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teleology in biology
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{{short description|The use of language of goal-directedness in the context of evolutionary adaptation}}{{good article}}File:Springbok pronk.jpg|thumb|"Behaviour with a purpose": a young springbok stotting.JOURNAL, The functions of stotting in Thomson's gazelles: Some tests of the predictions, Caro, TM, Tim Caro, Animal Behaviour, 1986, 34, 3, 663–684, 10.1016/S0003-3472(86)80052-5, A biologist might argue that this has the function of signalling to predators, helping the springbok to survive and allowing it to reproduce.]]Teleology in biology is the use of the language of goal-directedness in accounts of evolutionary adaptation, which some biologists and philosophers of science find problematic. The term teleonomy has also been proposed. Before Darwin, organisms were seen as existing because God had designed and created them; their features such as eyes were taken by natural theology to have been made to enable them to carry out their functions, such as seeing. Evolutionary biologists often use similar teleological formulations that invoke purpose, but these imply natural selection rather than actual goals, whether conscious or not. Dissenting biologists and religious thinkers held that evolution itself was somehow goal-directed (orthogenesis), and in vitalist versions, driven by a purposeful life force. Since such views are now discredited, with evolution working by natural selection acting on inherited variation, the use of teleology in biology has attracted criticism, and attempts have been made to teach students to avoid teleological language.Nevertheless, biologists still often write about evolution as if organisms had goals, and some philosophers of biology such as Francisco Ayala and biologists such as J. B. S. Haldane consider that teleological language is unavoidable in evolutionary biology.

Context

Teleology

Teleology, from Greek (wikt:τέλος|τέλος), telos "end, purpose"Eric Partridge, Origins: A Short Etymological Dictionary of Modern English, Routledge, 1977, p. 4187. and (wikt:-λογία|-λογία), logia, "a branch of learning", was coined by the philosopher Christian von Wolff in 1728.BOOK, Wolff, Christian von, Christian Wolff (philosopher), 1732, Philosophia Rationalis Sive Logica: Methodo Scientifica Pertractata Et Ad Usum Scientiarum Atque Vitae Aptata,weblink The concept derives from the ancient Greek philosophy of Aristotle, where the final cause (the purpose) of a thing is its function.WEB, Presuppositions of Aristotle's Politics,weblink Stanford Encyclopedia of Philosophy, 28 July 2016, However, Aristotle's biology does not envisage evolution by natural selection.BOOK, Leroi, Armand Marie, Armand Marie Leroi, The Lagoon: How Aristotle Invented Science, Aristotle's Lagoon, Bloomsbury, 2014, 978-1-4088-3622-4, 272–275, Phrases used by biologists like "a function of ... is to ..." or "is designed for" are teleological at least in language. The presence of real or apparent teleology in explanations of natural selection is a controversial aspect of the philosophy of biology, not least for its echoes of natural theology.BOOK, Futuyma, Douglas J., Douglas J. Futuyma, Evolutionary Biology, 1998, Sinauer Associates, 341, 342, 978-0-87893-189-7, File:John Ray from NPG.jpg|thumb|upright|The English natural theologian John Ray, and later William DerhamWilliam Derham

Natural theology

Before Darwin, natural theology both assumed the existence of God and used the appearance of function in nature to argue for the existence of God. The English parson-naturalist John Ray stated that his intention was "to illustrate the glory of God in the knowledge of the works of nature or creation".BOOK, The English Parson-Naturalist, Gracewing, Armstrong, Patrick, 2000, 46, 978-0-85244-516-7, Natural theology presented forms of the teleological argument or argument from design, namely that organs functioned well for their apparent purpose, so they were well-designed, so they must have been designed by a benevolent creator. For example, the eye had the function of seeing, and contained features like the iris and lens that assisted with seeing; therefore, ran the argument, it had been designed for that purpose.JOURNAL, Ayala, Francisco J., Francisco J. Ayala, 2006, Review of 'The Blasphemy of Intelligent Design: Creationism's Trojan Horse. The Wedge of Intelligent Design' by Barbara Forrest; Paul R. Gross, History and Philosophy of the Life Sciences, 28, 3, 409–421, 23334140, WEB, Rosen, Gideon,weblink The Argument from Design, Princeton University, 10 April 2017, BOOK, Paley, William, Paxton, James, Natural Theology: Or, Evidences of the Existence and Attributes of the Deity, Collected from the Appearances of Nature,weblink 1837, Gould, Kendall, and Lincoln, 18–,

Goal-directed evolution

Religious thinkers and biologists have repeatedly supposed that evolution was driven by some kind of life force, a philosophy known as vitalism, and have often supposed that it had some kind of goal or direction (towards which the life force was striving, if they also believed in that), known as orthogenesis or evolutionary progress. Such goal-directedness implies a long-term teleological force; some supporters of orthogenesis considered it to be a spiritual force, while others held that it was purely biological. For example, the Russian embryologist Karl Ernst von Baer believed in a teleological force in nature,Barbieri, Marcello. (2013). Biosemiotics: Information, Codes and Signs in Living Systems. Nova Science Publishers. p. 7. {{ISBN|978-1-60021-612-1}}Jacobsen, Eric Paul. (2005). From Cosmology to Ecology: The Monist World-view in Germany from 1770 to 1930. p. 100. Peter Lang. {{ISBN|978-0-8204-7231-7}} whereas the French spiritualist philosopher Henri Bergson linked orthogenesis with vitalism, arguing for a creative force in evolution known as élan vital in his book Creative Evolution (1907).Bowler, Peter J. (1992). The Eclipse of Darwinism: Anti-Darwinian Evolution Theories in the Decades around 1900. The Johns Hopkins University Press. pp. 116–117. {{ISBN|978-0-8018-4391-4}} The French biophysicist Pierre Lecomte du Noüy and the American botanist Edmund Ware Sinnott developed vitalist evolutionary philosophies known as telefinalism and telism respectively. Their views were heavily criticized as non-scientific;JOURNAL, Koch, Leo Francis, 1957, Vitalistic-Mechanistic Controversy, The Scientific Monthly, 85, 5, 245–255, the palaeontologist George Gaylord Simpson argued that Du Noüy and Sinnott were promoting religious versions of evolution.Simpson, George Gaylord. (1964). Evolutionary Theology: The New Mysticism. In This View of Life: The World of an Evolutionist. Harcourt, Brace & World. pp. 213–233 The Jesuit paleontologist Pierre Teilhard de Chardin argued that evolution was aiming for a supposed spiritual "Omega Point" in what he called "directed additivity".Lane, David H. (1996). The Phenomenon of Teilhard: Prophet for a New Age. Mercer University Press. pp. 60–64. {{ISBN|0-86554-498-0}}De Chardin, Pierre Teilhard. (2003, reprint edition). The Human Phenomenon. Sussex Academic Press. p. 65. {{ISBN|1-902210-30-1}} With the emergence of the modern evolutionary synthesis, in which the genetic mechanisms of evolution were discovered, the hypothesis of orthogenesis was largely abandoned by biologists,Levinton, Jeffrey S. (2001). Genetics, Paleontology, and Macroevolution. Cambridge University Press. pp. 14–16. {{ISBN|0-521-80317-9}}Montgomery, Georgina M; Largent, Mark A. (2015). A Companion to the History of American Science. Wiley. p. 218. {{ISBN|978-1-4051-5625-7}} "With the integration of Mendelian genetics and population genetics into evolutionary theory in the 1930s a new generation of biologists applied mathematical techniques to investigate how changes in the frequency of genes in populations combined with natural selection could produce species change. This demonstrated that Darwinian natural selection was the primary mechanism for evolution and that other models of evolution, such as neo-Lamarckism and orthogenesis, were invalid." especially with Ronald Fisher's argument in his 1930 book The Genetical Theory of Natural Selection.The Structure of Evolutionary Theory by Stephen Jay Gould, Chapter 7, section "Synthesis as Restriction"

Natural selection

Natural selection, introduced in 1859 as the central mechanism{{efn|Darwin also introduced a second mechanism, sexual selection, to explain traits such as the colourful "tail" train of the peacock.BOOK, Darwin, Charles, Charles Darwin, 1859, On the Origin of Species, 1st, 89,weblink }} of evolution by Charles Darwin, is the differential survival and reproduction of individuals due to differences in phenotype.BOOK, Zimmer, Carl, Carl Zimmer, Emlen, Douglas J., Douglas Emlen, 2013, Evolution: Making Sense of Life, 1st, Roberts and Company Publishers, 978-1-936221-17-2, The mechanism directly implies evolution, a change in heritable traits of a population over time.BOOK, Strickberger's Evolution, 4th, Hall, Brian K., Hallgrímsson, Benedikt, Jones and Bartlett, 2008, 4–6,weblink 978-1-4496-4722-3,

Adaptation

File:Sinornithosaurus IVPP V12811.jpg|thumb|upright=1.2|Feathers today serve the function of flight, but they were co-opted rather than adapted for this task, having evolved for an earlier purpose in theropods like Sinornithosaurus millenii, perhaps insulation.]]A trait which persists in a population is often assumed by biologists to have been selected for in the course of evolution, raising the question of how the trait achieves this. Biologists call any such mechanism the function of the trait, using phrases like "A function of stotting by antelopes is to communicate to predators that they have been detected",WEB, Teleological Notions in Biology,weblink Stanford Encyclopedia of Philosophy, 28 July 2016, 18 May 2003, or "The primate hand is designed (by natural selection) for grasping."WEB, Primates - marmosets, monkeys, apes, lemurs, humans,weblink NHPTV, 28 July 2016, The hands and feet of all primates, except for humans, are designed for grasping. Humans have hands designed for grasping, but not feet! Humans have opposable thumbs., An adaptation is an observable structure or other feature of an organism (for example, an enzyme) generated by natural selection to serve its current function. A biologist might propose the hypothesis that feathers are adaptations for bird flight. That would require three things: that the trait of having feathers is heritable; that the trait does serve the function of flight; and that the trait increases the fitness of the organisms that have it. Feathers clearly meet these three conditions in living birds. However, there is also a historical question, namely, did the trait arise at the same time as bird flight? Unfortunately for the hypothesis, this seems not to be so: theropod dinosaurs had feathers, but many of them did not fly.WEB, Understanding Evolution: Qualifying as an adaptation,weblink University of California at Berkeley, 29 July 2016, JOURNAL, Brusatte, Stephen L., Lloyd, Graeme T., Wang, Steve C., Norell, Mark A., 2014, Gradual assembly of avian body plan culminated in rapid rates of evolution across the dinosaur-bird transition, Current Biology, 24, 20, 2386–2392, 10.1016/j.cub.2014.08.034, 25264248, Feathers can be described as an exaptation, having been co-opted for flight but having evolved earlier for another purpose such as insulation. Biologists may describe both the co-option and the earlier adaptation in teleological language.JOURNAL, 4 April 2012, Whitfield, John, Largest feathered dinosaur yet discovered in China, Nature News Blog,weblink 4 April 2012, JOURNAL, Xu, X., Wang, K., Zhang, K., Ma, Q., Xing, L., Sullivan, C., Hu, D., Cheng, S., Wang, S., 2012, A gigantic feathered dinosaur from the Lower Cretaceous of China,weblink Nature, 484, 7392, 92–95, 10.1038/nature10906, etal, 22481363, yes,weblink" title="web.archive.org/web/20120417134949weblink">weblink 2012-04-17,

Status in evolutionary biology

Reasons for discomfort

Apparent teleology is a recurring issue in evolutionary biology,BOOK, Ruse, Michael, Michael Ruse, Travis, J., 2009, Evolution: The First Four Billion Years, The Belknap Press of Harvard University Press, 364, much to the consternation of some writers, and as an explanatory style it remains controversial.BOOK, 2004, Hanke, David, Teleology: The explanation that bedevils biology, 143–155, Explanations: Styles of explanation in science, Cornwell, John, Oxford University Press, 978-0-19-860778-6,weblink There are various reasons for discomfort with teleology among biologists.JOURNAL, Ribeiro, Manuel Gustavo Leitao, etal, On the debate about teleology in biology: the notion of "teleological obstacle", História, Ciências, Saúde-Manguinhos, 2015, 22, 4, 10.1590/S0104-59702015005000003, 25650703, 1321–1333, Firstly, the concept of adaptation is itself controversial, as it can be taken to imply, as the evolutionary biologists Stephen J. Gould and Richard Lewontin argued, that biologists agree with Voltaire's Doctor Pangloss in his 1759 satire Candide that this is "the best of all possible worlds", in other words that every trait is perfectly suited to its functions.JOURNAL, Gould, Stephen J., Stephen J. Gould, Lewontin, Richard, Richard Lewontin, 1979, The spandrels of San Marco and the Panglossian paradigm: a critique of the adaptationist programme, Proceedings of the Royal Society of London B, 205, 205, 581–598, 10.1098/rspb.1979.0086, However, all that evolutionary biology requires is the weaker claim that one trait is at least slightly better in a certain context than another, and hence is selected for.File:Louis George Meistergraveurin 1.jpg|thumb|The Watchmaker analogyWatchmaker analogySecondly, teleology is linked to the pre-Darwinian idea of natural theology, that the natural world gives evidence of the conscious design and beneficent intentions of a creator, as in the writings of John Ray. William Derham continued Ray's tradition with books such as his 1713 Physico-Theology and his 1714 Astro-Theology.BOOK, Weber, A. S., Nineteenth-Century Science: An Anthology, Broadview Press, 2000, 18, They in turn influenced William Paley who wrote a detailed teleological argument for God in 1802, Natural Theology, or Evidences of the Existence and Attributes of the Deity collected from the Appearances of Nature,BOOK, Paley, William, William Paley, Natural Theology, 2006, 1802, Oxford University Press, Oxford,weblink starting with the Watchmaker analogy.JOURNAL, Eddy, Matthew Daniel, Nineteenth Century Natural Theology, The Oxford Handbook of Natural Theology, 2013,weblink Such creationism, along with a vitalist life-force and directed orthogenetic evolution, has been rejected by most biologists.Thirdly, attributing purposes to adaptations risks confusion with popular forms of Lamarckism where animals in particular have been supposed to influence their own evolution through their intentions, though Lamarck himself spoke rather of habits of use, and the belief that his thinking was teleological has been challenged.JOURNAL, Burkhardt, Richard W., Lamarck, Evolution, and the Inheritance of Acquired Characters, Genetics, 194, 4, 2013, 793–805, 10.1534/genetics.113.151852, 23908372, BEDNARCZYK, A, [Jean-Baptiste de Lamarck (1744-1829). A dispute on the mechanism of evolution. On the bicentenary of the publication of Philosophie Zoologique (1809)]., Kwartalnik historii nauki i techniki : Kwartal'nyi zhurnal istorii nauki i tekhniki -, 54, 3-4, 2009, 0023-589X, 20481104, pl, 31–98, Fourthly, the teleological explanation of adaptation is uncomfortable because it seems to require backward causation, in which existing traits are explained by future outcomes; because it seems to attribute the action of a conscious mind when none is assumed to be present in an organism; and because, as a result, adaptation looks impossible to test empirically.A fifth reason concerns students rather than researchers: Gonzalez Galli argues that since people naturally imagine that evolution has a purpose or direction, then the use of teleological language by scientists may act as an obstacle to students when learning about natural selection. Such language, he argues, should be removed to make teaching more effective.JOURNAL, Gonzalez Galli, Leonardo Martin, Meinardi, Elsa N., The Role of Teleological Thinking in Learning the Darwinian Model of Evolution, Evolution: Education and Outreach, March 2011, 4, 1, 145–152, 10.1007/s12052-010-0272-7,

Removable teleological shorthand

Statements which imply that nature has goals, for example where a species is said to do something "in order to" achieve survival, appear teleological, and therefore invalid to evolutionary biologists. It is however usually possible to rewrite such sentences to avoid the apparent teleology. Some biology courses have incorporated exercises requiring students to rephrase such sentences so that they do not read teleologically. Nevertheless, biologists still frequently write in a way which can be read as implying teleology, even though that is not their intention.BOOK, Reiss, John O., 2009, Not by Design: Retiring Darwin's Watchmaker, University of California Press, John Reiss argues that evolutionary biology can be purged of apparent teleology by rejecting the pre-Darwinian watchmaker analogy for natural selection;JOURNAL, Depew, David J., Is Evolutionary Biology Infected with Invalid Teleological Reasoning? A Review of "Not By Design: Retiring Darwin's Watchmaker" by John O. Reiss, Philosophy in Theory and Biology, 2010, 2, 20160629, 10.3998/ptb.6959004.0002.005, other arguments against this analogy have also been promoted by writers such as the evolutionary biologist Richard Dawkins.BOOK, Dawkins, Richard, 1987, The Blind Watchmaker: Why the Evidence of Evolution Reveals a Universe Without Design, W W Norton, Some philosophers of biology such as James G. Lennox have argued that Darwin was a teleologist,JOURNAL, James G. Lennox, Lennox, James G., 1993, Darwin was a Teleologist, Biology and Philosophy, 8, 4, 409–421, 10.1007/bf00857687, while others like Michael Ghiselin described this claim as a myth promoted by misinterpretations of his discussions, and emphasized the distinction between using teleological metaphors and actually being teleological.JOURNAL, 10.1007/BF00850377, Ghiselin, Michael T., 1994, Darwin's language may seem teleological, but his thinking is another matter, Biology and Philosophy, 9, 4, 489–492, Michael Heads, on the other hand, describes a change in Darwin's thinking about evolution that can be traced from the first volume of On the Origin of Species to later volumes. For Heads, Darwin was originally a far more teleological thinker, but over time, "learned to avoid teleology." Heads cites a letter Darwin wrote in 1872, in which he downplayed the role of natural selection as a causal force on its own in explaining biological adaptation, and instead gave more weight to "laws of growth," that operate [without the aid of natural selection].JOURNAL, Heads, Michael, 2009-06, Darwin’s changing views on evolution: from centres of origin and teleology to vicariance and incomplete lineage sorting,weblink Journal of Biogeography, 36, 6, 1018–1026, 10.1111/j.1365-2699.2009.02127.x, 0305-0270, Andrew Askland, from the Sandra Day O'Connor College of Law claims that unlike transhumanism, an ideology that aims to improve the human condition, which he asserts is "wholly teleological", Darwinian evolution is not teleological.JOURNAL,weblink Askland, Andrew, The Misnomer of Transhumanism as Directed Evolution, International Journal of Emerging Technologies and Society, 9, 1, 2011, 71–78, Various commentators view the teleological phrases used in modern evolutionary biology as a type of shorthand for describing any function which offers an evolutionary advantage through natural selection. For example, the zoologist S. H. P. Madrell wrote that "the proper but cumbersome way of describing change by evolutionary adaptation [may be] substituted by shorter overtly teleological statements" for the sake of saving space, but that this "should not be taken to imply that evolution proceeds by anything other than from mutations arising by chance, with those that impart an advantage being retained by natural selection."JOURNAL, Madrell, S.H.P., 1998, Why are there no insects in the open sea?, The Journal of Experimental Biology, 201, 2461–2464,

Irreducible teleology

Other philosophers of biology argue instead that biological teleology is irreducible, and cannot be removed by any simple process of rewording.Francisco Ayala specified three separate situations in which teleological explanations are appropriate. First, if the agent consciously anticipates the goal of their own action; for example the behavior of picking up a pen can be explained by reference to the agent's desire to write. Ayala extends this type of teleological explanation to non-human animals by noting that A deer running away from a mountain lion. . . has at least the appearance of purposeful behavior."JOURNAL, Ayala, Francisco J., 1970-03, Teleological Explanations in Evolutionary Biology,weblink Philosophy of Science, 37, 1, 1–15, 10.1086/288276, 0031-8248, Second, teleological explanations are useful for systems that have a mechanism for self-regulation despite fluctuations in environment; for example, the self-regulation of body temperature in animals. Finally, they are appropriate "in reference to structures anatomically and physiologically designed to perform a certain function. "Ayala, relying on work done by philosopher Ernest Nagel, also rejects the idea that teleological arguments are inadmissible because they cannot be causal. For Nagel, teleological arguments must be consistent because they can always be reformulated as non-teleological arguments. The difference between the two is, for Ayala, merely one of emphasis. Nagel writes that while teleological arguments focus on "the consequences for a given system of a constituent part or process," the equivalent non-teleological arguments focus on ""some of the conditions ... under which the system persists in its characteristic organization and activities." BOOK,weblink The structure of science : Problems in the logic of scientific explanation, Nagel, E., 1961, Harcourt, Brace & World, 874878031, However, Francisco Ayala argued that that teleological statements are more explanatory and cannot be disposed of.BOOK, Ayala, Francisco J., Francisco J. Ayala, 1977, Teleological explanations, Dobzhansky, T., Theodosius Dobzhansky, Evolution, W.H. Freeman, 497–504, Ayala, Francisco (1998). "Teleological explanations in evolutionary biology." Nature's purposes: Analyses of Function and Design in Biology. The MIT Press. Karen Neander similarly argued that the modern concept of biological 'function' depends on natural selection. So, for example, it is not possible to say that anything that simply winks into existence, without going through a process of selection, actually has functions. We decide whether an appendage has a function by analysing the process of selection that led to it. Therefore, Neander argues, any talk of functions must be posterior to natural selection, function must be defined by reference to the history of a species, and teleology cannot be avoided.Neander, Karen (1998). "Functions as Selected Effects: The Conceptual Analyst's Defense," in C. Allen, M. Bekoff & G. Lauder (Eds.), Nature's Purposes: Analyses of Function and Design in Biology (pp. 313–333). The MIT Press. The evolutionary biologist Ernst Mayr likewise stated that "adaptedness ... is an a posteriori result rather than an a priori goal-seeking."JOURNAL, Ernst Mayr, Mayr, Ernst W., 1992, The idea of teleology, Journal of the History of Ideas, 53, 1, 117–135, 10.2307/2709913, 2709913, Angela Breitenbach, looking at the question of teleology in biology from a Kantian perspective, argues that teleology is important as "a heuristic in the search for causal explanations of nature and ... an inevitable analogical perspective on living beings." In her view of Kant, teleology implies something that cannot be explained by science, but only understood through analogy.JOURNAL, Breitenbach, Angela, Teleology in Biology : A Kantian Perspective, 2009, 31–56,weblink Kant Yearbook, 2009, 1, Colin Pittendrigh coined the similar term 'teleonomy' for apparently goal-directed biological phenomena. For Pittendrigh, the notion of 'adaptation' in biology, however it is defined, necessarily "connote that aura of design, purpose, or end-directedness, which has, since the time of Aristotle, seemed to characterize the living thing" JOURNAL, Angel, J. Lawrence, 1961-06, Behavior and evolution. By Anne Roe and George G. Simpson, eds. vii + 557 pp. Yale University Press, New Haven, 1958,weblink American Journal of Physical Anthropology, 19, 2, 218–219, 10.1002/ajpa.1330190215, 0002-9483, This association with Aristotle, however, is problematic, because it meant that the study of adaptation would inevitably be bound up with teleological explanations. Pittendrigh sought to preserve the aspect of design and purpose in biological systems, while denying that this design can be understood as a causal principle. The confusion, he says, would be removed if we described these systems "by some other term, like 'teleonomic,' in order to emphasize that the recognition and description of end-directedness does not carry a commitment to Aristotelian teleology as an efficient causal principle."BOOK, Pittendrigh, Colin S., Colin Pittendrigh, Adaptation, natural selection, and behavior, Behavior and Evolution, Roe, A., Simpson, George Gaylord, George Gaylord Simpson, New Haven, Yale University Press, 1958, 390–416, Ernst Mayr criticised Pittendrigh's confusion of Aristotle's four causes, arguing that evolution only involved the material and formal but not the efficient cause.{{efn|Both Pittendrigh and Mayr endorsed teleology in biology as an inherent part of evolutionary thinking.}} Mayr proposed to use the term only for "systems operating on the basis of a program of coded information."BOOK, Mayr, Ernst, Ernst Mayr, Cause and effect in biology, Lerner, D., Cause and effect, Free Press, New York, 1965, 33–50, William C. Wimsatt affirmed that the teleologicality of the language of biology and other fields derives from the logical structure of their background theories, and not merely from the use of teleological locutions such as "function" and "in order to". He stated that "To replace talk about function by talk about selection [...] is not to eliminate teleology but to rephrase it". However, Wimsatt argues that this thought does not mean an appeal to backwards causation, vitalism, entelechy, or anti-reductionist sentiments.JOURNAL, Wimsatt, W., William C. Wimsatt, 1972, Teleology and the logical structure of function statements, Studies in the History and Philosophy of Science, 3, 1–80,weblink 10.1016/0039-3681(72)90014-3, The biologist J. B. S. Haldane observed that "Teleology is like a mistress to a biologist: he cannot live without her but he's unwilling to be seen with her in public."BOOK, Hull, D., Philosophy of Biological Science, Prentice-Hall, 1973, Mayr, Ernst (1974) Boston Studies in the Philosophy of Science, Volume XIV, pages 91–117.

See also

Notes

{{notelist}}

References

{{reflist|30em}}{{Philosophy of biology}}

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