鲍德温效应

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模板:Distinguish

文件:Lamarck Compared to Darwin, Baldwin, Waddington.svg
The Baldwin effect compared to Lamarck's theory of evolution, Darwinian evolution, and Waddington's genetic assimilation. All the theories offer explanations of how organisms respond to a changed environment with adaptive inherited change.

Lamarck's theory of evolution, Darwinian evolution, and Waddington's genetic assimilation. All the theories offer explanations of how organisms respond to a changed environment with adaptive inherited change.]]

拉马克的进化论、达尔文进化论和华丁顿的基因同化论。[参考译文]所有的理论都解释了有机体是如何通过适应性遗传变化来应对环境变化的

In evolutionary biology, the Baldwin effect describes the effect of learned behavior on evolution. In brief, James Mark Baldwin and others suggested during the eclipse of Darwinism in the late 19th century that an organism's ability to learn new behaviors (e.g. to acclimatise to a new stressor) will affect its reproductive success and will therefore have an effect on the genetic makeup of its species through natural selection. Though this process appears similar to Lamarckian evolution, Lamarck proposed that living things inherited their parents' acquired characteristics. The Baldwin effect has been independently proposed several times, and today it is generally recognized as part of the modern synthesis.

In evolutionary biology, the Baldwin effect describes the effect of learned behavior on evolution. In brief, James Mark Baldwin and others suggested during the eclipse of Darwinism in the late 19th century that an organism's ability to learn new behaviors (e.g. to acclimatise to a new stressor) will affect its reproductive success and will therefore have an effect on the genetic makeup of its species through natural selection. Though this process appears similar to Lamarckian evolution, Lamarck proposed that living things inherited their parents' acquired characteristics. The Baldwin effect has been independently proposed several times, and today it is generally recognized as part of the modern synthesis.

在进化生物学中,鲍德温效应描述了学习行为对进化的影响。简而言之,在19世纪末达尔文主义的日蚀期间,詹姆斯·马克·鲍德温和其他人提出,生物体学习新行为的能力(例如:。为了适应新的压力)会影响它的繁殖成功,因此会通过自然选择影响其物种的基因组成。虽然这个过程与拉马克的进化过程相似,但拉马克认为生物继承了他们父母获得的特征。鲍德温效应已经被独立地提出了好几次,今天它被普遍认为是现代综合的一部分。


"A New Factor in Evolution"

The effect, then unnamed, was put forward in 1896 in a paper "A New Factor in Evolution" by American psychologist James Mark Baldwin. The paper proposed a mechanism for specific selection for general learning ability. As Robert Richards explains:[1]

The effect, then unnamed, was put forward in 1896 in a paper "A New Factor in Evolution" by American psychologist James Mark Baldwin. The paper proposed a mechanism for specific selection for general learning ability. As Robert Richards explains:

1896年,美国心理学家詹姆斯·马克·鲍德温在一篇名为《进化的新因素》的论文中提出了这一效应。本文提出了一种针对一般学习能力的具体选择机制。正如罗伯特 · 理查兹解释的那样:


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If animals entered a new environment—or their old environment rapidly changed—those

If animals entered a new environment—or their old environment rapidly changed—those

如果动物进入一个新的环境,或者它们原来的环境迅速发生变化,那么

that could flexibly respond by learning new behaviors or by ontogenetically adapting would

that could flexibly respond by learning new behaviors or by ontogenetically adapting would

可以通过学习新的行为或者通过个体遗传适应灵活地作出反应

be naturally preserved. This saved remnant would, over several generations, have the

be naturally preserved. This saved remnant would, over several generations, have the

自然保存。这些被保存下来的残余物,经过几代后,会有

opportunity to exhibit spontaneously congenital variations similar to their acquired traits and

opportunity to exhibit spontaneously congenital variations similar to their acquired traits and

有机会表现出自发的先天性变异,类似于他们获得的特征和

have these variations naturally selected. It would look as though the acquired traits had sunk

have these variations naturally selected. It would look as though the acquired traits had sunk

这些变化是自然选择的。这看起来好像后天获得的特征已经消失了

into the hereditary substance in a Lamarckian fashion, but the process would really be neo-Darwinian.

into the hereditary substance in a Lamarckian fashion, but the process would really be neo-Darwinian.

以拉马克的方式将其转化为遗传物质,但这个过程实际上是新达尔文主义的。

/ blockquote


Selected offspring would tend to have an increased capacity for learning new skills rather than being confined to genetically coded, relatively fixed abilities. In effect, it places emphasis on the fact that the sustained behavior of a species or group can shape the evolution of that species. The "Baldwin effect" is better understood in evolutionary developmental biology literature as a scenario in which a character or trait change occurring in an organism as a result of its interaction with its environment becomes gradually assimilated into its developmental genetic or epigenetic repertoire (Simpson, 1953; Newman, 2002). In the words of Daniel Dennett,[2]

Selected offspring would tend to have an increased capacity for learning new skills rather than being confined to genetically coded, relatively fixed abilities. In effect, it places emphasis on the fact that the sustained behavior of a species or group can shape the evolution of that species. The "Baldwin effect" is better understood in evolutionary developmental biology literature as a scenario in which a character or trait change occurring in an organism as a result of its interaction with its environment becomes gradually assimilated into its developmental genetic or epigenetic repertoire (Simpson, 1953; Newman, 2002). In the words of Daniel Dennett,

被选中的后代往往具有更强的学习新技能的能力,而不是局限于基因编码,相对固定的能力。实际上,它强调了一个事实,即一个物种或群体的持续行为可以塑造该物种的进化。在演化发育生物学文学中,“ Baldwin 效应”被更好地理解为一个场景,在这个场景中,一个有机体由于与其环境的相互作用而发生的性格或特征变化逐渐被同化到其发育的遗传或表观遗传剧目中(Simpson,1953; Newman,2002)。用丹尼尔 · 丹尼特的话来说,


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Thanks to the Baldwin effect, species can be said to pretest the efficacy of particular different designs by phenotypic (individual) exploration of the space of nearby possibilities. If a particularly winning setting is thereby discovered, this discovery will create a new selection pressure: organisms that are closer in the adaptive landscape to that discovery will have a clear advantage over those more distant.

Thanks to the Baldwin effect, species can be said to pretest the efficacy of particular different designs by phenotypic (individual) exploration of the space of nearby possibilities. If a particularly winning setting is thereby discovered, this discovery will create a new selection pressure: organisms that are closer in the adaptive landscape to that discovery will have a clear advantage over those more distant.

由于鲍德温效应,可以说物种通过表型(个体)探索附近空间的可能性,预先测试了特定不同设计的功效。如果一个特别成功的环境被发现,这个发现将创造一个新的选择压力: 在适应环境中更接近这个发现的有机体将比那些更遥远的有机体有明显的优势。

/ blockquote


An update to the Baldwin effect was developed by Jean Piaget, Paul Weiss, and Conrad Waddington in the 1960s–1970s. This new version included an explicit role for the social in shaping subsequent natural change in humans (both evolutionary and developmental), with reference to alterations of selection pressures.[3]

An update to the Baldwin effect was developed by Jean Piaget, Paul Weiss, and Conrad Waddington in the 1960s–1970s. This new version included an explicit role for the social in shaping subsequent natural change in humans (both evolutionary and developmental), with reference to alterations of selection pressures.

1960-1970年代,让 · 皮亚杰、保罗 · 维斯和康拉德 · 华丁顿对鲍德温效应进行了更新。这个新的版本包括了社会在塑造人类随后的自然变化(包括进化和发展)中的明确作用,并提到了选择压力的改变。


As is to be expected from Stigler's law, subsequent research shows that Baldwin was not the first to identify the process; Douglas Spalding mentioned it in 1873.[4]

As is to be expected from Stigler's law, subsequent research shows that Baldwin was not the first to identify the process; Douglas Spalding mentioned it in 1873.

正如斯蒂格勒定律所预期的那样,随后的研究表明,Baldwin 并不是第一个发现这个过程的人; 道格拉斯·斯普拉丁在1873年提到过这个过程。


Examples

Suppose a species is threatened by a new predator and there is a behavior that makes it more difficult for the predator to kill individuals of the species. Individuals who learn the behavior more quickly will obviously be at an advantage. As time goes on, the ability to learn the behavior will improve (by genetic selection), and at some point it will seem to be an instinct. Baldwin gives the following case involving cooperation: "Animals may be kept alive let us say in a given environment by social cooperation only; these transmit this social type of variation to posterity; thus social adaptation sets the direction of physical phylogeny and physical heredity is determined in part by this factor" (Baldwin, 1896, p. 553).

Suppose a species is threatened by a new predator and there is a behavior that makes it more difficult for the predator to kill individuals of the species. Individuals who learn the behavior more quickly will obviously be at an advantage. As time goes on, the ability to learn the behavior will improve (by genetic selection), and at some point it will seem to be an instinct. Baldwin gives the following case involving cooperation: "Animals may be kept alive let us say in a given environment by social cooperation only; these transmit this social type of variation to posterity; thus social adaptation sets the direction of physical phylogeny and physical heredity is determined in part by this factor" (Baldwin, 1896, p. 553).

假设一个物种受到一个新的捕食者的威胁,并且有一种行为使得捕食者更加难以杀死该物种的个体。更快学会这种行为的个体显然会处于有利地位。随着时间的推移,学习行为的能力将得到提高(通过基因选择) ,在某种程度上,这似乎是一种本能。鲍德温给出了以下涉及合作的案例: “动物可以保持活着,让我们说,在一个特定的环境中,只有通过社会合作; 这些传递这种社会类型的变异给子孙后代; 因此,社会适应确定了物理系统发育和物理遗传的方向,部分由这个因素决定”(鲍德温,1896年,第553页)。


The appearance of lactose tolerance (i.e., lactase persistence) in human populations with a long tradition of raising domesticated animals for milk production has been suggested as another example.[5] This argument holds that a feedback loop operates whereby a dairy culture increases the selective advantage from this genetic trait, while the average population genotype increases the collective rewards of a dairy culture.

The appearance of lactose tolerance (i.e., lactase persistence) in human populations with a long tradition of raising domesticated animals for milk production has been suggested as another example. This argument holds that a feedback loop operates whereby a dairy culture increases the selective advantage from this genetic trait, while the average population genotype increases the collective rewards of a dairy culture.

另一个例子是,在人类中出现了乳糖耐受性(即乳糖分解酶持续性) ,而人类在饲养家畜产奶方面有着悠久的传统。这种观点认为,乳制品文化通过一个反馈循环增加了这种遗传特性的选择优势,而普通群体的基因型增加了乳制品文化的集体回报。


Controversy and acceptance

Initially Baldwin's ideas were not incompatible with the prevailing, but uncertain, ideas about the mechanism of transmission of hereditary information and at least two other biologists put forward very similar ideas in 1896.[6][7] In 1901, Maurice Maeterlinck referred to behavioural adaptations to prevailing climates in different species of bees as ‘what had merely been an idea, therefore, and opposed to instinct, has thus by slow degrees become an instinctive habit’.[8] The Baldwin effect theory subsequently became more controversial, with scholars being split between "Baldwin boosters" and "Baldwin skeptics".[9] The theory was first called the "Baldwin effect" by George Gaylord Simpson in 1953.[9] Simpson "admitted that the idea was theoretically consistent, that is, not inconsistent with the modern synthesis",[9] but he doubted that the phenomenon occurred very often, or if so, could be proven to occur. In his discussion of the reception of the Baldwin-effect theory Simpson points out that the theory appears to provide a reconciliation between a neo-Darwinian and a neo-Lamarckian approach and that "Mendelism and later genetic theory so conclusively ruled out the extreme neo-Lamarckian position that reconciliation came to seem unnecessary".[10] In 1942, the evolutionary biologist Julian Huxley promoted the Baldwin effect as part of the modern synthesis, saying the concept had been unduly neglected by evolutionists.[11]

Initially Baldwin's ideas were not incompatible with the prevailing, but uncertain, ideas about the mechanism of transmission of hereditary information and at least two other biologists put forward very similar ideas in 1896. In 1901, Maurice Maeterlinck referred to behavioural adaptations to prevailing climates in different species of bees as ‘what had merely been an idea, therefore, and opposed to instinct, has thus by slow degrees become an instinctive habit’. The Baldwin effect theory subsequently became more controversial, with scholars being split between "Baldwin boosters" and "Baldwin skeptics". The theory was first called the "Baldwin effect" by George Gaylord Simpson in 1953. In 1942, the evolutionary biologist Julian Huxley promoted the Baldwin effect as part of the modern synthesis, saying the concept had been unduly neglected by evolutionists.

最初,鲍德温的想法并不与盛行的,但不确定的,关于遗传信息传递机制的想法不相容,至少其他两个生物学家在1896年提出了非常相似的想法。1901年,莫里斯·梅特林克提到不同种类的蜜蜂对当时气候的行为适应性,称之为仅仅是一种想法,因此,与本能相反,慢慢地变成了一种本能的习惯。鲍德温效应理论随后变得更具争议性,学者们被分为“鲍德温支持者”和“鲍德温怀疑论者”。这个理论在1953年首次被乔治·盖洛德·辛普森称为“ Baldwin 效应”。1942年,进化生物学家朱利安 · 赫胥黎宣称鲍德温效应是现代综合论的一部分,说这个概念被进化论者过度忽视了。


In the 1960s, the evolutionary biologist Ernst Mayr contended that the Baldwin effect theory was untenable because

In the 1960s, the evolutionary biologist Ernst Mayr contended that the Baldwin effect theory was untenable because

在20世纪60年代,进化生物学家恩斯特 · 迈尔声称鲍德温效应理论是站不住脚的,因为

  1. the argument is stated in terms of the individual genotype, whereas what is really exposed to the selection pressure is a phenotypically and genetically variable population;
the argument is stated in terms of the individual genotype, whereas what is really exposed to the selection pressure is a phenotypically and genetically variable population;

这个论点是从个体基因型的角度来阐述的,而真正暴露在选择压力下的是表型和遗传可变的种群;

  1. it is not sufficiently emphasized that the degree of modification of the phenotype is in itself genetically controlled;
it is not sufficiently emphasized that the degree of modification of the phenotype is in itself genetically controlled;

没有充分强调表型的改变程度本身是受遗传控制的;

  1. it is assumed that phenotypic rigidity is selectively superior to phenotypic flexibility.[12]
it is assumed that phenotypic rigidity is selectively superior to phenotypic flexibility.

假定表型刚性有选择地优于表型柔性。


In 1987 Geoffrey Hinton and Steven Nowlan demonstrated by computer simulation that learning can accelerate evolution, and they associated this with the Baldwin effect.[13][14][15]

In 1987 Geoffrey Hinton and Steven Nowlan demonstrated by computer simulation that learning can accelerate evolution, and they associated this with the Baldwin effect.

1987年,Geoffrey Hinton 和 Steven Nowlan 通过计算机模拟证明了学习可以加速进化,他们将其与 Baldwin 效应联系起来。


Paul Griffiths[16] suggests two reasons for the continuing interest in the Baldwin effect. The first is the role mind is understood to play in the effect. The second is the connection between development and evolution in the effect. Baldwin’s account of how neurophysiological and conscious mental factors may contribute to the effect[17][18][19] brings into focus the question of the possible survival value of consciousness.[20]

Paul Griffiths suggests two reasons for the continuing interest in the Baldwin effect. The first is the role mind is understood to play in the effect. The second is the connection between development and evolution in the effect. Baldwin’s account of how neurophysiological and conscious mental factors may contribute to the effect brings into focus the question of the possible survival value of consciousness.

保罗 · 格里菲斯提出了人们对鲍德温效应持续兴趣的两个原因。首先,我们可以理解心智在这种效果中所扮演的角色。二是发展与演变的关系。鲍德温关于神经生理学和有意识的心理因素如何促成这种效应的论述,使人们更加关注意识可能的生存价值问题。


文件:Carpodacus mexicanus -Madison, Wisconsin, USA-8.jpg
The house finch's colonisation of North America has provided empirical evidence of the Baldwin effect.[21]

The house finch's colonisation of North America has provided empirical evidence of the Baldwin effect.

[雀鸟在北美的殖民活动提供了鲍德温效应的经验证明。]


Still, observes David Depew, "it is striking that a rather diverse lot of contemporary evolutionary theorists, most of whom regard themselves as supporters of the Modern Synthesis, have of late become 'Baldwin boosters模板:'"[9] These Baldwin boosters

Still, observes David Depew, "it is striking that a rather diverse lot of contemporary evolutionary theorists, most of whom regard themselves as supporters of the Modern Synthesis, have of late become 'Baldwin boosters These Baldwin boosters

不过,戴维•迪普(David Depew)评论道: “令人惊讶的是,许多不同的当代进化论理论家,其中大多数人自认为是现代综合论(Modern Synthesis)的支持者,最近却成为了‘鲍德温的支持者,这些鲍德温


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are typically evolutionary psychologists who are searching for scenarios in which a population can get itself by behavioral trial and error onto a "hard to find" part of the fitness landscape in which human brain, language, and mind can rapidly coevolve. They are searching for what Daniel Dennett, himself a Baldwin booster, calls an "evolutionary crane," an instrument to do some heavy lifting fast.[9]

are typically evolutionary psychologists who are searching for scenarios in which a population can get itself by behavioral trial and error onto a "hard to find" part of the fitness landscape in which human brain, language, and mind can rapidly coevolve. They are searching for what Daniel Dennett, himself a Baldwin booster, calls an "evolutionary crane," an instrument to do some heavy lifting fast.

是典型的进化心理学家,他们正在寻找一个场景,在这个场景中,一个种群可以通过行为试验和错误到一个“难以找到”的适应景观的一部分,在这个景观中,人类的大脑,语言和思维可以迅速共同。他们正在寻找一种被丹尼尔 · 丹尼特称为“进化起重机”的工具,丹尼尔 · 丹尼特自己就是鲍德温的支持者。

/ blockquote


According to Daniel Dennett, recent work has rendered the Baldwin effect "no longer a controversial wrinkle in orthodox Darwinism".[2] Potential genetic mechanisms underlying the Baldwin effect have been proposed for the evolution of natural (genetically-determinant) antibodies.[22] In 2009, empirical evidence for the Baldwin effect was provided from the colonisation of North America by the house finch.[21]

According to Daniel Dennett, recent work has rendered the Baldwin effect "no longer a controversial wrinkle in orthodox Darwinism". Potential genetic mechanisms underlying the Baldwin effect have been proposed for the evolution of natural (genetically-determinant) antibodies. In 2009, empirical evidence for the Baldwin effect was provided from the colonisation of North America by the house finch.

丹尼尔•丹尼特(Daniel Dennett)表示,最近的工作已经使鲍德温效应“在正统达尔文主义中不再是一个有争议的问题”。潜在的遗传机制潜在的鲍德温效应已被提出的进化自然(遗传决定因素)抗体。2009年,经验证明在北美殖民时期为 Baldwin 效应提供了证据。


Comparison with genetic assimilation

模板:Further


The Baldwin effect has been confused with, and sometimes conflated with, a different evolutionary theory also based on phenotypic plasticity, C. H. Waddington's genetic assimilation. The Baldwin effect includes genetic accommodation, of which one type is genetic assimilation.[23]

The Baldwin effect has been confused with, and sometimes conflated with, a different evolutionary theory also based on phenotypic plasticity, C. H. Waddington's genetic assimilation. The Baldwin effect includes genetic accommodation, of which one type is genetic assimilation.

鲍德温效应与另一种进化理论相混淆,有时甚至混为一谈,这种进化理论也是基于表型可塑性的基因同化。鲍德温效应包括遗传调节,其中一种类型是遗传同化


See also


Notes

  1. Richards, Robert J. (1987). Darwin and the Emergence of Evolutionary Theories of Mind and Behavior. Chicago: The University of Chicago Press. p. 399. ISBN 978-0-226-71199-7. 
  2. 2.0 2.1 Dennett, Daniel (2003), "The Baldwin Effect: a Crane, not a Skyhook" in: Weber, Bruce H.; Depew, David J. (2003). Evolution and learning: The Baldwin effect reconsidered. Cambridge, MA: MIT Press. pp. 69–106. ISBN 978-0-262-23229-6. https://books.google.com/books?id=yBtRzBilw1MC&pg=PR9&dq=baldwin+effect#v=onepage&q&f=false. 
  3. Burman J. T. (2013). "Updating the Baldwin Effect: The biological levels behind Piaget's new theory". New Ideas in Psychology. 31 (3): 363–373. doi:10.1016/j.newideapsych.2012.07.003.
  4. Noble, R.; Noble, D. (2017) Was the Watchmaker Blind? Or Was She One-Eyed? Biology 2017, 6(4), 47; doi:10.3390/biology6040047, quoting Bateson, P. The adaptability driver: Links between behaviour and evolution. Biol. Theory 2006, 1, 342–345.
  5. Evolution and Learning: The Baldwin Effect Reconsidered
  6. Morgan, C. L. (1896). "On modification and variation". Science. 4 (99): 733–740. doi:10.1126/science.4.99.733. PMID 17735249.
  7. Osborne, H. F. (1896). "Ontogenic and phylogenic variation". Science. 4 (100): 786–789. doi:10.1126/science.4.100.786. PMID 17734840.
  8. Materlinck, Maurice (1901). The Life of the Bee. New York: Dodd, Mead and Co.. pp. Chapter VII section 102. 
  9. 9.0 9.1 9.2 9.3 9.4 Depew, David J. (2003), "Baldwin Boosters, Baldwin Skeptics" in: Weber, Bruce H.; Depew, David J. (2003). Evolution and learning: The Baldwin effect reconsidered. Cambridge, MA: MIT Press. pp. 3–31. ISBN 978-0-262-23229-6. https://books.google.com/books?id=yBtRzBilw1MC&pg=PR9&dq=baldwin+effect#v=onepage&q&f=false. 
  10. Simpson, George Gaylord (1953). "The Baldwin effect". Evolution. 7 (2): 110–117. doi:10.2307/2405746. JSTOR 2405746.
  11. Huxley, Julian (1942). Evolution: The Modern Synthesis. London: George Allen & Unwin Ltd. 
  12. Mayr, Ernst (1963). Animal Species and Evolution. Cambridge, MA: Harvard University Press. ISBN 978-0-674-03750-2. https://archive.org/details/animalspeciesevo00mayr. 
  13. Hinton, Geoffrey E.; Nowlan, Steven J. (1987). "How learning can guide evolution". Complex Systems. 1: 495–502.
  14. Maynard Smith, John (1987). "When learning guides evolution". Nature. 329 (6142): 761–762. doi:10.1038/329761a0. PMID 3670381.
  15. Puentedura, Ruben R. (2003). "The Baldwin effect in the age of computation". In Weber, Bruce H.. Evolution and Learning: The Baldwin Effect Reconsidered. Cambridge, MA: MIT press. pp. 219–234. 
  16. Griffiths, Paul E. (2003). "Beyond the Baldwin effect: James Mark Baldwin’s ‘social heredity,’ epigenetic inheritance, and niche construction". In Weber, Bruce H.. Evolution and Learning: The Baldwin Effect Reconsidered. Cambridge, MA: MIT press. pp. 193–215. 
  17. Baldwin, J. Mark (1896). "Heredity and instinct". Science. 3 (64): 438–441, 558–561. doi:10.1126/science.3.64.438. PMID 17780356.
  18. Baldwin, J. Mark (1896). "Consciousness and evolution". Psychological Review. 3 (3): 300–309. doi:10.1037/h0063996.
  19. Baldwin, J. Mark (1896). "A new factor in evolution". The American Naturalist. 30 (354): 441–451, 536–553. doi:10.1086/276408.
  20. Lindahl, B. I. B. (2001). "Consciousness, behavioural patterns and the direction of biological evolution: implications for the mind–brain problem". In Pylkkänen, Paavo. Dimensions of Conscious Experience. Amsterdam: John Benjamins. pp. 73–99. ISBN 978-90-272-5157-2. 
  21. 21.0 21.1 Badyaev, Alexander V. (March 2009). "Evolutionary significance of phenotypic accommodation in novel environments: an empirical test of the Baldwin effect". Phil. Trans. R. Soc. Lond. B. 364 (1520): 1125–1141. doi:10.1098/rstb.2008.0285. PMC 2666683. PMID 19324617.
  22. Anderson, Russell (1996). "How the adaptive antibodies facilitate the evolution of natural antibodies". Immunology and Cell Biology. 74 (2): 286–291. doi:10.1038/icb.1996.50. PMID 8799730.
  23. Crispo, Erika (2007). "The Baldwin effect and genetic assimilation: revisiting two mechanisms of evolutionary change mediated by phenotypic plasticity". Evolution. 61 (11): 2469–2479. doi:10.1111/j.1558-5646.2007.00203.x. PMID 17714500.


References

  • Cochran, Gregory; Harpending, Henry (2009). The 10,000 Year Explosion: How Civilization Accelerated Human Evolution. Basic Books. ISBN 978-0-465-00221-4. 
  • Hall, Brian K. (2001). "Organic Selection: Proximate Environmental Effects on the Evolution of Morphology and Behaviour". Biology and Philosophy. 16 (2): 215–237. doi:10.1023/a:1006773408919.
  • Richards, Robert J.; Darwin and the Emergence of Evolutionary Theories of Mind and Behavior, Chicago: The University of Chicago Press, 1987.


External links

Category:Evolutionary biology

分类: 进化生物学

Category:Selection

类别: 选择


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