascertainment, these results confirm that Northwest African genetic footprints are recognizable nowadays in the Spanish populations, particularly in Canary Islanders, and that the uneven African influences existing in these populations might increase the risk for false positives in association studies. Adjusting for population stratification assessed with a few dozen AIMs would be sufficient to control this effect.     

The nature and causes of annual fluctuations in numbers of Aphis fabae Scop. on field beans (Vicia faba)

Experiments for nine successive years showed that Aphis fabae Scop. populations on mid-March-sown field beans were either large with peak densities between late June and mid-July or very small with peak densities in early August. It is concluded that the largest populations develop when many plants have been colonized by primary migrants from Euonymus europaeus and temperature and radiation are above average during June and early July, as in the year 1957. Cold, dull weather slows multiplication and decreases the size of the peak population even when there is a large initial colonization, as in 1954. The peak population may also be less than predicted from the initial colonization when natural enemies are exceptionally abundant in early June, as in the year 1960. Yield losses of mid-March-sown crops in years of large A. fabae populations ranged from 53 % in 1954 (peak population of 1260 aphids per plant) to 100% in 1957 (6920 aphids per plant). Small summer populations with peak densities of about 0·2–85 aphids per plant developed on mid-March-sown plots in years when fewer than about 6% of the plants were colonized by primary migrants. Yield losses ranged from 6·3–13·6%. Three years' experiments indicated that crops sown in late April or May are relatively lightly infested in years when large populations develop on mid-March-sown crops. Conversely, they may be relatively heavily infested when the populations on these crops are small, as in 1955 when temperatures and sunshine during July and early August were above average. Small and large early summer populations tend to alternate in successive years. The alternation is upset by hot, sunny weather during July and August, and perhaps September and October, which compresses the population cycle Thus the large and small populations expected from this alternation in 1956 and 1960 developed instead during exceptionally fine weather in late summer 1955 and 1959, converting 1956 and 1960 to years of small and large populations respectively.     

Rapid evolution of goat and sheep globin genes following gene duplication

Statistical analyses of DNA sequences of globin genes (beta A, beta C, and gamma) from goat and sheep (including new sequence information for the second intron of sheep beta A and gamma, kindly provided by A. Davis and A. W. Nienhuis) indicate that the rates of nonsynonymous substitution in these genes have been greatly accelerated following the gene duplication separating gamma and the ancestor of beta A and beta C and the gene duplication separating beta A and beta C. In both cases the acceleration was apparently due to relaxation of purifying selection (functional constraints) rather than advantageous mutations because acceleration occurred only in less important parts of the beta globin chain. The rates of nonsynonymous substitution in these genes are estimated to be about 2.3 x 10(-9) per site per year, which is three times higher than that for the divergence between human beta and mouse beta major globin genes. Our analyses further suggest that the rate of synonymous substitution in functional genes and the rate of substitution in pseudogenes are approximately equal and are between 2.8 x 10(-9) and 5.0 x 10(-9) and that the rate of substitution in introns is about 3.0 x 10(-9). The divergence time between beta A and beta C and that between gamma and the beta A-beta C pair are about 12 and 30 million years, respectively. The proportion of transition mutations is estimated to be 64%, two times higher than expected under random mutation but considerably lower than the 96% estimated for animal mitochondrial DNA.      

GENETIC VARIABILITY AND POPULATION DIFFERENTIATION INFERRED FROM DNA FINGERPRINTING IN SILVEREYES (AVES: ZOSTEROPIDAE)

This study evaluated DNA fingerprinting as a tool for estimating population genetic diversity and differentiation by comparing minisatellite variation in island and mainland populations of silvereyes (Aves: Zosterops lateralis). Three populations with different recent histories were compared: (1) Heron Island and neighboring islands, colonized 3000 to 4000 yr ago; (2) Lady Elliot Island, colonized within the past two decades; and (3) an adjacent mainland population, which presumably has existed for thousands of years. The degree of genetic variability within the three populations reflected both their size and the time since their colonization. Minisatellite diversity was highest in the mainland population, intermediate in the Capricorn Island group (which was shown to represent a single admixture), and lowest in the Lady Elliot Island population, possibly because of a recent population bottleneck during colonization. Mean band sharing between any two populations was less than the mean within either of those populations, and four fingerprint bands common to island birds were rare or absent in the fingerprints of mainland birds. In the absence of significant gene flow between the mainland and the islands, the populations have apparently become distinct at minisatellite loci, as evidenced by differences in both allelic diversity and in the frequencies of specific fragments. Within the Heron Island population, cohort analyses demonstrated the temporal stability of the fingerprint profile over 6 yr. This study demonstrates that length polymorphisms at minisatellite loci may be stable enough over time to retain information about recent historical and demographic effects on the relative genetic variability and differentiation of small, closely related populations.     

Out of Africa and the evolution of human behavior

Twenty‐one years ago, a landmark exploration of mitochondrial DNA diversity popularized the idea of a recent African origin for all living humans. 1 The ancestral African population was estimated to have existed 200 ka (thousands of years ago) plus or minus a few tens of thousands of years. A corollary was that at some later date the fully modern African descendants of that population expanded to swamp or replace the Neanderthals and other nonmodern Eurasians. The basic concept soon became known as “Out of Africa,” after the Academy Award winning film (1985) that took its title, in turn, from Isak Dinesen's classic autobiography (1937). Many subsequent genetic analyses, including those of Ingman and coworkers 2 and Underhill and coworkers, 3 have reaffirmed the fundamental Out of Africa model. The fossil and archeological records also support it strongly. The fossil record implies that anatomically modern or near‐modern humans were present in Africa by 150 ka; the fossil and archeological records together indicate that modern Africans expanded to Eurasia beginning about 50 ka.     

Nonneutral admixture of immigrant genotypes in African Drosophila melanogaster populations from Zimbabwe

Drosophila melanogaster originated in Africa and colonized the rest of the world only recently (approximately 10,000 to 15,000 years ago). Using 151 microsatellite loci, we investigated patterns of gene flow between African D. melanogaster populations representing presumptive ancestral variation and recently colonized European populations. Although we detected almost no evidence for alleles of non-African ancestry in a rural D. melanogaster population from Zimbabwe, an urban population from Zimbabwe showed evidence for admixture. Interestingly, the degree of admixture differed among chromosomes. X chromosomes of both rural and urban populations showed almost no non-African ancestry, but the third chromosome in the urban population showed up to 70% of non-African alleles. When chromosomes were broken into contingent microsatellite blocks, even higher estimates of admixture and significant heterogeneity in admixture was observed among these blocks. The discrepancy between the X chromosome and the third chromosome is not consistent with a neutral admixture hypothesis. The higher number of European alleles on the third chromosome could be due to stronger selection against foreign alleles on the X chromosome or to more introgression of (beneficial) alleles on the third chromosome.     

Evolution of glucagon genes

Statistical analyses of DNA sequences of the preproglucagon genes from bovine, human, hamster, and anglerfish suggest that a gene duplication creating two anglerfish genes (AF I and II) occurred about 160 Myr ago, long after the separation of fish and mammals. The analyses further suggest that the internal duplication producing the glucagon and glucagon-like peptide II (GLP-II) regions occurred about 1.2 billion years ago, which would indicate that the GLP-II region was present in the ancestral anglerfish sequence but was silenced or deleted before the gene duplication separating AF I and II. The glucagon-like peptide I (GLP-I) was derived from a duplication of the ancestral glucagon region about 800 Myr ago. The rate of synonymous substitution in these genes is approximately 4.3 x 10(-9) substitutions per year per synonymous site. The rate of nonsynonymous substitution in the signal peptide region is about 1.1 x 10(-9) substitutions per year per nonsynonymous site, a high rate comparable to that in the C-peptide region of preproinsulin. The rate of nonsynonymous substitution in the glicentin-related pancreatic polypeptide (GRPP) region is 0.63 x 10(-9) for the comparisons between mammalian species and 1.8 x 10(-9) for the comparisons between fish and mammals; the moderate rate in mammals suggests a physiological role for GRPP. The glucagon region is extremely conservative; no nonsynonymous substitution is observed in the mammalian genes, and a nonsynonymous rate of 0.18 x 10(-9) was obtained from the comparisons between fish and mammals. In the GLP-I region, the rate of nonsynonymous substitution was estimated to be 0.08 x 10(-9) for the comparisons between mammalian species and 0.30 x 10(- 9) for the comparisons between fish and mammals. In the GLP-II region, the rate was estimated to be 0.25 x 10(-9) for the comparisons between mammalian species. Thus, GLP-I and II are also very conservative, which suggests an important physiological role for these peptides.