104516585 Are Humans Still Evolving Recent and Ongoing Human Evolution

Embed Size (px)

Citation preview

  • 7/28/2019 104516585 Are Humans Still Evolving Recent and Ongoing Human Evolution

    1/4

    1

    Are humans still evolving? Recent and ongoing human evolutionLast edited 10 November 2011; Submitted for grade 17 November 2011, submitted to UA 26 April 2012

    1 ABSTRACTIt is easy to think that, on or shortly following the appearance of

    Homo sapiens, humans stopped evolving. After all, the ability of

    humans to discover, refine, and pass on techniques of tool use has

    provided an ever-increasing degree of protection from the threats

    of the natural world, from the earliest fires ability to ward off

    potential predators and lessen the effects of cold weather to mod-

    ern medical practices which make disease almost irrelevant to

    some segments of human society. However, numerous studies have

    shown quite conclusively that this is not the case: the evolution of

    mankind has never halted nor slowed, and nothing indicates that it

    ever will. Human culture itself has a powerful ability to alter and

    even create selection, such as the selection for adult lactase persis-

    tence conferred by the development of dairy farming, or the cultur-

    al transmission of mate-choice preferences.

    Selection since the radiation ofH. sapiens from Africa is easy to

    prove by detecting evidence of strong (and ethnically varying)

    selective sweeps in the human genome, and even in modern times,

    evolution can be demonstrated over only a few generations. In

    recent years, human resistance to heart disease has risen, and fe-

    cundity has increased markedly due to decreases in maternal age at

    first birth, shorter inter-birth interval, and later age at menopause.

    This change in fecundity is also associated with changes in growth:

    in developed nations, female height is increasing, while in devel-

    oping nations it decreases, due to a well-established tradeoff be-

    tween growth and reproduction, and varying food availability.

    2 INTRODUCTIONAre humans still evolving? Though at one point in time, this was

    denied by some of the top scientific minds of the age (Talt 1869),

    now the popular consensus is established to such an extent that

    even popular news media sources (Harrel 2009; Bootle 2011) are

    joining in stating that, yes, Homo sapiens is still subject to the

    process of evolution by natural selection despite advances in

    medicine and technology that mean most humans are able to

    survive and reproduce.

    The reason for this is simple: while most humans may survive

    and reproduce, some will reproduce more than others, and any

    heritable traits that improve reproductive success will be passed on

    to the next generation at a disproportionately high rate, causing the

    population to evolve.

    This paper will provide proof for the claim that humans are

    indeed evolving, citing studies demonstrating human evolution on

    macro and micro time scales and exploring in humans the

    processes of natural and sexual selection with regard to the

    characteristically human trait, culture.

    3 GENE-CULTURE COEVOLUTIONBefore exploring cases and evidence of human evolution, we

    must first define and address the issue of gene-culture coevolution,

    a process that has had great impact on human evolutionary change.

    Richerson et al define culture as all of the information that

    individuals acquire from other individuals in a variety of social

    learning processes including teaching and imitation (2010).

    Culture serves as a form of information parallel to genetic

    information, and unlike genetic information, culture can be

    transmitted in multiple ways, horizontally from related or unrelated

    individuals or groups as well as vertically from parent to child

    (Feldman and Laland 1996).

    Gene-culture coevolution is the impact on genetic evolution of

    culture, which can create changes in either human preferences or

    their environmental conditions. These changes will increase anddecrease the relative fitness of different genotypes, thereby

    describing selection and causing evolution.

    It should be noted that cultural change does not always increase

    selection; culture can also serve to soften selection. For example,

    humans moving into a cold environment adapted to this change by

    wearing more clothes and building fires and shelters to keep warm

    rather than evolving physiological tolerance to the cold (Laland

    2008). But selection cannot be eliminated entirely, and even in this

    specific example there is evidence for human genetic adaptation to

    different environments (Williamson et al 2007), though none

    would ague that cultural change is not the crucial factor in human

    adaptation to hostile environments. As well, the use of advanced

    medicine unquestionably softens selective pressures that might

    otherwise favor humans with greater disease resistance, though as

    will be discussed later, this merely creates alternative selective

    pressures.

    Despite this dampening effectwhich is arguably the most

    potent adaptation humans possessculture does enact vast

    changes on the human genome via niche-construction and sexual

    selection. It has been argued by several groups that gene-culture

    coevolution may in fact be the dominant form of human evolution

    (Laland et al 2010, Richerson et al 2010), likely affecting hominid

    evolution since before the divergence from the other great apes

    (Richerson et al 2010) and possibly even serving to accelerate

    human evolution (Hawks et al 2007).

    3.1 Culture can affect genetic selection by changingthe environment

    Perhaps the most pervasive way in which gene-culture

    coevolution can act is by cultural change enacting a similar change

    on the surrounding environment, which then affects genetic

    selection.

  • 7/28/2019 104516585 Are Humans Still Evolving Recent and Ongoing Human Evolution

    2/4

    09007659

    2

    To use a favorite example of gene-culture coevolution

    proponents, evidence indicates that the spread of the lactase gene,

    which allows adult humans to retain an enzyme that allows more

    efficient digestion of dairy products, was preceded by the spread of

    cattle domestication and therefore dairy consumption (Tishkoff

    2007). The implications of this are clear: the cultural trait of cattle

    domestication, which provided a reliable food source to early

    humans, spread throughout populations in Europe, North Africa,

    and the Middle East. At some point, a genetic mutation arose

    which allowed possessors to get even more nutrients out of the

    presence of domesticated cattle, and it too spread throughout the

    region.

    The case of the lactase gene is one of great importance to the

    question of contemporary human evolution, and will be addressed

    further in section 3. Relevant to this section is the fact that the

    lactase gene spread as a result of the cultural development of dairy

    farming, a clear-cut example of culture affecting changes in the

    genome.

    3.2 Culture can affect genetic selection by changingmating preferences

    In addition to altering the selection based on the environment,

    culture may also affect sexual selection in humans. Recent studies

    have experimentally demonstrated mate-choice copying in humans

    (Little et al 2011), showing that human mate choice is dependent

    on social learning. The link between cultural groups and mate

    preference is well established (Boyd and Richerson 1985),

    showing that any two arbitrarily selected cultures will have

    different patterns of mate preference. These studies, and other

    modeling done by Laland (1994) indicate that any arbitrary mating

    preference for a heritable and phenotypically obvious trait such as

    hair or eye color can spread throughout a population via culturaltransmission, producing sexual selection. In addition, Ihara et al

    (2003) have shown that according to models of genetic and cultural

    transmission, female mate choice and male traits may link and

    produce runaway sexual selection that could drastically alter the

    phenome of the species.

    4 RECENT HUMAN EVOLUTIONOver the past several thousand years, human adaptive change

    has not slowed or stopped but accelerated (Hawks et al 2007);

    recent studies into the human genome have detected literally

    thousands of genes that have been subject to detectable positive

    selection sweeps (Nielsen et al 2007). Notable among these arestudies that show increased numbers of selection sweeps among

    non-African populations, probably a result of adaptation after the

    colonization of new environments (Williamson et al 2007),

    increased numbers of complete sweeps in non-African populations

    (Coop et al 2009), and studies showing the evidence of powerful

    selection sweeps in well-studied genes conferring malaria

    resistance and lactose digestion (Sabeti et al 2006). Additionally,

    these studies do not provide the whole picture, and much more

    investigation is needed to conclusively state how much of the

    human genome has been subjected to selection (Wang et al 2006).

    In the case of the lactase gene, we see a gene that was not only

    subjected to one of the highest rates of positive selection known

    (Bersagleri et al 2004), but one that is linked to ethnic background.

    The lactase gene is most prominent among peoples of European

    ancestry, and occurs at rates approaching zero in East Asian groups

    (Bersagleri et al 2004), indicating that the selective pressure for the

    lactase gene occurred after the separation of populations that

    colonized Europe and East Asia, and after the advent of dairy

    farming. This claim is supported by other studies linking the

    degree of presence of the lactase gene with cultural use of

    fermented and unfermented dairy products (Laland 2010).

    Clearly, humans have evolved over the past ten thousand years,

    and in patterns that vary with location, as one would expect for any

    other organism that has experienced changing climates and

    colonized new habitats.

    5 ONGOING HUMAN EVOLUTIONAlthough the typical argument stating that humans are not

    evolving hinges upon the massive advances in medicine and

    technology that have taken place over the past few centuries

    (McKie 2002), numerous studies have detected measureable

    evolution over even decades, in consistent patterns the world over.

    5.1 Improving cardiovascular healthDespite advances in medicine, cardiovascular disease kills an

    estimated 17.3 million people per year, most of who live in

    developing nations (World Health Organization 2011). Like any

    other common fatal health risk, this creates selective pressure on

    the species, favoring those individuals who are less prone tocardiovascular diseases. Though any overall improvement in

    cardiovascular health could be attributed to improving public

    knowledge and healthcare, studies taking into account these

    confounding factors indicate that systolic blood pressure and LDL

    cholesterol levels (two predictors of cardiovascular disease) have

    dropped over the past 60 years (Byars et al 2010, Stearns et al

    2010). It is unclear to what degree this improvement in

    cardiovascular health is truly due to genetic evolution rather than

    scientific advancement, but sources indicate that at least part of the

    improvement in health has a genetic basis (Stearns et al 2010).

    5.2 Changes in heightAdult height is associated in humans with levels of nutrition

    (Frisch 1978), and with reproductive success, though differently in

    males and females. In male humans, height is a common sexually

    selected trait (Hensley 1994) that correlates with male reproductive

    success: the men with the peak reproductive success are above the

    populations mean height (Nettle 2002a). The inverse is true in

    females, where there is a tradeoff between high adult height and

    early sexual maturity, so females of a short stature (and more

  • 7/28/2019 104516585 Are Humans Still Evolving Recent and Ongoing Human Evolution

    3/4

    Are humans still evolving? Recent and ongoing human evolution

    3

    importantly, early maturation) have greater lifetime reproductive

    success (Helle 2008), though the directional selection on height is

    less pronounced for women than it is for men (Nettle 2002b).

    Studies in the developing world yield conflicting results: height is

    positively correlated with female reproductive success (Allal 2004;

    Sear et al 2004), likely due to varying levels in nutrition. In these

    nations, there is a higher infant mortality rate for the offspring of

    short women. This variation in selection by location implies that

    changes in height are implicitly tied to levels of nutrition, and that

    selection will only act to increase male height and decrease female

    height if the population has the resources to do so.

    5.3 Widening reproductive windowBy far the most consistent finding among all studies into human

    evolution is that the female reproductive window is widening.

    Short-period investigations into changes from generation to

    generation in populations in Quebec (Milot et al 2011), Finland

    (Helle et al 2005), Gambia (Allal et al 2004), the USA (Byars et al

    2010), Australia (Kirk et al 2001), Kenya (Borgehoff 1989), Nepal(Gubhaju 1983), and Sweden (Low 1991) have all yielded

    consistent results: over the past century low maternal age at first

    birth correlates strongly with lifetime reproductive success. This

    not only affects women: a 1996 study by Kr et al showed that in

    monogamous societies the age of a mans wife was by far the

    greatest indicator of his lifetime reproductive success.

    As well as early reproduction, studies have linked shorter inter-

    birth interval (Stearns et al 2010) and later age at menopause with

    lifetime reproductive success (Kirk et al 2001; Byars et al 2010),

    though the correlation between these traits and fitness is far lower

    than that between first reproduction and fitness (Stearns et al

    2010).

    As with selection on height, reproductive ability is intrinsicallylinked to food availability (Frisch 1978), which has increased

    tremendously in recent generations. Any organism with unlimited

    access to resources will reproduce as rapidly as possible, and over

    the past centuries humans have not only been reproducing at a

    maximum rate, but that maximum rate increases as humans evolve

    to reproduce more often and from an earlier age to a later age.

    6 CONCLUSIONThe idea that humankind has somehow removed itself from the

    effects of natural selection seems rooted in the vain idea that we

    are better than other species, and above the effects that drive

    changes in lesser animals. And yet, investigating the questionillustrates quite clearly that we are subject to the same processes

    that govern the evolution of every other organism: natural selection

    causes genes that allow digestion of a readily available food source

    or increase resistance to common diseases to spread amongst a

    population, sexual selection causes the proliferation of arbitrary

    male-trait and female-choice genes, and when food is available in

    excess and the likelihood of death by disease is at an all-time low,

    the humans which reproduce fastest win themselves a greater stake

    in the genetic composition of the next generation. Even culture, a

    feature of the species lauded as something that somehow separates

    us from the animals, has a tremendous effect on the selective

    pressures that drive human evolution. Numerous studies have

    proven that humans have continued to evolve over the past five

    thousand years, and that we will continue to do so with no sign or

    indication that that will ever change.

    REFERENCES

    Allal N, Sear R, Prentice A M, Mace R. (2004) An evolutionary model of statue. Age

    at first birth and reprodictove success in Gambian women. Proceedings of the

    Royal Society of London: Series B271 465-470

    Bersagleri T, Sabeti PC, Patterson N, Vanderploeg T, Scaffner SF, Drake JA, Rhodes

    M, Reich DE, Hirschhorn JN. (2004) Genetic signatures of strong recent positive

    selection at the lactase gene.American Journal of Human Genetics74 1111-1120

    Bootle O (2011) Are humans still evolving by Darwins natural selection? BBC

    Science and Environment, 1 March 2011

    Boyd R, Richerson PJ (1985) Culture and the evolutionary process. Chicago, IL:

    Chicago University Press.

    Borgehoff MM. (1989) Early maturing Kipsigis women have higher reproductive

    success than late maturing women and cost more to marry. Behavioral Ecology and

    Sociobiology 24 145-153Byars SG, Ewbank D, Govindaraju DH, Stearns SC. (2010) Natural selection in a

    contemporary human population.Proceedings of the National Academy of Science

    USA107 1787-1792.

    Coop G, Pickrell JK, Novembre J, Kudaravalli S, Li J, Absher D, Myers R M, Cavalli-

    Sforza LL, Feldman MW, Pritchard JK. (2009) The role of geography in human

    adaptation.PLoS Genetics5 (6)

    Feldman MW, Laland KN. (1996) Gene-culture coevolutionary theory. Trends in

    Ecology and Evolution 11 (11)

    Frisch RE 1978. Population, food intake, and fertility. There is historical evidence for

    a direct effect of nutrition on reproductive ability. Science 199 22-30

    Gubhaju B. (1983) Fertility differentials in Nepal. Journal of Biosociological Science

    15 325-331

    Harrel E (2009) Darwin lives! Modern humans are still evolving. Time Science, 23

    October 2009

    Hawks J, Wang ET, Cochran GM, Harpending HC, Moysiz RK. (2007) Recent

    acceleration of human adaptive evolution. Proceedings of the National Academy of

    Science USA104 (52) 20753-20758

    Helle S, Lummaa V, Jokela J. (2005) Are reproductive and somatic senescence

    coupled in humans? Late, but not early, reproduction correlated with longevity in

    historical Sami women.Proceedings of the Royal Society of Biology272

    Helle S. (2008) A tradeoff between reproduction and growth in contemporary Finnish

    women.Evolution and Human Behavior29

    Hensley WE (1994) Height as a basis for interpersonal attraction. Adolescence 29

    (114) 469-474

    Ihara Y, Aoki K, Feldman MW (2003) Runaway sexual selection with paternal

    transmission of the male trait and gene-culture determination of the female

    preference. Theoretical Population Biology63 (1) 53-62

    Kr P, Jokela J, Helle T, Kojola I. (1996) Direct and correlative phenotypic selection

    on life-history traits in three pre-industrial human populations. Proceedings of the

    Royal Society of London263 1475-1480

    Kirk KM, Blomberg SP, Duffy DL, Heath AC, Owens IPF (2001)Natural se lection

    and quantitative genetics of life-history traits in western women: a twin study.

    Evolution55 (2) 423-435Laland KN (1994) Sexual selection with a culturally transmitted mating preference.

    Theoretical Population Biology 45 (1) 1-15

    Laland KN (2008) Exploring gene-culture interactions: insights from handedness,

    sexual selection and niche-construction case studies. Philosophical Transactions of

    the Royal Society363 3577-3589

    Laland KN, Odling-Smee J, Myles S (2010) How culture has shaped the human

    genome: bringing genetics and the human sciences together. Nature Reviews

    Genetics11 137-148

    Little AC, Jones BC, DeBruine LM, Caldwell CA (2011) Social learning and human

    mate preferences: a potential mechanism for generating and maintaining between-

  • 7/28/2019 104516585 Are Humans Still Evolving Recent and Ongoing Human Evolution

    4/4

    09007659

    4

    population diversity in attract ion.Philosophical Transactions of The Royal Society

    Biological Sciences366 (1563) 366-375

    Low BS. (1991) Reproductive life in nineteenth century Sweden: an evolutionary

    perspective on demographic phenomena.Ethnology and Sociobiology12 411-448

    Milot E, Mayer F M, Nussey H, Boisvert M, Pelletier F, Rale D. (2011) Evidence for

    evolution in response to natural selection in a contemporary human population.

    Proceedings of the National Academy of Science USA108 (41) 17040-17045

    McKie R (2002) Is human evolution finally over? The Observer, 3 February 2002.

    Nielsen R, Hellmann I, Hubisz M, Bustamante C, Clark AG. (2007) Recent and

    ongoing selection in the human genome.Nature Reviews Genetics8.Nettle D. (2002) Height and reproductive success in a cohort of Br itish men. Human

    Nature 13 (4) 473-491

    Nettle D (2002) Womens height, reproductive success and the evolution of sexual

    dimorphism in modern humans. Proceedings of the Royal Society London 269

    1919-1923

    Richerson PJ, Boyd R, Henrich J (2010) Gene-culture coevolution in the age of

    genomics. Proceedings of the National Academy of Science USA 107 (2) 8982-

    8992

    Sear R, Allal N, Mace R, McGregor I. (2004) Height and reproductive success among

    Gambian women.American Journal of Human Biology 16 223-223.

    Stearns SC, Byars SG, Diddahally RG, Ewbank D. (2010) Measuring selection in

    contemporary human populations.Nature Reviews Genetics 11 611-622

    Sabeti PC, Schaffner SF, Fry B, Lohmueller J, Varilly P, Shamovsky O, Palma A,

    Mikkelsen TS, Altshuler D, Lander ES. (2006) Positive natural selection in the

    human lineage. Science312

    Talt L. (1869) Has the law of natural selection by survival of the fittest failed in thecase of man?Dublin Quarterly Journal of Medical Science47 102-113

    Tishkoff SA (2007) Convergent adaptation of human lactase persistence in Africa and

    Europe.Nature Genetics39 31-40

    Wang ET, Kodama G, Baldi P, Moyzis R K. (2006) Global landscape of recent

    inferred Darwinian selection for Homo sapiens. Proceedings of the National

    Academy of Science USA103 (1) 135-140

    Williamson SH, Hubisz MJ, Clark AG, Payseur BA, Bustamante CD (2007)

    Localizing recent adaptive evolution in the human genome. PLoS Genetics3 (6)

    World Health Organization, 2011. Cardiovascular disease [Online] Available at

    http://www.who.int/cardiovascular_diseases/en/. [Accessed 10 November 2011]

    http://www.who.int/cardiovascular_diseases/en/http://www.who.int/cardiovascular_diseases/en/http://www.who.int/cardiovascular_diseases/en/