结论

随着现代人类扩张并走出非洲,爆发了一场创新。在扩张之前,在非洲就有这类改变的迹象,然而在解剖学意义上的现代人类遭遇并取代尼安德特人之后,这一迹象大约两万年后在欧洲大大强化。从考古学记录上看,由创新过渡到全面的行为现代性不规则地发生在许多地方,之后最终在亚欧大陆的其他地区完成。我们提出的观点是,即便从尼安德特人(或其他远古人类)身上流向解剖学意义上的现代人类的基因流很有限,这也有助于现代人类获取大部分他们所需要的优势等位基因。我们相信这短时间内大量涌入的适应性基因组对“人类革命”所需的能力的贡献良多,我们也同样相信来自远古人类群体的基因渗入会被证实在现代人类起源的故事中有中心的地位。

所以截至大约4万年前,人类在身体结构上和行为上都变得现代(这不是说他们就完全像今天的人们一样了)。他们有着比他们的祖先远为强大的创新力量,很可能得益于从他们的尼安德特表亲身上偷得的基因。文化改变的速度增长了几十倍,而当冰川后退之后新的机会涌现,改变的速度就更快了。

[1]Roy C.Anderson,“The Ecological Relationships of Meningeal Worm and Native Cervids in North America,”Journal of Wildlife Diseases 8,no.4(1972):304—310.

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[2]Daniel M.Tompkins et al.,“Parapoxvirus Causes a Deleterious Disease in Red Squirrels Associated with UK Population Declines,”Proceedingsof the Royal Society,B:Biological Sciences 269,no.1490(2002):529.

[3]Richard Klein,The Human Career(Chicago:University of Chicago Press,1999),524.

[4]Richard Klein,The Dawn of Human Culture(New York:Wiley,2002),270.

[5]Christopher S.Henshilwood et al.,“Emergence of Modern Human Behavior:Middle Stone Age Engravings from South Africa,”Science 295,no.5558(2002):1278—1280.

[6]Carleton S.Coon,The Origin of Races(New York:Alfred A.Knopf,1963).

[7]Chris Stringer and Peter Andrews,“Genetic and Fossil Evidence for the Origin of Modern Humans,”Science 239,no.4845(1988):1263—1268.

[8]D.Curnoe et al.,“Timing and Tempo of Primate Speciation,”Journal of Evolutionary Biology 19,no.1(2006):59—65.

[9]Hilde Vervaecke and Linda Van Elsacker,“Hybrids between Common Chimpanzees(Pan troglodytes)and Pygmy Chimpanzees(Pan paniscus)in Captivity,”Mammalia(Paris)56,no.4(1992):667—669.

[10]Trenton W.Holliday,“Speciation by Distance and Temporal Overlap:A New Approach to Understanding Neanderthal Evolution,”in Neanderthals Revisited:New Approaches and Perspectives,edited by T.Harrison and K.Harvati(New York:Sinauer,2006).

[11]J.Sweeney and T.Samansky,“Elements of Successful Facility Design:Marine Mammals,”in Conservation of Endangered Species in Captivity:An Interdisciplinary Approach,edited by E.F.Gibbons,B.Durrant,and J.Demarest(New York:State University of New York Press,1995).

[12]关于混合尼安德特人基因被认为不显著这一点,可见(举例):Chris Stringer and Peter Andrews,The Complete World of Human Evolution(London:Thames and Hudson,2005)。或可见:Klein,The Dawn of Human Culture.

[13]Milford Wolpoff et al.,“Modern Human Ancestry at the Peripheries:A Test of the Replacement Theory,”Science 291,no.5502(2001):293—297.

[14]可以想到的就有芝加哥大学的玛莎·汉博林和安娜·迪·里恩佐在少数意大利人身上发现的达菲(Duffy)血型基因附近一个区的极端特殊形式,与我们在所有其他人类身上看到的都很不一样。M.T.Hamblin and A.Di Rienzo,“Detectionof the Signature of Natural Selection in Humans:Evidence from the Duffy Blood Group Locus,”American Journal of Human Genetics 66,no.5(2000):1669–1679.

[15]Amos Zayed and Charles W.Whitfield,“A Genome-Wide Signature of Positive Selection in Ancient and Recent Invasive Expansions of the Honey Bee Apis mellifera,”Proceedings of the National Academy of Sciences 105,no.9(2008):3421.

[16]H.C.Stutz and L.K.Thomas,“Hybridization and Introgression in Cowania and Purshia,”Evolution 18,no.2(1964):183—195.

[17]Brian Hare et al.,“The Domestication of Social Cognition in Dogs,”Science 298,no.5598(2002):1634—1636.

[18]Henry Harpending and Jay Sobus,“Sociopathy as an Adaptation,”Ethology and Sociobiology 8,no.3(1987):63—72.

[19]Richard W.Bulliet,The Camel and the Wheel(Cambridge:Harvard University Press,1975).

[20]David W.Frayer,“Evolution at the European Edge:Neanderthal and Upper Paleolithic Relationships,”Préhistoire Européenne 2(1993):9—69;David W.Frayer,“Perspectives on Neandertals as Ancestors,”in Conceptual Issues in Modern Human Origins Research,edited by G.A.Clark and C.M.Willermet(New York:Aldine de Gruyter,1998),220—234.

[21]Cidália Duarte et al.,“The Early Upper Paleolithic Human Skeleton from the Abrigo do Lagar Velho(Portugal)and Modern Human Emergence in Iberia,”Proceedings of the National Academy of Sciences 96(1999):7604—7609;Eric Trinkaus,“Early Modern Humans,”Annual Review of Anthropology 34(2005):207—230.

[22]D.Serre et al.,“No Evidence of Neandertal mtDNA Contributionto Early Modern Humans,”PLoS Biology 2,no.3(2004):313—317;M.Currat and L.Excoffier,“Modern Humans Did Not Admix with Neanderthals during Their Range Expansion into Europe,”PLoS Biology 2,no.12(2004):e21;T.D.Weaver and C.C.Roseman,“Ancient DNA,Late Neandertal Survival,and Modern-Human–Neandertal Genetic Admixture,”Current Anthropology 46,no.4(2005):677—683.

[23]Daniel Garrigan et al.,“Deep Haplotype Divergence and Long-Range Linkage Disequilibrium at Xp21.1Provides Evidence That Humans Descend from a Structured Ancestral Population,”Genetics 170(2005a):1849—1856;Daniel Garrigan et al.,“Evidence for Archaic Asian Ancestry on the Human X Chromosome,”Molecular Biology and Evolution 22(2005b):189—192;J.Hardy et al.,“Evidence Suggesting That Homo neanderthalensis Contributed the H2MAPT Haplotype to Homo sapiens,”Biochemical Society Transactions 33(2005):582—585;Vincent Plagnol and Jeffrey D.Wall,“Possible Ancestral Structure in Human Populations,”PLoS Genetics 2(2006):e105;P.D.Evans et al.,“Microcephalin,a Gene Regulating Brain Size,Continues to Evolve Adaptively in Humans,”Science 309,no.5741(2005):1717—1720;P.D.Evans et al.,“Evidence That the Adaptive Allele of the Brain Size Gene Microcephalin Introgressed into Homo sapiens from an Archaic Homo Lineage,”Proceedings of the National Academy of Sciences 103,no.48(2006):18178.

[24]Vincent Plagnol and Jeffrey D.Wall,“Possible Ancestral Structurein Human Populations,”PLoS Genetics 2(2006):e105.

[25]Patrick D.Evans et al.,“Evidence That the Adaptive Allele of the Brain Size Gene Microcephalin Introgressed into Homo sapiens from an Archaic Homo Lineage,”Proceedings of the National Academy of Sciences USA 103,no.48(2006):18178.

[26]Graham K.Coop et al.,“The Timing of Selection at the Human FOXP2Gene,”Molecular Biology and Evolution 25,no.7(2008):1257.

[27]Johannes Krause et al.,“The Derived FOXP2Variant of Modern Humans Was Shared with Neandertals,”Current Biology 17,no.21(2007):1908—1912.

[28]James O’Connell and James Allen,“Dating the Colonization of Sahul(Pleistocene Australia—New Guinea):A Review of Recent Research,”Journal of Archaeological Science 31,no.6(2004):835—853.