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Hominin Interbreeding and the Evolution of Human Variation Kwang Hyun Ko*

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Hominin Interbreeding and the Evolution of Human Variation Kwang Hyun Ko* Ko J of Biol Res-Thessaloniki (2016) 23:17 DOI 10.1186/s40709-016-0054-7 Journal of Biological Research-Thessaloniki REVIEW Open Access Hominin interbreeding and the evolution of human variation Kwang Hyun Ko* Abstract Mitochondrial Eve confirms the “out of Africa” theory, but the evidence also supports interbreeding between Homo sapiens and other hominins: Neanderthals, Denisovans, and Homo heidelbergensis. This article explains how inter- breeding between early H. sapiens and archaic hominins occurred. The availability of edible insects in East Asia aided the spread of the unaggressive, highly cooperative Neanderthals, who interbred with H. sapiens in Asia, resulting in a higher admixture of Neanderthal DNA in East Asian populations. Geographical variation in degree of interbreed- ing between H. sapiens and Neanderthals likely contributed to neurological and behavioral differences in modern humans. Similarly, people with Denisovan genetic admixture were better able to dwell in mountainous regions, allow- ing their genetic legacy to cross the Himalayas and persist in Southeast Asian and Oceanian H. sapiens. In the Sub- Saharan region, unaffected by Denisovan or Neanderthal interbreeding, H. sapiens interbred with H. heidelbergensis, because high humidity militated against fire-making and allowed the survival of these non-fire-making hominins. Keywords: Neanderthal, Homo sapiens, Denisovan, Homo heidelbergensis, Hominin interbreeding Background to make up a single, continuously gradient (as distinct There are two branching hypotheses on the origin of the from categorically separate) human species. human species. The most widely accepted is the “out of DNA analysis demonstrating the existence of “Mito- Africa” (OOA) theory, which holds that archaic Homo chondrial Eve” has strongly corroborated the recent Afri- sapiens evolved into anatomically modern humans solely can origin model of OOA by providing crucial support to in Africa between 200,000 and 60,000 years ago [1]. This the theory that H. sapiens moved from Africa to replace hypothesis further proposes that members of one branch residing hominin populations elsewhere [4]. Mitochon- of H. sapiens left Africa at some point between 125,000 drial Eve is the most recent matrilineal common ancestor and 60,000 years ago, and that over a long period, these of all humans currently alive. Women pass along mito- H. sapiens replaced more “primitive” populations of other chondrial DNA unchanged during sexual reproduction, hominins in Asia or Europe, such as Homo neandertha- and the DNA of this most recent woman from whom all lensis and Homo erectus [2]. currently living humans descend through an unbroken The competing theory is the multiregional evolu- line on their mother’s side proves that modern humans tion hypothesis [3], which argues that some or all of the only evolved once, most likely in East Africa, sometime genetic variation among contemporary human races is between 150,000 and 200,000 years ago. attributable to genetic inheritance from either other H. At the same time, however, studies suggest that Nean- sapiens subspecies or from other hominid species. In the derthals, our closest-known evolutionary relatives, coex- multiregional model, all archaic human forms worldwide, isted with H. sapiens on Earth for more than 5000 years such as H. erectus and Neanderthals, as well as modern and frequently interbred with modern humans [5]. forms, subsequently evolved together into the diverse According to researchers, at least one-fifth of the Nean- populations of modern H. sapiens, which are considered derthal genome may lurk within modern humans, influ- encing traits including the appearance of the skin and hair people have today and the diseases they get. This *Correspondence: [email protected]; [email protected] Hanyang University, Seoul, South Korea finding indicates that a true “extinction” of Neanderthals © 2016 The Author(s). This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The Creative Commons Public Domain Dedication waiver (http://creativecommons.org/ publicdomain/zero/1.0/) applies to the data made available in this article, unless otherwise stated. Ko J of Biol Res-Thessaloniki (2016) 23:17 Page 2 of 9 may not have occurred [6], but that they may have been of Denisovan ancestry in the islands of Oceania, particu- absorbed into H. sapiens. Genetic evidence shows that larly Papua New Guinea, and some parts of mainland other archaic hominins, such as the Denisovans, also Asia, such as Tibet. interbred with H. sapiens [7]. The most current version of the OOA hypothesis emphasizes the African origin Why does Neanderthal ancestry appear to a higher degree of most human populations but allows for the possibil- in Asia? ity of local contributions/interbreeding between humans All H. sapiens living today have interbred to some degree and other hominins [8]. Consequently, this article mainly with Neanderthals, Denisovans, or other hominins, and discusses Neanderthal–human interbreeding, while also as outlined above, we know that these hominin groups explaining other admixtures of archaic humans with lasted longer and interbred more in some parts of the hominins who were their contemporaries, such as Den- world than in other areas. isovans and H. heidelbergensis. Most hominins other than Denisovans and Neander- thals were simply replaced by H. sapiens that migrated Origin of race: human interbreeding with Neanderthals out of Africa, but sufficient interbreeding occurred with and Denisovans Denisovans and Neanderthals in Eurasia to leave a sig- Neanderthals are an extinct species of human (in the nificant mark on modern human DNA [15]. Because genus Homo), related to modern humans [9]. Traces left Neanderthals ranged only from Europe to West Asia, the by Neanderthals include bone and stone tools, which question of why there were two waves of interbreeding have been found all over Eurasia, from Western Europe between East Asian H. sapiens and Neanderthals remains to central and northern Asia. Neanderthals are generally a mystery. The answer to this question lies in differences classified by biologists as H. neanderthalensis, and some- in behavior, and in particular aggressiveness, between times as Homo sapiens neanderthalensis. groups of hominins. Denisovans are another extinct species of humans, Homo heidelbergensis, which exhibited proto-Nean- similar to Neanderthals. The Denisova Cave is located derthal traits, existed in Eurasia as early as 350,000– in southwestern Siberia, in the Altai Mountains near the 600,000 years ago, while the first Neanderthals appeared Russian border with China and Mongolia [10]. Research between 200,000 and 250,000 years ago [16]. At varying shows that Denisovans shared a common origin with times, Neanderthals inhabited the region from Western Neanderthals but were genetically distinct. Europe to Central Asia; their eastern and northern range Recent genetic studies have shown a higher Neander- extended to Okladnikov in the Altai region and Byzovaya thal admixture in East Asians compared with Europeans in the Ural region of present-day Russia [17]. Neander- [11], most likely indicating that at least two independent thals started to disappear/interbreed with H. sapiens gene-flow events must have taken place in early mod- from the time the latter migrated to Europe. Fossil find- ern humans and that the early ancestors of East Asians ings have indicated brutality and violence among H. sapi- experienced more admixture than those of Europeans ens living 10,000 years ago [18]. The evidence has shown after the divergence of these two groups [12]; to put it in that in addition to interbreeding, Neanderthals were another way, studies seeking to explain why East Asians also very often killed by H. sapiens, and in related find- inherited 15–30 % more Neanderthal DNA than Euro- ings, genetic studies have shown that the mutations in peans have concluded that East Asians interbred with ADSL, GLDC, and SLITRK1 genes, which are associated Neanderthals in two waves [13]. with hyperactivity and aggressive behavior in modern The first interbreeding with Neanderthals occurred humans, were not found in Neanderthals [19–21]. Thus, in the Middle East before the ancestors of modern non- by multiple methods, H. sapiens were responsible for the Africans spread out across Eurasia. The ancestors of extinction of Neanderthals, who were more cooperative modern Europeans and Asians then split out of this and less aggressive than H. sapiens according to studies migrant group [12], and the ancestors of East Asians from various fields. interbred again with Neanderthals after the split. The first It should be noted that compared to Africa, Eurasia humans with proto-Neanderthal traits are believed to lacks predators that could have presented a threat to have existed in Eurasia as early as 350,000–600,000 years hominin species. The increase in species richness or bio- ago, with the first “true Neanderthals” appearing between diversity that occurs from the poles to the tropics is often 200,000 and 250,000 years ago. referred to as the latitudinal diversity gradient (LDG), As this implies, Neanderthals and Denisovans were and the greatest biodiversity
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