Gene flow from archaic humans to Homo sapiens

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In this family tree hypothesis (for the time up to 2 million years ago), its author, Chris Stringer , emphasizes in 2012 the central position of Homo heidelbergensis as a link between Neanderthals , Denisovans and Homo sapiens ; other paleoanthropologists assign the African finds identified here as heidelbergensis to Homo erectus . On the far right, Stringer indicates that some genetic abnormalities have been detected in Africa which - besides Neanderthals and Denisovans - seem to point to a third gene flow from a previously unexplained pre -human population to anatomically modern humans. In the Asian Homo erectus , Stringer emphasizes the separation into Peking man and Java man , and he interprets Homo antecessor as an early European branch of Homo erectus . The origin of Homo floresiensis is unclear.

Gene flow from archaic human to anatomically modern humans ( Homo sapiens ) came to current state of research at least twice in the history of the genus Homo : on the one of Neanderthals , on the other hand from the as Denisova people designated population . A small proportion, an estimated 1% to 4%, of the DNA of Eurasians and North Africans is not “ modern ” and agrees with that of the Neanderthals, while these genetic markers are found in Sub-Saharan Africans (i.e. in the members of the Yoruba and the San ) were undetectable.

In Melanesians an additional portion comes from estimated 4% to 6% of their DNA with the interpretations of the researchers, according to relatives of tight Asian Neanderthals, the Denisova people . A DNA analysis carried out in 2011 also points to a third gene flow in Africans from a previously unknown, extinct archaic population.

Neanderthals

In the course of the 20th century, different hypotheses about a possible pairing of Neanderthals and anatomically modern humans in the Middle Paleolithic were discussed on the basis of anatomical features ; in the 2000s also with reference to genetics .

  • In the meantime, genetic analysis studies published between 2013 and 2015 on the Homo sapiens finds from Peştera cu oasis in Romania and Ust-Ischim in Siberia have provided evidence of Neanderthal DNA in both fossils. Accordingly, a mating between Neanderthals and Homo sapiens can be assumed not only in the Levant, but also in Eastern Europe and in Siberia.

The genetic analyzes indicate that gene flow between Neanderthals and Homo sapiens occurred several times and independently of one another . It is believed that a gene flow between the two species occurred more than 270,000 years ago; it has been speculated that this event may have occurred in the Middle East with the participation of a homo sapiens population from Africa .

Research history

The mixing hypothesis (also: hybridization theory) has been discussed in specialist circles since the discovery of the first Neanderthal remains in the 19th century. At first it was believed that the Neanderthals were the direct ancestors of modern man; he was considered a predecessor from which the modern present-day man developed. Thomas Huxley suspected that many Europeans still had traces of their Neanderthal ancestors, namely negative traits such as primitivism . Furthermore, the further development from Neanderthals to present-day people took place in parallel for all ethnicities - then declared as races : a hypothesis that in the 20th century was referred to as the "multiregional model" .

In 1907, the Danish ethnographer Hans Peder Steensby, in his article Race Studies in Denmark, rejected the interpretation that the Neanderthals were ape-like or inferior. Rather, all people today are of mixed origin. A cross between humans and Neanderthals is the best available explanation for the majority of the findings known at the time.

In the first half of the twentieth century Carleton S. Coon took the view that the so-called Caucasian race arose from a double mixture. Initially, in the Paleolithic Age, Neanderthals and Homo sapiens were mixed . These hybrids were later mixed again with Homo sapiens from the Mediterranean region. He repeated his theory in his 1962 book The Origin of Races .

The British psychologist Stan Gooch also developed a theory about the alleged hybridization of Neanderthals and Cro-Magnons in his works Personality and Evolution (1973) and The Neanderthal Question (1977) . This theory, however, was not based on a comparison of the anatomical features of both populations, but relied largely on Gooch's interpretation of modern human psychology and society, which he believed were at least partially traceable to Neanderthal culture. Although his theories were rejected by scientific paleoanthropology , he developed his approach further in Cities of Dreams (1989) and The Neanderthal Legacy (2008).

The focus of the debate shifted from the study of anatomy to the field of archaeogenetics in the 2000s , after advances in this area from 2006 onwards raised hopes for a deciphering of the Neanderthal genome.

anatomy

One of the loudest proponents of the anatomical hybridization thesis was Erik Trinkaus of the University of Washington. Trinkaus interpreted numerous fossils as hybrids , including the child of Lagar Velho , a skeleton found in Lagar Velho (Portugal) and dated around 24,000 years ago. In another publication from 2006, co-authored by Trinkaus, the fossils found in 1952 in the Peştera Muierii cave (Romania) were also interpreted as hybrids.

In his work "Neanderthal", Paul Jordan points out that certain functions on some "modern" skulls of Eastern European Cro-Magnons are difficult to explain without mixing them. In another study recently in Peştera Muierilor, Romania , the researchers found remains of people dated to an age of 30,000 years. These had anatomical features that are mostly diagnosed as "modern". At the same time, however, according to the researchers' interpretations, they have characteristics that are typical of Neanderthals, cannot be explained as a parallel evolution and were not developed in early Homo sapiens in Africa. This included: a large bulge on the back of the skull, a stronger attachment at the elbow joint and a narrow base at the shoulder joint. With the analysis of the shoulder of an individual it was shown that this, like a Neanderthal, probably did not have the full capacity to throw a javelin.

On the theses based on the anatomy of Iberian Neanderthal fossils

The paleontological analysis of the development of modern man in Europe has shifted from looking at the Neanderthals to the biology and chronology of the earliest modern man in western Eurasia. This focus, which includes morphologically modern humans 28,000 years ago, shows increasing evidence that they represent a variable mosaic of modern humans, archaic humans, and Neanderthal traits. Studies of fossils from the upper plains of the Sima de las Palomas, Murcia (Spain), dated 40,000 years ago, show a late persistence of the Neanderthals on the Iberian Peninsula. This supports the conclusion that the Neanderthals were not only displaced or even swept away by the advance of modern humans.

In addition, the Neanderthals 'Palomas' have a number of features of modern humans that were rarely or not at all trained in earlier Neanderthals. Either they developed independently to the modern-human-looking features or, what is more likely, they had contact with early modern humans around the Pyrenees. If the latter is the case, it implies that their persistence in the Middle Paleolithic in the Iberian Peninsula was a matter of choice, rather than cultural retardation.

genetics

The DNA sequence of the Neanderthals published by researchers from Svante Pääbo's group in May 2010 shows that non-African populations of anatomically modern humans share some genetic characteristics with Neanderthals. According to the authors of this study, this is best explained by a gene flow from Neanderthals to anatomically modern humans, which is believed to have occurred in the Middle East . An estimated 1 to 4% of the DNA of Europeans and Asians (subjects of French, Chinese and Papuan descent) matches the DNA of Neanderthals. These genes could not be detected in test persons from the regions south of the Sahara (in test persons from the Yoruba and the San). Some researchers even suspect a gene flow of 3.4–7.9% in Eurasian populations.

In more recent studies, a higher proportion of gene flow from Neanderthals to Homo sapiens was found in East Asians compared to Europeans. From this it was concluded on the one hand that there were at least two different gene flow events and on the other hand that the second gene flow occurred in East Asians after the two groups had separated. Further explanations for the difference are discussed: Due to a possible later emigration of the ancestors of the Europeans, a dilution of this proportion could have occurred and in the case of East Asians there was a lower selection caused by evolutionary pressure. It was also found that there is a small but significant range of variation in the Neanderthal proportion within European populations, which is not found in East Asian populations.

  • In May 2015, at a symposium in Cold Spring Harbor , New York , the preliminary results of the sequencing of the DNA of the fossil Oase 1 , dated to an age of 40,000 years and from the oasis cave in Romania, were presented. The genetic analysis of Oasis 1 showed that this fossil has between six and nine percent of the genes of the Neanderthal. The research team led by Qiaomei Fu estimated that the connection between Homo sapiens and a Neanderthal man was no more than four to six generations ago in this fossil . This study shows that Homo sapiens and Neanderthals were successfully mated not only in the Middle East, but also in Europe.
  • When Ust-Ishim was found in Siberia , 2 percent of Neanderthal DNA was found. The time of gene flow was dated to around 7,000 to 13,000 years before the individual's lifetime (around 45,000 years ago) - with genetic proximity to the people living in Eurasia.

No evidence of Neanderthal genes had previously been found in the mitochondrial DNA of modern Europeans, suggesting that no direct maternal line of Neanderthal origin would have survived into modern times.

Changes in the human microcephalin gene, which has an important influence on brain size, was also cited as possible evidence of gene flow. The loss of function of this gene due to harmful mutations can lead to microcephaly . One type of this gene, called the (modern) "D", has a worldwide distribution of 70%. However, it has a young coalescence age. The youngest common ancestor of the carriers of this gene lived about 37,000 years ago. The remaining species of this gene, referred to as "Non-D" or "Non-D", coalesced to about 990,000 years ago. The separation of the D and non-D genes is estimated to have occurred 1,100,000 years ago. It is possible that the D gene conferred an evolutionary advantage on its carriers; but this is controversial.

Since the distribution of the D allele is large outside of Africa, but lower in the region south of the Sahara at 29%, it was suggested to infer the involvement of an archaic Eurasian population group. Estimates of the divergence time of modern humans and Neanderthals, based on mitochondrial DNA, suggest that this archaic population is the Neanderthals, from whom this gene entered the gene pool of modern humans through introgression . However, an examination of the DNA from the Neanderthal fossils from the Vindija cave (Croatia) by Svante Pääbo showed that it contained the non-D allele for microcephalin and that there was no evidence of gene flow or crossbreeding. However, a study published in May 2010 showed that one of the Neanderthals from the Mezzana site ( Monti Lessini , Italy) carried the older version of the D allele, which is common in Eurasian modern humans. This study did not rule out crossbreeds between Neanderthals and Homo sapiens , but stated that the individual DNA findings could not support the thesis that the D allele was passed on from Neanderthals to modern humans.

Based on a 2001 study of the gene that causes redhead in humans, it has been speculated that the Neanderthals may have had red hair and may have passed this trait on to redheaded and freckled modern humans. A 2007 study to analyze Neanderthal DNA found that some Neanderthals were actually red-haired, but that this was due to a mutation in the MC1R gene that could not be detected in modern humans.

Genes that affect keratins , however, have been found to have flowed from Neanderthals to modern East Asians and Europeans, and are believed to have led to better morphological adaptation of skin and hair in environmental conditions outside of Africa. This is also assumed for various genes that are associated with diseases such as type 2 diabetes , primary biliary cirrhosis , the autoimmune diseases lupus erythematosus and Crohn's disease or smoking behavior.

It has also been argued that some genes affecting the immune system may have come from Neanderthals. The "HLA-C * 0702" gene was found in Neanderthal fossils and is common in modern Europeans and Asians, but is extremely rare in people of African descent. It was therefore assumed that this immune gene entered the gene pool of Homo sapiens after leaving Africa and enabled him to survive against diseases outside of Africa, against which the Neanderthals were already armed.

A study published in early 2017 examined the extent to which genes assigned to the Neanderthals are active in gene expression in today's modern humans . One finding was that cis-acting effects could be determined for a quarter of the haplotypes introgressed by the Neanderthals . Gene regulatory variants introgressed by the Neanderthals have a phenotypic effect . However, Neanderthal alleles are expressed differently depending on the type of tissue. These are often downregulated when genes are expressed in the brain and testes. These are suppressed very effectively, especially in the cerebellum and the basal ganglia .

In October 2017, the almost complete genome of a Neanderthal woman who lived in the Vindija cave around 52,000 years ago was reconstructed . A comparison with the only similar completely reconstructed Neanderthal genome from the Denissowa Cave in the Altai Mountains revealed differences of only 1.6 deviations per 1000 base pairs , which means that these two individuals were genetically closer to each other than any two, is the case with people living today. From this it was concluded that the Neanderthals lived in a very small population. At the same time, the proportion of Neanderthal DNA in the genetic make-up of people living outside Africa was now shown to be 1.8 to 2.6 percent, and the continued activity of some genes that had passed over to anatomically modern humans was described.

Some of the genes introduced by the Neanderthals into the non - African Homo sapiens populations have also been detected in today's African Homo sapiens populations; from this finding it was concluded that these genes come from the gene pool of the common ancestors of Neanderthals and Homo sapiens . The emigration of relatively small Homo sapiens groups from Africa led to a reduction in their genetic diversity ( genetic bottleneck ), and the introgression of Neanderthal DNA later brought some of the previously lost genes back into the homo sapiens populations outside of Africa .

Denisova people

Tests comparing the genome of Denisovans with those of six people living today - a "! Kung" from South Africa, a Nigerian, a French, a Papuan, a bougainviller and a Han Chinese - have shown that between 4% and 6% of the genome of the Melanesians (represented by the Papuan and the Bougainville Islanders) match that of the Denisova people. This DNA may have been introduced into the local population of anatomically modern humans during the early migration to Melanesia . This interpretation fits in with the results of the other comparative tests, which showed an increase in the alleles shared by Denisovans and Australian Aborigines; other Eurasian and African populations served as a basis for comparison. It has also been observed that the Papuan genome in Papua New Guinea has more allele matches with the Denisova people than that of the Aborigines in Australia. Adding Neanderthal DNA and Denisova DNA shows that the Melanesians have the largest share of the genome of archaic people worldwide, at around 8%.

The gene flow to the ancestors of the Melanesians and other populations in Oceania presumably occurred independently of that to the ancestors of the people of East Asia .

David Reich (2010) from Harvard University, in collaboration with Mark Stoneking (1997) from the team at the Max Planck Institute for Evolutionary Anthropology, also found genetic evidence that the Aborigines and smaller scattered groups of people in Southeast Asia, so-called Negrito peoples like the Mamanwa in the Philippines, also have the DNA of the Denisova people. However, not all Negrito peoples have DNA from the Denisova people; the Onge , the indigenous people of the island of Little Andaman, and the Malaysian Jehai , for example, were found to have no traceable Denisovan genetic makeup. These data have been interpreted to mean that the gene flow may have occurred in mainland Southeast Asia and suggest that the Denisova people were once widespread in East Asia.

The immune system alleles of the HLA genes have attracted particular attention in an attempt to identify genes that may have come from archaic humans. Although it is not included in the Denisova genome, the distribution pattern and divergence of the "HLA-B * 73" allele from other HLA alleles has led to the assumption that it is through introgression from Denisova humans into populations Western Asia was brought in. In fact, half of the HLA alleles in modern Eurasier are archaic HLA haplotypes; from which it was concluded that they could have come from Denisovans or from Neanderthals. The frequency of these alleles in Homo sapiens suggests a positive selection pressure for their distribution and maintenance.

It is possible that in Asia, in addition to the introgression on the part of the Denisovans, there was also gene flow from two other, so far not identified populations to anatomically modern humans.

Unknown sub-Saharan African man

In 2011 Michael Hammer et al. from the University of Arizona DNA of two African hunter-gatherer groups, the Biaka (also Aka ), a pygmy people , and the San , as well as the West African, agricultural Mandinka . They concluded that around two percent of the genetic material in these modern African populations was introduced into the genome of the hunter-gatherer groups about 35,000 years ago. They also came to the conclusion that these sequences must have come from a taxon of the genus Homo , which is now extinct and has not yet been fossilized , which split off from the lineage leading to anatomically modern humans around 700,000 years ago.

In 2012, Sarah Tishkoff et al. another study published by the University of Pennsylvania. They tested three sub-Saharan African populations - Pygmies, from Cameroon, and the Hadza and Sandawe , both from Tanzania. The team found evidence that ancestors of the hunter-gatherers had mixed up with members of previously unknown hominini populations, likely more than 40,000 years ago. The Iwo-Eleru skull is also interpreted as possible evidence of the survival of a population with partly archaic features.

A study published in 2017 also supported the hypothesis of an introgression of genes in an archaic hominini population that had not yet been fossilized. Ultimately, this could mean that Homo sapiens in Africa did not emerge from a single, regional stem population, but from loosely connected groups that lived all over Africa and - after being isolated from one another for longer periods - repeatedly exchanged genes.

In 2019 and 2020, the hypothesis was confirmed again in two studies. From the results of the studies published in 2019, it was concluded “that the estimated proportion of the Neanderthal genome in Eurasians is strongly influenced by the presence of the ghost population” and the amount of DNA that comes from Neanderthals in Eurasians could therefore be up to three times higher than previous models suggested. In 2020, it was also not ruled out that several populations contributed to the introgression.

Super archaic influences

In February 2020, a study was published, according to which gene flow from a previously unidentified, archaic homo- population to the common ancestors of Neanderthals and Denisovans, whose genetic markers increase today - as a result of later gene flow, was published Homo sapiens - are also detectable in anatomically modern humans.

literature

  • Rajiv C. McCoy, Jon Wakefield, and Joshua M. Akey: Impacts of Neanderthal-Introgressed Sequences on the Landscape of Human Gene Expression. In: Cell . Volume 168, No. 5, 2017, pp. 916–927.e12, doi: 10.1016 / j.cell.2017.01.038

See also

Individual evidence

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