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Structures and functions in the evolution of morality

       Emanuele Serrelli
       University of Milano Bicocca, Italy




                                              Erice, Sicily (IT)
                                             17-22 June 2012
Evolutionary explanations
Evolutionary explanations




  “Panglossian”
         or
  “just-so story”
    explanation

  (Stephen Jay Gould &
   Richard C. Lewontin,
“spandrels” paper, 1979)
Evolutionary explanations




  “Panglossian”
         or
  “just-so story”
    explanation

  (Stephen Jay Gould &     A trait T
   Richard C. Lewontin,
“spandrels” paper, 1979)
Evolutionary explanations




  “Panglossian”
         or
  “just-so story”
    explanation

  (Stephen Jay Gould &     A trait T   A function
   Richard C. Lewontin,
“spandrels” paper, 1979)
Evolutionary explanations




  “Panglossian”
         or
  “just-so story”
    explanation

  (Stephen Jay Gould &     A trait T   A function
   Richard C. Lewontin,
“spandrels” paper, 1979)
Evolutionary explanations




  “Panglossian”
         or
  “just-so story”
    explanation

  (Stephen Jay Gould &      A trait T   A function
   Richard C. Lewontin,
“spandrels” paper, 1979)
                           Morality?
Evolutionary explanations




  “Panglossian”
         or
  “just-so story”
    explanation

  (Stephen Jay Gould &      A trait T                      A function
   Richard C. Lewontin,
“spandrels” paper, 1979)
                           Morality?
                             “Why do you evolutionists always try to
                            assess the value of anything even before
                                      knowing what it is?”
                                    (Francis Crick, in Gould 1991)
Moral traits - what is morality?
Moral traits - what is morality?

• Joyce: PRECISIFICATION OF MORAL SENSE
Moral traits - what is morality?

• Joyce: PRECISIFICATION OF MORAL SENSE
• Kitcher: “ETHICS” OR “MORALITY”? “JUST A TECHNICAL
  PHILOSOPHICAL PROBLEM”?
Moral traits - what is morality?

• Joyce: PRECISIFICATION OF MORAL SENSE
• Kitcher: “ETHICS” OR “MORALITY”? “JUST A TECHNICAL
  PHILOSOPHICAL PROBLEM”?
      lack of precisifications in the “evolution of morality” field?
Moral traits - what is morality?

• Joyce: PRECISIFICATION OF MORAL SENSE
• Kitcher: “ETHICS” OR “MORALITY”? “JUST A TECHNICAL
  PHILOSOPHICAL PROBLEM”?
      lack of precisifications in the “evolution of morality” field?
                still trying to get a grip?
Moral traits - what is morality?

• Joyce: PRECISIFICATION OF MORAL SENSE
• Kitcher: “ETHICS” OR “MORALITY”? “JUST A TECHNICAL
  PHILOSOPHICAL PROBLEM”?
      lack of precisifications in the “evolution of morality” field?
                still trying to get a grip?
                an explanatory strategy?
Moral traits - what is morality?

• Joyce: PRECISIFICATION OF MORAL SENSE
• Kitcher: “ETHICS” OR “MORALITY”? “JUST A TECHNICAL
  PHILOSOPHICAL PROBLEM”?
      lack of precisifications in the “evolution of morality” field?
                still trying to get a grip?
                an explanatory strategy?
• Joyce: INDETERMINATE CONCEPTS
Moral traits - what is morality?

• Joyce: PRECISIFICATION OF MORAL SENSE
• Kitcher: “ETHICS” OR “MORALITY”? “JUST A TECHNICAL
  PHILOSOPHICAL PROBLEM”?
      lack of precisifications in the “evolution of morality” field?
                still trying to get a grip?
                an explanatory strategy?
• Joyce: INDETERMINATE CONCEPTS
• Churchland: RADIAL STRUCTURE AND FUZZY BOUNDARIES OF
  CONCEPTS
Moral traits - what is morality?

• Joyce: PRECISIFICATION OF MORAL SENSE
• Kitcher: “ETHICS” OR “MORALITY”? “JUST A TECHNICAL
  PHILOSOPHICAL PROBLEM”?
      lack of precisifications in the “evolution of morality” field?
                still trying to get a grip?
                an explanatory strategy?
• Joyce: INDETERMINATE CONCEPTS
• Churchland: RADIAL STRUCTURE AND FUZZY BOUNDARIES OF
  CONCEPTS
      BUT scientific need for concepts
Moral traits - what is morality?

• Joyce: PRECISIFICATION OF MORAL SENSE
• Kitcher: “ETHICS” OR “MORALITY”? “JUST A TECHNICAL
  PHILOSOPHICAL PROBLEM”?
      lack of precisifications in the “evolution of morality” field?
                still trying to get a grip?
                an explanatory strategy?
• Joyce: INDETERMINATE CONCEPTS
• Churchland: RADIAL STRUCTURE AND FUZZY BOUNDARIES OF
  CONCEPTS
      BUT scientific need for concepts
                at least, pragmatically, contextualizing results in certain premises
Moral traits - what is morality?

 EVOLUTIONARY
 CRITERIA
 for trait recognition:

 • HERITABILITY
 • INTER-INDIVIDUAL
   VARIATION

 • FITNESS FUNCTION
Moral traits - what is morality?

                          The evolutionary relevance
 EVOLUTIONARY
 CRITERIA                 of a trait depends
 for trait recognition:   on its inheritance pattern
 • HERITABILITY
 • INTER-INDIVIDUAL
   VARIATION

 • FITNESS FUNCTION
Moral traits - what is morality?

                                                    The evolutionary relevance
 EVOLUTIONARY
 CRITERIA                                           of a trait depends
 for trait recognition:                             on its inheritance pattern
 • HERITABILITY
 • INTER-INDIVIDUAL
     VARIATION

 • FITNESS FUNCTION




Picture credits: "Autosomal recessive disorders" http://www.brusselsgenetics.be/p_453.htm // Wright (1931)
Moral traits - what is morality?

                                                    The evolutionary relevance
 EVOLUTIONARY
 CRITERIA                                           of a trait depends
 for trait recognition:                             on its inheritance pattern
 • HERITABILITY
 • INTER-INDIVIDUAL
     VARIATION

 • FITNESS FUNCTION




Picture credits: "Autosomal recessive disorders" http://www.brusselsgenetics.be/p_453.htm // Wright (1931)
Moral traits - what is morality?

                                                    The evolutionary relevance
 EVOLUTIONARY
 CRITERIA                                           of a trait depends
 for trait recognition:                             on its inheritance pattern
 • HERITABILITY
 • INTER-INDIVIDUAL
     VARIATION

 • FITNESS FUNCTION


                                                                        Genes are all which is inheritable...
                                                                            (The Modern Synthesis, 1920s-40s)

Picture credits: "Autosomal recessive disorders" http://www.brusselsgenetics.be/p_453.htm // Wright (1931)
Moral traits - what is morality?

 EVOLUTIONARY
 CRITERIA
 for trait recognition:

 • HERITABILITY
 • INTER-INDIVIDUAL
   VARIATION

 • FITNESS FUNCTION




                          http://blogs.scientificamerican.com/science-sushi/2011/10/05/a-moral-gene/
Moral traits - what is morality?

                          Genes do not necessarily
 EVOLUTIONARY
 CRITERIA                 lose their importance
 for trait recognition:
                          (e.g., Churchland 2011)
 • HERITABILITY
                          (e.g. kin selection
                          models of altruism evolution)
 • INTER-INDIVIDUAL
   VARIATION

 • FITNESS FUNCTION
Moral traits - what is morality?

                                                   Genes do not necessarily
 EVOLUTIONARY
 CRITERIA                                          lose their importance
 for trait recognition:
                                                   (e.g., Churchland 2011)
 • HERITABILITY
                                                   (e.g. kin selection
                                                   models of altruism evolution)
 • INTER-INDIVIDUAL
   VARIATION

 • FITNESS FUNCTION                                but heritability has a
                                                   larger scope today:
                                                   the heritability pattern of a
                                                   dynamic phenotype

   Picture credit: Fusco (2011), in Evoluzione Modelli e Processi, ed. by Ferraguti
   & Castellacci, Pearson, Italy.
Moral traits - what is morality?

 EVOLUTIONARY
 CRITERIA
 for trait recognition:

 • HERITABILITY
 • INTER-INDIVIDUAL
   VARIATION

 • FITNESS FUNCTION




                          Picture credits: Flack & de Waal (2000), Journal of Consciousness Studies, p. 6.
Moral traits - what is morality?

 EVOLUTIONARY
 CRITERIA
 for trait recognition:

 • HERITABILITY
 • INTER-INDIVIDUAL
   VARIATION
      “Twin and family studies
 • FITNESS FUNCTION of
       suggest a heritability
 psychopathy in the range of about
 70%; childhood conditions such as
abuse and neglect may contribute to
those who are genetically disposed”
        Churchland 2011, p. 41)

                                      Picture credits: Flack & de Waal (2000), Journal of Consciousness Studies, p. 6.
Moral traits - what is morality?

                          We are studying evolutionarily
 EVOLUTIONARY
 CRITERIA                 relevant traits only if we can
 for trait recognition:
                          assess their reliable
 • HERITABILITY           reappearance
 • INTER-INDIVIDUAL
   VARIATION

 • FITNESS FUNCTION
Moral traits - what is morality?

                          We are studying evolutionarily
 EVOLUTIONARY
 CRITERIA                 relevant traits only if we can
 for trait recognition:
                          assess their reliable
 • HERITABILITY           reappearance
 • INTER-INDIVIDUAL
   VARIATION              “Biometrical population genetics”
                          (cf. Serrelli 2010, Lewontin 1980)

 • FITNESS FUNCTION
Moral traits - what is morality?

                          We are studying evolutionarily
 EVOLUTIONARY
 CRITERIA                 relevant traits only if we can
 for trait recognition:
                          assess their reliable
 • HERITABILITY           reappearance
 • INTER-INDIVIDUAL
   VARIATION              “Biometrical population genetics”
                          (cf. Serrelli 2010, Lewontin 1980)

 • FITNESS FUNCTION
                          QTL - Quantitative Traits Locis
                          (cf. Hartl & Clark 2007, chp. 8)
Moral traits - what is morality?

                          We are studying evolutionarily
 EVOLUTIONARY
 CRITERIA                 relevant traits only if we can
 for trait recognition:
                          assess their reliable
 • HERITABILITY           reappearance
 • INTER-INDIVIDUAL
   VARIATION              “Biometrical population genetics”
                          (cf. Serrelli 2010, Lewontin 1980)

 • FITNESS FUNCTION
                          QTL - Quantitative Traits Locis
                          (cf. Hartl & Clark 2007, chp. 8)


                          Trait = inheritable pattern
Moral traits - what is morality?

                                                       Trait = inheritable pattern
 EVOLUTIONARY
 CRITERIA
 for trait recognition:

 • HERITABILITY
 • INTER-INDIVIDUAL
     VARIATION

 • FITNESS FUNCTION

Picture credits:
chimps: http://angelorganic.org/darwin-mis-
interpreted/
birds: http://news.sciencemag.org/sciencenow/
2010/08/cheatin-hearts-get-stuck-with-th.html
ants: http://www.ridelust.com/traffic-congestion-and-
social-insects-we-can-learn-how-to-drive-from-ants/
V. Bulatov (2011), http://bulatov.org/math/1101/
webtalk.html
Moral traits - what is morality?

                                                       Trait = inheritable pattern
 EVOLUTIONARY
 CRITERIA
 for trait recognition:                                Need for models
 • HERITABILITY
 • INTER-INDIVIDUAL
     VARIATION

 • FITNESS FUNCTION

Picture credits:
chimps: http://angelorganic.org/darwin-mis-
interpreted/
birds: http://news.sciencemag.org/sciencenow/
2010/08/cheatin-hearts-get-stuck-with-th.html
ants: http://www.ridelust.com/traffic-congestion-and-
social-insects-we-can-learn-how-to-drive-from-ants/
V. Bulatov (2011), http://bulatov.org/math/1101/
webtalk.html
Moral traits - what is morality?

                                                       Trait = inheritable pattern
 EVOLUTIONARY
 CRITERIA
 for trait recognition:                                Need for models
 • HERITABILITY
 • INTER-INDIVIDUAL                                    Homology:
     VARIATION
                                                       pattern shows up
 • FITNESS FUNCTION                                    in different taxa
Picture credits:
chimps: http://angelorganic.org/darwin-mis-
interpreted/
birds: http://news.sciencemag.org/sciencenow/
2010/08/cheatin-hearts-get-stuck-with-th.html
ants: http://www.ridelust.com/traffic-congestion-and-
social-insects-we-can-learn-how-to-drive-from-ants/
V. Bulatov (2011), http://bulatov.org/math/1101/
webtalk.html
Moral traits - what is morality?

                                                       Trait = inheritable pattern
 EVOLUTIONARY
 CRITERIA
 for trait recognition:                                Need for models
 • HERITABILITY
 • INTER-INDIVIDUAL                                    Homology:
     VARIATION
                                                       pattern shows up
 • FITNESS FUNCTION                                    in different taxa
Picture credits:
chimps: http://angelorganic.org/darwin-mis-


                                                       Trait of...
interpreted/
birds: http://news.sciencemag.org/sciencenow/
2010/08/cheatin-hearts-get-stuck-with-th.html
ants: http://www.ridelust.com/traffic-congestion-and-
social-insects-we-can-learn-how-to-drive-from-ants/
V. Bulatov (2011), http://bulatov.org/math/1101/
webtalk.html
                                                       a relevant unit
The function of morality




 In Neodarwinism not
    all effects are
functions. Function is
   bound to Natural
        Selection              A trait T                     function?
                                                                  flight?
   (cf. Williams 1966)
                                                      effect
                         ...      effect        e.g. thermoregulation?
                                 e.g. mating?
The function of morality

                         (1) ΔqT = [sqT(1−qT)]/
                         [1−s(1−qT)]
                         (cf. Wright 1931)

 In Neodarwinism not
    all effects are
functions. Function is
   bound to Natural
        Selection

   (cf. Williams 1966)
The function of morality

                         (1) ΔqT = [sqT(1−qT)]/
                         [1−s(1−qT)]
                         (cf. Wright 1931)

 In Neodarwinism not
    all effects are      Need to specify a “problem
functions. Function is
   bound to Natural      background” (cf. Kitcher 2011)
        Selection

   (cf. Williams 1966)
The function of morality

                         (1) ΔqT = [sqT(1−qT)]/
                         [1−s(1−qT)]
                         (cf. Wright 1931)

 In Neodarwinism not
    all effects are      Need to specify a “problem
functions. Function is
   bound to Natural      background” (cf. Kitcher 2011)
        Selection

   (cf. Williams 1966)   (2) w = f (A, B, … T)
The function of morality

                         (1) ΔqT = [sqT(1−qT)]/
                         [1−s(1−qT)]
                         (cf. Wright 1931)

 In Neodarwinism not
    all effects are      Need to specify a “problem
functions. Function is
   bound to Natural      background” (cf. Kitcher 2011)
        Selection

   (cf. Williams 1966)   (2) w = f (A, B, … T)
The function of morality

                         (1) ΔqT = [sqT(1−qT)]/
                         [1−s(1−qT)]
                         (cf. Wright 1931)

 In Neodarwinism not
    all effects are      Need to specify a “problem
functions. Function is
   bound to Natural      background” (cf. Kitcher 2011)
        Selection

   (cf. Williams 1966)   (2) w = f (A, B, … T)
The function of morality

                         (1) ΔqT = [sqT(1−qT)]/
                         [1−s(1−qT)]
                         (cf. Wright 1931)

 In Neodarwinism not
    all effects are      Need to specify a “problem
functions. Function is
   bound to Natural      background” (cf. Kitcher 2011)
        Selection

   (cf. Williams 1966)   (3) T = f (F, r)
The function of morality

                         (1) ΔqT = [sqT(1−qT)]/
                         [1−s(1−qT)]
                         (cf. Wright 1931)

 In Neodarwinism not
    all effects are      Need to specify a “problem
functions. Function is
   bound to Natural      background” (cf. Kitcher 2011)
        Selection

   (cf. Williams 1966)   (3) T = f (F, r)
The function of morality

                         (1) ΔqT = [sqT(1−qT)]/
                         [1−s(1−qT)]
                         (cf. Wright 1931)

 In Neodarwinism not
    all effects are      Need to specify a “problem
functions. Function is
   bound to Natural      background” (cf. Kitcher 2011)
        Selection

                         (3) T = f (F, r)        Functioning
   (cf. Williams 1966)
                                                 and roles are
                                                 not causal
                                                 upon traits
                                                 and structures
The function of morality


   (3) T = f (F, r)


      “...the palette of
 neurochemicals... affecting
  neurons and muscles is
   substantially the same
   across vertebrates and
      invertebrates...”
     (Churchland 2011, p. 45)

                                Picture credit: McPartland et al. (2006), Gene 370:64-74. http://
                                www.sciencedirect.com/science/article/pii/S0378111905007067
The function of morality


  (3) T = f (F, r)




  oxytocin is found in all
       vertebrates
    (Churchland 2011, p. 45)
                               Picture credit: McPartland et al. (2006), Gene 370:64-74. http://
                               www.sciencedirect.com/science/article/pii/S0378111905007067
The function of morality


  (3) T = f (F, r)




“Oxytocin ... is at the hub of
  the intricate network of
mammalian adaptations for
     caring for others”
     (Churchland 2011, p. 14)
                                 Picture credit: McPartland et al. (2006), Gene 370:64-74. http://
                                 www.sciencedirect.com/science/article/pii/S0378111905007067
The function of morality


  (3) T = f (F, r)




“...modest modifications in
exiting neural structures ...
can lead to new outcomes”
    (Churchland 2011, p. 14)
                                Picture credit: McPartland et al. (2006), Gene 370:64-74. http://
                                www.sciencedirect.com/science/article/pii/S0378111905007067
The function of morality


  (3) T = f (F, r)


 “a large... range of social
  patterns found among
mammals..., but underlying
them are probably different
arrangements of receptors
  for oxytocin and other
       hormones and
      neurochemicals”
    (Churchland 2011, p. 14)   Picture credit: McPartland et al. (2006), Gene 370:64-74. http://
                               www.sciencedirect.com/science/article/pii/S0378111905007067
The function of morality


  (3) ∆T = f (∆F, ∆r)


          “Biological evolution does not achieve adaptations by
      designing a whole new mechanism from scratch, but modifies
       what is already in place, little bit by little bit. Social emotions,
          values, and behavior are not the result of a wholly new
           engineering plan, but rather an adaptation of existing
      arrangements and mechanisms that are intimately linked with
      the self-preserving circuitry for fighting, freezing, and flight, on
           the one hand, and for rest and digest, on the other”
                              (Churchland 2011, p. 46)
Evolutionary explanations of morality

• Explanatory diversity and integration
Evolutionary explanations of morality

• Explanatory diversity and integration
• Traits/structures and functions are needed,
  but don’t come for free
Evolutionary explanations of morality

• Explanatory diversity and integration
• Traits/structures and functions are needed,
  but don’t come for free
• Traits as inheritable patterns
Evolutionary explanations of morality

• Explanatory diversity and integration
• Traits/structures and functions are needed,
  but don’t come for free
• Traits as inheritable patterns
         models
Evolutionary explanations of morality

• Explanatory diversity and integration
• Traits/structures and functions are needed,
  but don’t come for free
• Traits as inheritable patterns
         models
         homologies avoiding circularity
Evolutionary explanations of morality

• Explanatory diversity and integration
• Traits/structures and functions are needed,
  but don’t come for free
• Traits as inheritable patterns
         models
         homologies avoiding circularity
         units of evolution
Evolutionary explanations of morality

• Explanatory diversity and integration
• Traits/structures and functions are needed,
  but don’t come for free
• Traits as inheritable patterns
         models
         homologies avoiding circularity
         units of evolution
Evolutionary explanations of morality

• Explanatory diversity and integration
• Traits/structures and functions are needed,
  but don’t come for free
• Traits as inheritable patterns
         models
         homologies avoiding circularity
         units of evolution

• Functions
      conceptual problems (e.g. Cummins 1975, Wright 1973, Millikan
      1984, 1989, Griffiths 1993, Godfrey-Smith 1994, cf. Casebeer
      2003)




                  Picture credit: http://www.azoft.com/spotlight/2011/05/05/15-bizarre-dead-tech-products.html
Evolutionary explanations of morality

• Explanatory diversity and integration
• Traits/structures and functions are needed,
  but don’t come for free
• Traits as inheritable patterns
          models
          homologies avoiding circularity
          units of evolution

• Functions
      conceptual problems (e.g. Cummins 1975, Wright 1973, Millikan
      1984, 1989, Griffiths 1993, Godfrey-Smith 1994, cf. Casebeer
      2003)
      sights for identifying traits




                   Picture credit: http://www.azoft.com/spotlight/2011/05/05/15-bizarre-dead-tech-products.html
Evolutionary explanations of morality

• Explanatory diversity and integration
• Traits/structures and functions are needed,
  but don’t come for free
• Traits as inheritable patterns
          models
          homologies avoiding circularity
          units of evolution

• Functions
      conceptual problems (e.g. Cummins 1975, Wright 1973, Millikan
      1984, 1989, Griffiths 1993, Godfrey-Smith 1994, cf. Casebeer
      2003)
      sights for identifying traits
      crucial for Darwinian explanations




                   Picture credit: http://www.azoft.com/spotlight/2011/05/05/15-bizarre-dead-tech-products.html
Evolutionary explanations of morality

• Explanatory diversity and integration
• Traits/structures and functions are needed,
  but don’t come for free
• Traits as inheritable patterns
          models
          homologies avoiding circularity
          units of evolution

• Functions
      conceptual problems (e.g. Cummins 1975, Wright 1973, Millikan
      1984, 1989, Griffiths 1993, Godfrey-Smith 1994, cf. Casebeer
      2003)
      sights for identifying traits
      crucial for Darwinian explanations
      functions / effects


                   Picture credit: http://www.azoft.com/spotlight/2011/05/05/15-bizarre-dead-tech-products.html
THANK YOU!

Picture credits: http://www.azoft.com/spotlight/2011/05/05/15-bizarre-dead-tech-products.html
REFERENCES:
• Boehm C (2000), Conflict and the evolution of social control. Journal of Consciousness Studies 7(1-2), “Evolutionary Origins of Morality. Cross-Disciplinary
   Perspectives”, pp. 79-101.
• Churchland PS (2011). Braintrust: What Neuroscience Tells Us About Morality. Princeton: Princeton University Press.
• Cummins R (1975). Functional analysis. Journal of Philosophy 72:741-765.
• Downes SM (2010). Heritability. The Stanford Encyclopedia of Philosophy (Fall 2010 Edition), Edward N. Zalta (ed.), URL = http://plato.stanford.edu/archives/fall2010/
   entries/heredity/.
• Eisenberg N, Strayer J (1987) Empathy and its development. Cambridge UP, Cambridge.
• Ferraguti M, Castellacci C, eds. (2011), Evoluzione: modelli e processi. Pearson, Milano-Torino.
• Flack JC, de Waal FBM (2000), ‘Any animal whatever’: Darwinian building blocks of morality in monkeys and apes. Journal of Consciousness Studies 7(1-2),
   “Evolutionary Origins of Morality. Cross-Disciplinary Perspectives”, pp. 1-29.
• Godfrey-Smith P (1994). A modern history theory of functions. Reprinted in: Allen C, Bekoff M, Lauder G, eds. Nature’s Purposes - Analyses of Function and Design in
   Biology. Cambridge, MA: MIT Press, 1998, pp. 453-477.
• Gould SJ (1991), “Capezzoli maschili e glande clitorideo”, in trad. it. Bravo Brontosauro, Milano, Feltrinelli Universale Economica, 2002, pp. 125‐139.
• Gould SJ, Lewontin RC (1979), The spandrels of San Marco and the Panglossian paradigm: a critique of the adaptationist programme, Proc. R. Soc. Lond. B, 205,
   581-598. Trad. it. di M. Ferraguti, I pennacchi di San Marco e il paradigma di Pangloss: critica al programma adattazionista. Piccola biblioteca on line, Einaudi, Torino,
   2001. http://www.einaudi.it/einaudi/ita/pdf/gould-lewontin.pdf
• Griffiths PE (1993). Functional analysis and proper functions. British Journal of the Philosophy of Science 44:409-422.
• Griffiths PE, Brigandt I, eds. (2007). “The Importance of Homology for Biology and Philosophy”, Biology and Philosophy 22.
• Hartl DL, Clark AG (2007). Principles of Population Genetics, Fourth ed. Sunderland, Mass.: Sinauer Associates.
• Kitcher P (2011). The Ethical Project, Harvard University Press, Cambridge, Mass. & London, UK.
• Jablonka E, Lamb MJ (2005). Evolution in Four Dimensions. Cambridge (MA): The MIT Press, revised in 2010.
• Millikan R (1984). Language, Thought and Other Biological Categories. Cambridge, Mass: MIT Press.
• Millikan R (1989). In defence of proper functions. Philosophy of Science 56:288-302.
• Müller G.B., 2010. Epigenetic innovation. In Pigliucci M, Müller GB, eds. Evolution The Extended Synthesis. Cambridge-London: MIT Press, pp. 307-332.
• Odling-Smee J, Laland K, Feldman MW (2003). Niche Construction. Princeton: Princeton University Press.
• McPartland JM, Matias I, Di Marzo V, GlassM (2006). Evolutionary origins of the endocannabinoid system, Gene 370:64-74, 10.1016/j.gene.2005.11.004.
•   Pievani T, Serrelli E (2011). Exaptation in human evolution: how to test adaptive vs exaptive evolutionary hypotheses. Journal of Anthropological Sciences 89, pp. 9-23.
    [DOI 10.4436/jass.89015]
•   Serrelli E (2011). The structure of population genetics. Poster at the meeting of the International Society for History, Philosophy, and Social Studies of Biology (ISHPSSB),
    Salt Lake City (Utah, USA) July 10th-16th 2011. [http://hdl.handle.net/10281/28245]
•   Sober E (1984). The Nature of Selection. Evolutionary Theory in Philosophical Focus. University of Chicago Press, Chicago.
•   Sober E, Wilson DS (1998). Unto Others: the Evolution and Psychology of Unselfish Behavior. Harvard University Press, Cambridge (MA).
•   Williams GC (1966). Adaptation and Natural Selection. Princeton: Princeton University Press.
•   Wright L (1973). Functions. Philosophical Review 82:139-168.
•   Wright S (1931). Evolution in mendelian populations. Genetics, 16 (97): 97–159.

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Emanuele Serrelli - Structures and functions in the evolution of morality

  • 1. Structures and functions in the evolution of morality Emanuele Serrelli University of Milano Bicocca, Italy Erice, Sicily (IT) 17-22 June 2012
  • 3. Evolutionary explanations “Panglossian” or “just-so story” explanation (Stephen Jay Gould & Richard C. Lewontin, “spandrels” paper, 1979)
  • 4. Evolutionary explanations “Panglossian” or “just-so story” explanation (Stephen Jay Gould & A trait T Richard C. Lewontin, “spandrels” paper, 1979)
  • 5. Evolutionary explanations “Panglossian” or “just-so story” explanation (Stephen Jay Gould & A trait T A function Richard C. Lewontin, “spandrels” paper, 1979)
  • 6. Evolutionary explanations “Panglossian” or “just-so story” explanation (Stephen Jay Gould & A trait T A function Richard C. Lewontin, “spandrels” paper, 1979)
  • 7. Evolutionary explanations “Panglossian” or “just-so story” explanation (Stephen Jay Gould & A trait T A function Richard C. Lewontin, “spandrels” paper, 1979) Morality?
  • 8. Evolutionary explanations “Panglossian” or “just-so story” explanation (Stephen Jay Gould & A trait T A function Richard C. Lewontin, “spandrels” paper, 1979) Morality? “Why do you evolutionists always try to assess the value of anything even before knowing what it is?” (Francis Crick, in Gould 1991)
  • 9. Moral traits - what is morality?
  • 10. Moral traits - what is morality? • Joyce: PRECISIFICATION OF MORAL SENSE
  • 11. Moral traits - what is morality? • Joyce: PRECISIFICATION OF MORAL SENSE • Kitcher: “ETHICS” OR “MORALITY”? “JUST A TECHNICAL PHILOSOPHICAL PROBLEM”?
  • 12. Moral traits - what is morality? • Joyce: PRECISIFICATION OF MORAL SENSE • Kitcher: “ETHICS” OR “MORALITY”? “JUST A TECHNICAL PHILOSOPHICAL PROBLEM”? lack of precisifications in the “evolution of morality” field?
  • 13. Moral traits - what is morality? • Joyce: PRECISIFICATION OF MORAL SENSE • Kitcher: “ETHICS” OR “MORALITY”? “JUST A TECHNICAL PHILOSOPHICAL PROBLEM”? lack of precisifications in the “evolution of morality” field? still trying to get a grip?
  • 14. Moral traits - what is morality? • Joyce: PRECISIFICATION OF MORAL SENSE • Kitcher: “ETHICS” OR “MORALITY”? “JUST A TECHNICAL PHILOSOPHICAL PROBLEM”? lack of precisifications in the “evolution of morality” field? still trying to get a grip? an explanatory strategy?
  • 15. Moral traits - what is morality? • Joyce: PRECISIFICATION OF MORAL SENSE • Kitcher: “ETHICS” OR “MORALITY”? “JUST A TECHNICAL PHILOSOPHICAL PROBLEM”? lack of precisifications in the “evolution of morality” field? still trying to get a grip? an explanatory strategy? • Joyce: INDETERMINATE CONCEPTS
  • 16. Moral traits - what is morality? • Joyce: PRECISIFICATION OF MORAL SENSE • Kitcher: “ETHICS” OR “MORALITY”? “JUST A TECHNICAL PHILOSOPHICAL PROBLEM”? lack of precisifications in the “evolution of morality” field? still trying to get a grip? an explanatory strategy? • Joyce: INDETERMINATE CONCEPTS • Churchland: RADIAL STRUCTURE AND FUZZY BOUNDARIES OF CONCEPTS
  • 17. Moral traits - what is morality? • Joyce: PRECISIFICATION OF MORAL SENSE • Kitcher: “ETHICS” OR “MORALITY”? “JUST A TECHNICAL PHILOSOPHICAL PROBLEM”? lack of precisifications in the “evolution of morality” field? still trying to get a grip? an explanatory strategy? • Joyce: INDETERMINATE CONCEPTS • Churchland: RADIAL STRUCTURE AND FUZZY BOUNDARIES OF CONCEPTS BUT scientific need for concepts
  • 18. Moral traits - what is morality? • Joyce: PRECISIFICATION OF MORAL SENSE • Kitcher: “ETHICS” OR “MORALITY”? “JUST A TECHNICAL PHILOSOPHICAL PROBLEM”? lack of precisifications in the “evolution of morality” field? still trying to get a grip? an explanatory strategy? • Joyce: INDETERMINATE CONCEPTS • Churchland: RADIAL STRUCTURE AND FUZZY BOUNDARIES OF CONCEPTS BUT scientific need for concepts at least, pragmatically, contextualizing results in certain premises
  • 19. Moral traits - what is morality? EVOLUTIONARY CRITERIA for trait recognition: • HERITABILITY • INTER-INDIVIDUAL VARIATION • FITNESS FUNCTION
  • 20. Moral traits - what is morality? The evolutionary relevance EVOLUTIONARY CRITERIA of a trait depends for trait recognition: on its inheritance pattern • HERITABILITY • INTER-INDIVIDUAL VARIATION • FITNESS FUNCTION
  • 21. Moral traits - what is morality? The evolutionary relevance EVOLUTIONARY CRITERIA of a trait depends for trait recognition: on its inheritance pattern • HERITABILITY • INTER-INDIVIDUAL VARIATION • FITNESS FUNCTION Picture credits: "Autosomal recessive disorders" http://www.brusselsgenetics.be/p_453.htm // Wright (1931)
  • 22. Moral traits - what is morality? The evolutionary relevance EVOLUTIONARY CRITERIA of a trait depends for trait recognition: on its inheritance pattern • HERITABILITY • INTER-INDIVIDUAL VARIATION • FITNESS FUNCTION Picture credits: "Autosomal recessive disorders" http://www.brusselsgenetics.be/p_453.htm // Wright (1931)
  • 23. Moral traits - what is morality? The evolutionary relevance EVOLUTIONARY CRITERIA of a trait depends for trait recognition: on its inheritance pattern • HERITABILITY • INTER-INDIVIDUAL VARIATION • FITNESS FUNCTION Genes are all which is inheritable... (The Modern Synthesis, 1920s-40s) Picture credits: "Autosomal recessive disorders" http://www.brusselsgenetics.be/p_453.htm // Wright (1931)
  • 24. Moral traits - what is morality? EVOLUTIONARY CRITERIA for trait recognition: • HERITABILITY • INTER-INDIVIDUAL VARIATION • FITNESS FUNCTION http://blogs.scientificamerican.com/science-sushi/2011/10/05/a-moral-gene/
  • 25. Moral traits - what is morality? Genes do not necessarily EVOLUTIONARY CRITERIA lose their importance for trait recognition: (e.g., Churchland 2011) • HERITABILITY (e.g. kin selection models of altruism evolution) • INTER-INDIVIDUAL VARIATION • FITNESS FUNCTION
  • 26. Moral traits - what is morality? Genes do not necessarily EVOLUTIONARY CRITERIA lose their importance for trait recognition: (e.g., Churchland 2011) • HERITABILITY (e.g. kin selection models of altruism evolution) • INTER-INDIVIDUAL VARIATION • FITNESS FUNCTION but heritability has a larger scope today: the heritability pattern of a dynamic phenotype Picture credit: Fusco (2011), in Evoluzione Modelli e Processi, ed. by Ferraguti & Castellacci, Pearson, Italy.
  • 27. Moral traits - what is morality? EVOLUTIONARY CRITERIA for trait recognition: • HERITABILITY • INTER-INDIVIDUAL VARIATION • FITNESS FUNCTION Picture credits: Flack & de Waal (2000), Journal of Consciousness Studies, p. 6.
  • 28. Moral traits - what is morality? EVOLUTIONARY CRITERIA for trait recognition: • HERITABILITY • INTER-INDIVIDUAL VARIATION “Twin and family studies • FITNESS FUNCTION of suggest a heritability psychopathy in the range of about 70%; childhood conditions such as abuse and neglect may contribute to those who are genetically disposed” Churchland 2011, p. 41) Picture credits: Flack & de Waal (2000), Journal of Consciousness Studies, p. 6.
  • 29. Moral traits - what is morality? We are studying evolutionarily EVOLUTIONARY CRITERIA relevant traits only if we can for trait recognition: assess their reliable • HERITABILITY reappearance • INTER-INDIVIDUAL VARIATION • FITNESS FUNCTION
  • 30. Moral traits - what is morality? We are studying evolutionarily EVOLUTIONARY CRITERIA relevant traits only if we can for trait recognition: assess their reliable • HERITABILITY reappearance • INTER-INDIVIDUAL VARIATION “Biometrical population genetics” (cf. Serrelli 2010, Lewontin 1980) • FITNESS FUNCTION
  • 31. Moral traits - what is morality? We are studying evolutionarily EVOLUTIONARY CRITERIA relevant traits only if we can for trait recognition: assess their reliable • HERITABILITY reappearance • INTER-INDIVIDUAL VARIATION “Biometrical population genetics” (cf. Serrelli 2010, Lewontin 1980) • FITNESS FUNCTION QTL - Quantitative Traits Locis (cf. Hartl & Clark 2007, chp. 8)
  • 32. Moral traits - what is morality? We are studying evolutionarily EVOLUTIONARY CRITERIA relevant traits only if we can for trait recognition: assess their reliable • HERITABILITY reappearance • INTER-INDIVIDUAL VARIATION “Biometrical population genetics” (cf. Serrelli 2010, Lewontin 1980) • FITNESS FUNCTION QTL - Quantitative Traits Locis (cf. Hartl & Clark 2007, chp. 8) Trait = inheritable pattern
  • 33. Moral traits - what is morality? Trait = inheritable pattern EVOLUTIONARY CRITERIA for trait recognition: • HERITABILITY • INTER-INDIVIDUAL VARIATION • FITNESS FUNCTION Picture credits: chimps: http://angelorganic.org/darwin-mis- interpreted/ birds: http://news.sciencemag.org/sciencenow/ 2010/08/cheatin-hearts-get-stuck-with-th.html ants: http://www.ridelust.com/traffic-congestion-and- social-insects-we-can-learn-how-to-drive-from-ants/ V. Bulatov (2011), http://bulatov.org/math/1101/ webtalk.html
  • 34. Moral traits - what is morality? Trait = inheritable pattern EVOLUTIONARY CRITERIA for trait recognition: Need for models • HERITABILITY • INTER-INDIVIDUAL VARIATION • FITNESS FUNCTION Picture credits: chimps: http://angelorganic.org/darwin-mis- interpreted/ birds: http://news.sciencemag.org/sciencenow/ 2010/08/cheatin-hearts-get-stuck-with-th.html ants: http://www.ridelust.com/traffic-congestion-and- social-insects-we-can-learn-how-to-drive-from-ants/ V. Bulatov (2011), http://bulatov.org/math/1101/ webtalk.html
  • 35. Moral traits - what is morality? Trait = inheritable pattern EVOLUTIONARY CRITERIA for trait recognition: Need for models • HERITABILITY • INTER-INDIVIDUAL Homology: VARIATION pattern shows up • FITNESS FUNCTION in different taxa Picture credits: chimps: http://angelorganic.org/darwin-mis- interpreted/ birds: http://news.sciencemag.org/sciencenow/ 2010/08/cheatin-hearts-get-stuck-with-th.html ants: http://www.ridelust.com/traffic-congestion-and- social-insects-we-can-learn-how-to-drive-from-ants/ V. Bulatov (2011), http://bulatov.org/math/1101/ webtalk.html
  • 36. Moral traits - what is morality? Trait = inheritable pattern EVOLUTIONARY CRITERIA for trait recognition: Need for models • HERITABILITY • INTER-INDIVIDUAL Homology: VARIATION pattern shows up • FITNESS FUNCTION in different taxa Picture credits: chimps: http://angelorganic.org/darwin-mis- Trait of... interpreted/ birds: http://news.sciencemag.org/sciencenow/ 2010/08/cheatin-hearts-get-stuck-with-th.html ants: http://www.ridelust.com/traffic-congestion-and- social-insects-we-can-learn-how-to-drive-from-ants/ V. Bulatov (2011), http://bulatov.org/math/1101/ webtalk.html a relevant unit
  • 37. The function of morality In Neodarwinism not all effects are functions. Function is bound to Natural Selection A trait T function? flight? (cf. Williams 1966) effect ... effect e.g. thermoregulation? e.g. mating?
  • 38. The function of morality (1) ΔqT = [sqT(1−qT)]/ [1−s(1−qT)] (cf. Wright 1931) In Neodarwinism not all effects are functions. Function is bound to Natural Selection (cf. Williams 1966)
  • 39. The function of morality (1) ΔqT = [sqT(1−qT)]/ [1−s(1−qT)] (cf. Wright 1931) In Neodarwinism not all effects are Need to specify a “problem functions. Function is bound to Natural background” (cf. Kitcher 2011) Selection (cf. Williams 1966)
  • 40. The function of morality (1) ΔqT = [sqT(1−qT)]/ [1−s(1−qT)] (cf. Wright 1931) In Neodarwinism not all effects are Need to specify a “problem functions. Function is bound to Natural background” (cf. Kitcher 2011) Selection (cf. Williams 1966) (2) w = f (A, B, … T)
  • 41. The function of morality (1) ΔqT = [sqT(1−qT)]/ [1−s(1−qT)] (cf. Wright 1931) In Neodarwinism not all effects are Need to specify a “problem functions. Function is bound to Natural background” (cf. Kitcher 2011) Selection (cf. Williams 1966) (2) w = f (A, B, … T)
  • 42. The function of morality (1) ΔqT = [sqT(1−qT)]/ [1−s(1−qT)] (cf. Wright 1931) In Neodarwinism not all effects are Need to specify a “problem functions. Function is bound to Natural background” (cf. Kitcher 2011) Selection (cf. Williams 1966) (2) w = f (A, B, … T)
  • 43. The function of morality (1) ΔqT = [sqT(1−qT)]/ [1−s(1−qT)] (cf. Wright 1931) In Neodarwinism not all effects are Need to specify a “problem functions. Function is bound to Natural background” (cf. Kitcher 2011) Selection (cf. Williams 1966) (3) T = f (F, r)
  • 44. The function of morality (1) ΔqT = [sqT(1−qT)]/ [1−s(1−qT)] (cf. Wright 1931) In Neodarwinism not all effects are Need to specify a “problem functions. Function is bound to Natural background” (cf. Kitcher 2011) Selection (cf. Williams 1966) (3) T = f (F, r)
  • 45. The function of morality (1) ΔqT = [sqT(1−qT)]/ [1−s(1−qT)] (cf. Wright 1931) In Neodarwinism not all effects are Need to specify a “problem functions. Function is bound to Natural background” (cf. Kitcher 2011) Selection (3) T = f (F, r) Functioning (cf. Williams 1966) and roles are not causal upon traits and structures
  • 46. The function of morality (3) T = f (F, r) “...the palette of neurochemicals... affecting neurons and muscles is substantially the same across vertebrates and invertebrates...” (Churchland 2011, p. 45) Picture credit: McPartland et al. (2006), Gene 370:64-74. http:// www.sciencedirect.com/science/article/pii/S0378111905007067
  • 47. The function of morality (3) T = f (F, r) oxytocin is found in all vertebrates (Churchland 2011, p. 45) Picture credit: McPartland et al. (2006), Gene 370:64-74. http:// www.sciencedirect.com/science/article/pii/S0378111905007067
  • 48. The function of morality (3) T = f (F, r) “Oxytocin ... is at the hub of the intricate network of mammalian adaptations for caring for others” (Churchland 2011, p. 14) Picture credit: McPartland et al. (2006), Gene 370:64-74. http:// www.sciencedirect.com/science/article/pii/S0378111905007067
  • 49. The function of morality (3) T = f (F, r) “...modest modifications in exiting neural structures ... can lead to new outcomes” (Churchland 2011, p. 14) Picture credit: McPartland et al. (2006), Gene 370:64-74. http:// www.sciencedirect.com/science/article/pii/S0378111905007067
  • 50. The function of morality (3) T = f (F, r) “a large... range of social patterns found among mammals..., but underlying them are probably different arrangements of receptors for oxytocin and other hormones and neurochemicals” (Churchland 2011, p. 14) Picture credit: McPartland et al. (2006), Gene 370:64-74. http:// www.sciencedirect.com/science/article/pii/S0378111905007067
  • 51. The function of morality (3) ∆T = f (∆F, ∆r) “Biological evolution does not achieve adaptations by designing a whole new mechanism from scratch, but modifies what is already in place, little bit by little bit. Social emotions, values, and behavior are not the result of a wholly new engineering plan, but rather an adaptation of existing arrangements and mechanisms that are intimately linked with the self-preserving circuitry for fighting, freezing, and flight, on the one hand, and for rest and digest, on the other” (Churchland 2011, p. 46)
  • 52. Evolutionary explanations of morality • Explanatory diversity and integration
  • 53. Evolutionary explanations of morality • Explanatory diversity and integration • Traits/structures and functions are needed, but don’t come for free
  • 54. Evolutionary explanations of morality • Explanatory diversity and integration • Traits/structures and functions are needed, but don’t come for free • Traits as inheritable patterns
  • 55. Evolutionary explanations of morality • Explanatory diversity and integration • Traits/structures and functions are needed, but don’t come for free • Traits as inheritable patterns models
  • 56. Evolutionary explanations of morality • Explanatory diversity and integration • Traits/structures and functions are needed, but don’t come for free • Traits as inheritable patterns models homologies avoiding circularity
  • 57. Evolutionary explanations of morality • Explanatory diversity and integration • Traits/structures and functions are needed, but don’t come for free • Traits as inheritable patterns models homologies avoiding circularity units of evolution
  • 58. Evolutionary explanations of morality • Explanatory diversity and integration • Traits/structures and functions are needed, but don’t come for free • Traits as inheritable patterns models homologies avoiding circularity units of evolution
  • 59. Evolutionary explanations of morality • Explanatory diversity and integration • Traits/structures and functions are needed, but don’t come for free • Traits as inheritable patterns models homologies avoiding circularity units of evolution • Functions conceptual problems (e.g. Cummins 1975, Wright 1973, Millikan 1984, 1989, Griffiths 1993, Godfrey-Smith 1994, cf. Casebeer 2003) Picture credit: http://www.azoft.com/spotlight/2011/05/05/15-bizarre-dead-tech-products.html
  • 60. Evolutionary explanations of morality • Explanatory diversity and integration • Traits/structures and functions are needed, but don’t come for free • Traits as inheritable patterns models homologies avoiding circularity units of evolution • Functions conceptual problems (e.g. Cummins 1975, Wright 1973, Millikan 1984, 1989, Griffiths 1993, Godfrey-Smith 1994, cf. Casebeer 2003) sights for identifying traits Picture credit: http://www.azoft.com/spotlight/2011/05/05/15-bizarre-dead-tech-products.html
  • 61. Evolutionary explanations of morality • Explanatory diversity and integration • Traits/structures and functions are needed, but don’t come for free • Traits as inheritable patterns models homologies avoiding circularity units of evolution • Functions conceptual problems (e.g. Cummins 1975, Wright 1973, Millikan 1984, 1989, Griffiths 1993, Godfrey-Smith 1994, cf. Casebeer 2003) sights for identifying traits crucial for Darwinian explanations Picture credit: http://www.azoft.com/spotlight/2011/05/05/15-bizarre-dead-tech-products.html
  • 62. Evolutionary explanations of morality • Explanatory diversity and integration • Traits/structures and functions are needed, but don’t come for free • Traits as inheritable patterns models homologies avoiding circularity units of evolution • Functions conceptual problems (e.g. Cummins 1975, Wright 1973, Millikan 1984, 1989, Griffiths 1993, Godfrey-Smith 1994, cf. Casebeer 2003) sights for identifying traits crucial for Darwinian explanations functions / effects Picture credit: http://www.azoft.com/spotlight/2011/05/05/15-bizarre-dead-tech-products.html
  • 63. THANK YOU! Picture credits: http://www.azoft.com/spotlight/2011/05/05/15-bizarre-dead-tech-products.html
  • 64. REFERENCES: • Boehm C (2000), Conflict and the evolution of social control. Journal of Consciousness Studies 7(1-2), “Evolutionary Origins of Morality. Cross-Disciplinary Perspectives”, pp. 79-101. • Churchland PS (2011). Braintrust: What Neuroscience Tells Us About Morality. Princeton: Princeton University Press. • Cummins R (1975). Functional analysis. Journal of Philosophy 72:741-765. • Downes SM (2010). Heritability. The Stanford Encyclopedia of Philosophy (Fall 2010 Edition), Edward N. Zalta (ed.), URL = http://plato.stanford.edu/archives/fall2010/ entries/heredity/. • Eisenberg N, Strayer J (1987) Empathy and its development. Cambridge UP, Cambridge. • Ferraguti M, Castellacci C, eds. (2011), Evoluzione: modelli e processi. Pearson, Milano-Torino. • Flack JC, de Waal FBM (2000), ‘Any animal whatever’: Darwinian building blocks of morality in monkeys and apes. Journal of Consciousness Studies 7(1-2), “Evolutionary Origins of Morality. Cross-Disciplinary Perspectives”, pp. 1-29. • Godfrey-Smith P (1994). A modern history theory of functions. Reprinted in: Allen C, Bekoff M, Lauder G, eds. Nature’s Purposes - Analyses of Function and Design in Biology. Cambridge, MA: MIT Press, 1998, pp. 453-477. • Gould SJ (1991), “Capezzoli maschili e glande clitorideo”, in trad. it. Bravo Brontosauro, Milano, Feltrinelli Universale Economica, 2002, pp. 125‐139. • Gould SJ, Lewontin RC (1979), The spandrels of San Marco and the Panglossian paradigm: a critique of the adaptationist programme, Proc. R. Soc. Lond. B, 205, 581-598. Trad. it. di M. Ferraguti, I pennacchi di San Marco e il paradigma di Pangloss: critica al programma adattazionista. Piccola biblioteca on line, Einaudi, Torino, 2001. http://www.einaudi.it/einaudi/ita/pdf/gould-lewontin.pdf • Griffiths PE (1993). Functional analysis and proper functions. British Journal of the Philosophy of Science 44:409-422. • Griffiths PE, Brigandt I, eds. (2007). “The Importance of Homology for Biology and Philosophy”, Biology and Philosophy 22. • Hartl DL, Clark AG (2007). Principles of Population Genetics, Fourth ed. Sunderland, Mass.: Sinauer Associates. • Kitcher P (2011). The Ethical Project, Harvard University Press, Cambridge, Mass. & London, UK. • Jablonka E, Lamb MJ (2005). Evolution in Four Dimensions. Cambridge (MA): The MIT Press, revised in 2010. • Millikan R (1984). Language, Thought and Other Biological Categories. Cambridge, Mass: MIT Press. • Millikan R (1989). In defence of proper functions. Philosophy of Science 56:288-302. • Müller G.B., 2010. Epigenetic innovation. In Pigliucci M, Müller GB, eds. Evolution The Extended Synthesis. Cambridge-London: MIT Press, pp. 307-332. • Odling-Smee J, Laland K, Feldman MW (2003). Niche Construction. Princeton: Princeton University Press. • McPartland JM, Matias I, Di Marzo V, GlassM (2006). Evolutionary origins of the endocannabinoid system, Gene 370:64-74, 10.1016/j.gene.2005.11.004. • Pievani T, Serrelli E (2011). Exaptation in human evolution: how to test adaptive vs exaptive evolutionary hypotheses. Journal of Anthropological Sciences 89, pp. 9-23. [DOI 10.4436/jass.89015] • Serrelli E (2011). The structure of population genetics. Poster at the meeting of the International Society for History, Philosophy, and Social Studies of Biology (ISHPSSB), Salt Lake City (Utah, USA) July 10th-16th 2011. [http://hdl.handle.net/10281/28245] • Sober E (1984). The Nature of Selection. Evolutionary Theory in Philosophical Focus. University of Chicago Press, Chicago. • Sober E, Wilson DS (1998). Unto Others: the Evolution and Psychology of Unselfish Behavior. Harvard University Press, Cambridge (MA). • Williams GC (1966). Adaptation and Natural Selection. Princeton: Princeton University Press. • Wright L (1973). Functions. Philosophical Review 82:139-168. • Wright S (1931). Evolution in mendelian populations. Genetics, 16 (97): 97–159.

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