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Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biologyEmanuele SerrelliFinal discuss...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biologyEmanuele SerrelliFinal discuss...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biologycf. I.2.2                     ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Adaptive landscapes as a cas...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Adaptive landscapes as a cas...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Adaptive landscapes as a cas...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• The Modern Synthesis (MS):  ...
Adaptive landscapes: A case study of metaphors, models, and synthesis in evolutionary biology                             ...
Adaptive landscapes: A case study of metaphors, models, and synthesis in evolutionary biology                             ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biologycf. I.3                       ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Introducing the idea of a mi...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Introducing the idea of a mi...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Introducing the idea of a mi...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Introducing the idea of a mi...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology                              ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biologycf. I.1.3 - I.1.1 - I.1.2     ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Three common ideas to adapti...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Three common ideas to adapti...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Three common ideas to adapti...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Three common ideas to adapti...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Three common ideas to adapti...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Three common ideas to adapti...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biologycf. I.3.3                     ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• If MS was the building of a ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• If MS was the building of a ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• If MS was the building of a ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• If MS was the building of a ...
cf. I.2.3,                                                            Fig. I.6 p. 28,                                     ...
cf. I.4                                              nimbios.org    Adaptive landscapes: A casestudy of metaphors, models,...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology                              ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Proposal to clarify the rela...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Proposal to clarify the rela...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Proposal to clarify the rela...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biologycf. I.5, Fig. I.8 p. 63       ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• I proposed:    • Adaptive la...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• I proposed:    • Adaptive la...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• I proposed:    • Adaptive la...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• I proposed:    • Adaptive la...
cf. II.1• Which population genetic model(s) are adaptive  landscapes metaphors of? Unclear in literature.       14
cf. II.1• Which population genetic model(s) are adaptive  landscapes metaphors of? Unclear in literature.       14
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology                              ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Which model(s) are adaptive ...
cf. II.1,                                                                    Fig. II.3 p. 79,                             ...
cf. II.1,                                                                             Fig. II.3 p. 79,                    ...
cf. II.1,                                                                             Fig. II.3 p. 79,                    ...
cf. II.1,                                                                              Fig. II.3 p. 79,                   ...
cf. II.1,                                                                              Fig. II.3 p. 79,                   ...
cf. II.1,                                                                              Fig. II.3 p. 79,                   ...
Proposal:             Term                        Kind of object                          Synonyms                        ...
Proposal:             Term                        Kind of object                          Synonyms                        ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biologycf. III.2                     ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Today’s landscape models:cf....
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Today’s landscape models:   ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Today’s landscape models:   ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Today’s landscape models:   ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Today’s landscape models:   ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Today’s landscape models:   ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Today’s landscape models:   ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Today’s landscape models:   ...
Adaptive landscapes: A case study of metaphors,  models, and synthesis in evolutionary biology                     Mount I...
Adaptive landscapes: A case study of metaphors,  models, and synthesis in evolutionary biology                     Mount I...
Adaptive landscapes: A case study of metaphors,  models, and synthesis in evolutionary biology                     Mount I...
Adaptive landscapes: A case study of metaphors,  models, and synthesis in evolutionary biology                     Mount I...
Adaptive landscapes: A case study of metaphors,  models, and synthesis in evolutionary biology                           M...
Adaptive landscapes: A case study of metaphors,   models, and synthesis in evolutionary biologyGavrilets (2004)  Research ...
Adaptive landscapes: A case study of metaphors,   models, and synthesis in evolutionary biology                           ...
Adaptive landscapes: A case study of metaphors,   models, and synthesis in evolutionary biology                           ...
Adaptive landscapes: A case study of metaphors,   models, and synthesis in evolutionary biology                           ...
Adaptive landscapes: A case study of metaphors,   models, and synthesis in evolutionary biology                           ...
Adaptive landscapes: A case study of metaphors,   models, and synthesis in evolutionary biology                           ...
Adaptive landscapes: A case study of metaphors,   models, and synthesis in evolutionary biology                           ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biologycf. III.3.2 - III.3.3         ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• The landscape example (and p...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• The landscape example (and p...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• The landscape example (and p...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• The landscape example (and p...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• The landscape example (and p...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• The landscape example (and p...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• The landscape example (and p...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology                              ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Pigliucci’s contemporary int...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Pigliucci’s contemporary int...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Pigliucci’s contemporary int...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Pigliucci’s contemporary int...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Pigliucci’s contemporary int...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Pigliucci’s contemporary int...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Pigliucci’s contemporary int...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Pigliucci’s contemporary int...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology                              ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Challenges for the Extended ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Challenges for the Extended ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Challenges for the Extended ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Challenges for the Extended ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Challenges for the Extended ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Challenges for the Extended ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Challenges for the Extended ...
Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biologyEmanuele SerrelliFinal discuss...
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Adaptive landscapes: A case study of metaphors, models, and synthesis in evolutionary biology

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Emanuele Serrelli

Final discussion, XXIII cycle, January 17th 2011
PhD School in Human Sciences
University of Milano Bicocca
Coordinator: prof. Ottavia Albanese
Advisor: prof. Dietelmo Pievani

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Adaptive landscapes: A case study of metaphors, models, and synthesis in evolutionary biology

  1. 1. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biologyEmanuele SerrelliFinal discussion, XXIII cycle, January 2011PhD School in Human SciencesUniversity of Milano BicoccaCoordinator: prof. Ottavia AlbaneseAdvisor: prof. Dietelmo Pievani 1
  2. 2. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biologyEmanuele SerrelliFinal discussion, XXIII cycle, January 2011PhD School in Human SciencesUniversity of Milano BicoccaCoordinator: prof. Ottavia AlbaneseAdvisor: prof. Dietelmo PievaniIntroduction 1
  3. 3. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biologycf. I.2.2 2
  4. 4. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Adaptive landscapes as a case of scientific metaphorcf. I.2.2 2
  5. 5. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Adaptive landscapes as a case of scientific metaphor• Metaphor: nature and role in sciencecf. I.2.2 2
  6. 6. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Adaptive landscapes as a case of scientific metaphor• Metaphor: nature and role in science ‣ Consideration of the historical- scientific context of its spreading: the Modern Synthesiscf. I.2.2 2
  7. 7. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• The Modern Synthesis (MS): 1. a two-steps process (1910s-1930s) central in shaping today’s evolutionary biology; 2. its essence is communication among separate fields (e.g. paleontology, Ernst Mayr (1904-2005) theoretical and experimental genetics, Wikimedia Commons & PLOS Biology morphology, zoology, botanics and so on).cf. I.3.2 3
  8. 8. Adaptive landscapes: A case study of metaphors, models, and synthesis in evolutionary biology Mayr’s view of MS pbs.org paleontology «...to give a grand sense of unity George Gaylord Simpson pbs.org embracing genes, phenotypes with their adaptations to different environments, speciations, micro experimental genetics and macroevolution, singular Theodosius Dobzhansky episodes and the general pattern of evolution» (p. 208). ornithology, the naturalists Ernst Mayrsciencemag.org theoretical genetics evolutionary theory skeptic.com Sewall Wright Julian Huxley ... cf. I.3.2 huxley.net 4
  9. 9. Adaptive landscapes: A case study of metaphors, models, and synthesis in evolutionary biology Mayr’s view of MS pbs.org paleontology George Gaylord Simpson pbs.org experimental genetics Theodosius Dobzhansky ornithology, the naturalists Ernst Mayrsciencemag.org theoretical genetics evolutionary theory skeptic.com Sewall Wright Julian Huxley ... cf. I.3.2 huxley.net 5
  10. 10. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biologycf. I.3 6
  11. 11. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Introducing the idea of a migrant metaphor:cf. I.3 6
  12. 12. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Introducing the idea of a migrant metaphor: • NOT a “one-shot”, all-inclusive metaphor of evolution;cf. I.3 6
  13. 13. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Introducing the idea of a migrant metaphor: • NOT a “one-shot”, all-inclusive metaphor of evolution; • NOT a “ready-to-use” universal, context-independent tool;cf. I.3 6
  14. 14. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Introducing the idea of a migrant metaphor: • NOT a “one-shot”, all-inclusive metaphor of evolution; • NOT a “ready-to-use” universal, context-independent tool; • the migrant metaphor is found in different fields of biology, but declined differently in each.cf. I.3 6
  15. 15. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology George Gaylord Simpson (1944) Theodosius Dobzhansky (1937) Tempo and Mode in Evolution Genetics and the Origin of Species paleontologycf. I.2.1, Fig. I.2 p. 9 - I.2.2, Fig. I.3 p. 17 - I.2.4 7
  16. 16. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biologycf. I.1.3 - I.1.1 - I.1.2 8
  17. 17. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Three common ideas to adaptive landscapes:cf. I.1.3 - I.1.1 - I.1.2 8
  18. 18. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Three common ideas to adaptive landscapes: • “Higher is better” - altitude corresponds to better solutions (peaks), and worst possibilities are in valleys;cf. I.1.3 - I.1.1 - I.1.2 8
  19. 19. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Three common ideas to adaptive landscapes: • “Higher is better” - altitude corresponds to better solutions (peaks), and worst possibilities are in valleys; • “better” and “worst” are relative to a given environment which is not represented in the diagram but influences its structure;cf. I.1.3 - I.1.1 - I.1.2 8
  20. 20. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Three common ideas to adaptive landscapes: • “Higher is better” - altitude corresponds to better solutions (peaks), and worst possibilities are in valleys; • “better” and “worst” are relative to a given environment which is not represented in the diagram but influences its structure; • landscape is the whole set of possibilities for an evolving entity, and evolution is conceived as the realization of potentialities through time.cf. I.1.3 - I.1.1 - I.1.2 8
  21. 21. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Three common ideas to adaptive landscapes: • “Higher is better” - altitude corresponds to better solutions (peaks), and worst possibilities are in valleys; • “better” and “worst” are relative to a given environment which is not represented in the diagram but influences its structure; • landscape is the whole set of possibilities for an evolving entity, and evolution is conceived as the realization of potentialities through time.cf. I.1.3 - I.1.1 - I.1.2 8
  22. 22. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Three common ideas to adaptive landscapes: • “Higher is better” - altitude corresponds to better solutions (peaks), and worst possibilities are in valleys; • “better” and “worst” are relative to a given environment which is not represented in the diagram but influences Population adaptation as “climbing” its structure; • landscape is the whole set of possibilities for an evolving entity, and evolution is conceived as the realization of potentialities through time.cf. I.1.3 - I.1.1 - I.1.2 8
  23. 23. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biologycf. I.3.3 9
  24. 24. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• If MS was the building of a common language among separate fields - as opposed to an extension of mathematical models of population genetics - here we can see an important role for the adaptive landscape metaphor as:cf. I.3.3 9
  25. 25. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• If MS was the building of a common language among separate fields - as opposed to an extension of mathematical models of population genetics - here we can see an important role for the adaptive landscape metaphor as: • a figure of speech, generator of common language (peak, valley, ruggedness etc.);cf. I.3.3 9
  26. 26. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• If MS was the building of a common language among separate fields - as opposed to an extension of mathematical models of population genetics - here we can see an important role for the adaptive landscape metaphor as: • a figure of speech, generator of common language (peak, valley, ruggedness etc.); • a migrant metaphor (µεταϕοσά, “to carry over”, “to trasfer from”).cf. I.3.3 9
  27. 27. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• If MS was the building of a common language among separate fields - as opposed to an extension of mathematical models of population genetics - here we can see an important role for the adaptive landscape metaphor as: • a figure of speech, generator of common language (peak, valley, ruggedness etc.); • a migrant metaphor (µεταϕοσά, “to carry over”, “to trasfer from”).• An clue of the metaphor’s communicational effectiveness: Dawkins (1996)cf. I.3.3 9
  28. 28. cf. I.2.3, Fig. I.6 p. 28, I.2.4 froes.dds.nl Adaptive landscapes: A casestudy of metaphors, models, and Richard Dawkins (1996) synthesis in evolutionary biology Climbing Mount Improbable 10
  29. 29. cf. I.4 nimbios.org Adaptive landscapes: A casestudy of metaphors, models, and Sergey Gavrilets (1997 sgg.) synthesis in evolutionary biology Holey landscapes 11
  30. 30. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology Sewall Wright (1932) Sergey Gavrilets (1997)cf. I.5 - II.1, II.2 12
  31. 31. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Proposal to clarify the relationship between holey and rugged landscapes it is necessary to consider: the native (as opposed to migrant) adaptive landscape metaphor. Sewall Wright (1932) Sergey Gavrilets (1997)cf. I.5 - II.1, II.2 12
  32. 32. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Proposal to clarify the relationship between holey and rugged landscapes it is necessary to consider: the native (as opposed to migrant) adaptive landscape metaphor. • native theoretical context: Mendelian population genetics (presented to biologists of other specializations); Sewall Wright (1932) conceived and proposed by Sewall Wright in 1932 (paper “The roles of mutation, inbreeding, crossbreeding and selection in evolution” at the 6th International Congress of Genetics); Sergey Gavrilets (1997)cf. I.5 - II.1, II.2 12
  33. 33. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Proposal to clarify the relationship between holey and rugged landscapes it is necessary to consider: the native (as opposed to migrant) adaptive landscape metaphor. • native theoretical context: Mendelian population genetics (presented to biologists of other specializations); Sewall Wright (1932) conceived and proposed by Sewall Wright in 1932 (paper “The roles of mutation, inbreeding, crossbreeding and selection in evolution” at the 6th International Congress of Genetics); • the same native context is shared by holey landscapes. Sergey Gavrilets (1997)cf. I.5 - II.1, II.2 12
  34. 34. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biologycf. I.5, Fig. I.8 p. 63 13
  35. 35. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• I proposed: • Adaptive landscapes, in their native context, are a metaphor already, but a metaphor in a different sense:cf. I.5, Fig. I.8 p. 63 13
  36. 36. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• I proposed: • Adaptive landscapes, in their native context, are a metaphor already, but a metaphor in a different sense:cf. I.5, Fig. I.8 p. 63 13
  37. 37. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• I proposed: • Adaptive landscapes, in their native context, are a metaphor already, but a metaphor in a different sense: • Could this open to a more general reflection about the recursive nature of knowledge (involving the migrant metaphor as well)?cf. I.5, Fig. I.8 p. 63 13
  38. 38. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• I proposed: • Adaptive landscapes, in their native context, are a metaphor already, but a metaphor in a different sense: • Could this open to a more general reflection about the recursive nature of knowledge (involving the migrant metaphor as well)? • Holey and rugged landscapes are metaphors for some population genetics model. Which one?cf. I.5, Fig. I.8 p. 63 13
  39. 39. cf. II.1• Which population genetic model(s) are adaptive landscapes metaphors of? Unclear in literature. 14
  40. 40. cf. II.1• Which population genetic model(s) are adaptive landscapes metaphors of? Unclear in literature. 14
  41. 41. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology Mendelian population (high dimensionality) EPISTEMOLOGICAL GAP equations METAPHOR adaptive surfacecf. II.2.1-3, Figg. II.1-2 p. 78 - II.2.4 15
  42. 42. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Which model(s) are adaptive landscapes metaphors of? Unclear in literature: 1. Analysis of the theoretical structure of population genetics; 2. result: in its native context, adaptive landscape was metaphor of a really fundamental formal object: Mendelian population, i.e. a combination space provided with fitness; 3. the space has too many dimensions to be treated directly with mathematical equations (epistemological gap): ‣ I criticized many authors who considered landscapes as a metaphor of “too complicated equations”; ‣ I analyzed how Wright probably did arrive to know something about the combination space (intuition, limited experience, and heuristics).cf. II.2.1-3, Figg. II.1-2 p. 78 - II.2.4 16
  43. 43. cf. II.1, Fig. II.3 p. 79, Population genetics II.3 Mendelian Biometrical• A further source of confusion in literature: what is to be intended with “model”? 17
  44. 44. cf. II.1, Fig. II.3 p. 79, Population genetics II.3 Mendelian Biometrical Many notions of models are available• A further source of confusion in literature: what is to be intended with “model”? 17
  45. 45. cf. II.1, Fig. II.3 p. 79, Population genetics II.3 Mendelian Biometrical Many notions of models are available ⇓• A further source of confusion in literature: what is to be intended with “model”? 17
  46. 46. cf. II.1, Fig. II.3 p. 79, Population genetics II.3 Mendelian Biometrical Many notions of models are available ⇓ Variability in the semantic extension of the term “model”• A further source of confusion in literature: what is to be intended with “model”? 17
  47. 47. cf. II.1, Fig. II.3 p. 79, Population genetics II.3 Mendelian Biometrical Many notions of models are available ⇓ Variability in the semantic extension of the term “model” ⇓• A further source of confusion in literature: what is to be intended with “model”? 17
  48. 48. cf. II.1, Fig. II.3 p. 79, Population genetics II.3 Mendelian Biometrical Many notions of models are available ⇓ Variability in the semantic extension of the term “model” ⇓ Need for a pragmatic approach in philosophy of biology• A further source of confusion in literature: what is to be intended with “model”? 17
  49. 49. Proposal: Term Kind of object Synonyms Mendelian Model as «stable target of explanation», - Combination / a pragmatically- population similar to model organisms in genetic / metric space stated vocabulary experimental biology - Space for debating (with fitness) Mendelian Theoretical structure Wrightian population adaptive population genetics genetics (opposed to landscapes. Fisherian or biometrical) Population Dynamical system Realized (N individual) subset of the model Function of time Environment, Compressed self-evident truths phenotypes, and interactions among them Population genetics Equations of gene frequencies (one or Equations equations few diallelic loci, frequency spectrums) Landscape language Metaphorical verbal language, Includes peaks, valleys, vocabulary to communicate features of ruggedness, ridges, the model climbing etc. Landscape models Combination spaces built on imitation of Landscape spaces, Mendelian population. These models are combination spaces, «stable targets of explanation» in their spaces own right. Adaptive surface 3D visualization - Fitness surface - metaphorical in case of high- - Surface dimensional spaces - Surface picture - exact in case of low-dimensional spaces - Landscape picturecf. Table II.1 p. 116 18 Table II.1. Pragmatically-stated vocabulary for the debate on adaptive landscapes.
  50. 50. Proposal: Term Kind of object Synonyms Mendelian Model as «stable target of explanation», - Combination / a pragmatically- population similar to model organisms in genetic / metric space stated vocabulary experimental biology - Space for debating (with fitness) Mendelian Theoretical structure Wrightian population adaptive population genetics genetics (opposed to landscapes. Fisherian or biometrical) Population Dynamical system Realized (N individual) subset of the model Function of time Environment, Compressed self-evident truths phenotypes, and interactions among them Population genetics Equations of gene frequencies (one or Equations equations few diallelic loci, frequency spectrums) Landscape language Metaphorical verbal language, Includes peaks, valleys, vocabulary to communicate features of ruggedness, ridges, the model climbing etc. Landscape models Combination spaces built on imitation of Landscape spaces, Mendelian population. These models are combination spaces, «stable targets of explanation» in their spaces own right. Adaptive surface 3D visualization - Fitness surface - metaphorical in case of high- - Surface dimensional spaces - Surface picture - exact in case of low-dimensional spaces - Landscape picturecf. Table II.1 p. 116 18 Table II.1. Pragmatically-stated vocabulary for the debate on adaptive landscapes.
  51. 51. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biologycf. III.2 19
  52. 52. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Today’s landscape models:cf. III.2 19
  53. 53. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Today’s landscape models: • surface is not essential;cf. III.2 19
  54. 54. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Today’s landscape models: • surface is not essential; • they are combination spaces (thus similar to Mendelian population) to which landscape metaphorical language is applied;cf. III.2 19
  55. 55. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Today’s landscape models: • surface is not essential; • they are combination spaces (thus similar to Mendelian population) to which landscape metaphorical language is applied; • they vary by:cf. III.2 19
  56. 56. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Today’s landscape models: • surface is not essential; • they are combination spaces (thus similar to Mendelian population) to which landscape metaphorical language is applied; • they vary by: ‣ nature of factors which are combined;cf. III.2 19
  57. 57. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Today’s landscape models: • surface is not essential; • they are combination spaces (thus similar to Mendelian population) to which landscape metaphorical language is applied; • they vary by: ‣ nature of factors which are combined; ‣ dimensionality;cf. III.2 19
  58. 58. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Today’s landscape models: • surface is not essential; • they are combination spaces (thus similar to Mendelian population) to which landscape metaphorical language is applied; • they vary by: ‣ nature of factors which are combined; ‣ dimensionality; ‣ method of fitness assignment;cf. III.2 19
  59. 59. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Today’s landscape models: • surface is not essential; • they are combination spaces (thus similar to Mendelian population) to which landscape metaphorical language is applied; • they vary by: ‣ nature of factors which are combined; ‣ dimensionality; ‣ method of fitness assignment; ‣ nature of dynamics.cf. III.2 19
  60. 60. Adaptive landscapes: A case study of metaphors, models, and synthesis in evolutionary biology Mount Improbable pbs.org Richard Dawkins paleontology George Gaylord Simpson pbs.org experimental genetics Theodosius Dobzhanskysciencemag.org skeptic.com theoretical genetics Sewall Wright cf. III.1 huxley.net 20
  61. 61. Adaptive landscapes: A case study of metaphors, models, and synthesis in evolutionary biology Mount Improbable pbs.org Richard Dawkins paleontology George Gaylord Simpson pbs.org experimental genetics Theodosius Dobzhansky Very conservative, but inconsistentsciencemag.org skeptic.com theoretical genetics Sewall Wright cf. III.1 huxley.net 20
  62. 62. Adaptive landscapes: A case study of metaphors, models, and synthesis in evolutionary biology Mount Improbable pbs.org Richard Dawkins paleontology George Gaylord Simpson pbs.org Innovative (phenotypic), consistent experimental genetics Theodosius Dobzhansky Very conservative, but inconsistentsciencemag.org skeptic.com theoretical genetics Sewall Wright cf. III.1 huxley.net 20
  63. 63. Adaptive landscapes: A case study of metaphors, models, and synthesis in evolutionary biology Mount Improbable pbs.org Richard Dawkins Rhetorical, paleontology abstracting George Gaylord Simpson pbs.org Innovative (phenotypic), consistent experimental genetics Theodosius Dobzhansky Very conservative, but inconsistentsciencemag.org skeptic.com theoretical genetics Sewall Wright cf. III.1 huxley.net 20
  64. 64. Adaptive landscapes: A case study of metaphors, models, and synthesis in evolutionary biology Model = Mendelian population (= high-dimensional combination space) Research interest:NEUTRAL CHANGE Research interest: ADAPTATION cf. III.2.3, III.3 21
  65. 65. Adaptive landscapes: A case study of metaphors, models, and synthesis in evolutionary biologyGavrilets (2004) Research interest:SPECIATION Model = Mendelian population (= high-dimensional combination space) Research interest:NEUTRAL CHANGE Research interest: ADAPTATION cf. III.2.3, III.3 21
  66. 66. Adaptive landscapes: A case study of metaphors, models, and synthesis in evolutionary biology A patchwork of formal- mathematical tools:Gavrilets (2004) Statistical analyses Low-dimensional spaces Equations of gene frequencies Research interest:SPECIATION Model = Mendelian population (= high-dimensional combination space) Research interest:NEUTRAL CHANGE Research interest: ADAPTATION cf. III.2.3, III.3 21
  67. 67. Adaptive landscapes: A case study of metaphors, models, and synthesis in evolutionary biology “Holey” combination A patchwork of formal- spaces mathematical tools:Gavrilets (2004) Statistical analyses Low-dimensional spaces Equations of gene frequencies Research interest:SPECIATION Model = Mendelian population (= high-dimensional combination space) Research interest:NEUTRAL CHANGE Research interest: ADAPTATION cf. III.2.3, III.3 21
  68. 68. Adaptive landscapes: A case study of metaphors, models, and synthesis in evolutionary biology “Holey” combination A patchwork of formal- spaces mathematical tools: PercolationGavrilets (2004) analysis Statistical analyses Low-dimensional spaces Equations of gene frequencies Research interest:SPECIATION Model = Mendelian population (= high-dimensional combination space) Research interest:NEUTRAL CHANGE Research interest: ADAPTATION cf. III.2.3, III.3 21
  69. 69. Adaptive landscapes: A case study of metaphors, models, and synthesis in evolutionary biology “Holey” combination A patchwork of formal- spaces mathematical tools: PercolationGavrilets (2004) analysis Statistical analyses Low-dimensional spaces Equations of gene frequencies Research interest:SPECIATION Model = Mendelian population (= high-dimensional combination space) Research interest:NEUTRAL CHANGE Research interest: ADAPTATION cf. III.2.3, III.3 21
  70. 70. Adaptive landscapes: A case study of metaphors, models, and synthesis in evolutionary biology “Holey” combination A patchwork of formal- spaces mathematical tools: PercolationGavrilets (2004) analysis Statistical analyses Visual, metaphorical representation Low-dimensional spaces Equations of gene frequencies Research interest:SPECIATION Model = Mendelian population (= high-dimensional combination space) Research interest:NEUTRAL CHANGE Research interest: ADAPTATION cf. III.2.3, III.3 21
  71. 71. Adaptive landscapes: A case study of metaphors, models, and synthesis in evolutionary biology “Holey” combination A patchwork of formal- spaces mathematical tools: PercolationGavrilets (2004) analysis Statistical analyses Visual, metaphorical representation Low-dimensional spaces Equations of gene frequencies Research interest:SPECIATION Model = Mendelian population (= high-dimensional combination space) Research interest:NEUTRAL CHANGE Research interest: ADAPTATION cf. III.2.3, III.3 21
  72. 72. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biologycf. III.3.2 - III.3.3 22
  73. 73. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• The landscape example (and particularly the analysis of Gavrilets’s Fitness Landscapes and the Origin of Species, 2004) shows:cf. III.3.2 - III.3.3 22
  74. 74. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• The landscape example (and particularly the analysis of Gavrilets’s Fitness Landscapes and the Origin of Species, 2004) shows: • advancement of modeling is not trivially a better and more complete description of the world;cf. III.3.2 - III.3.3 22
  75. 75. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• The landscape example (and particularly the analysis of Gavrilets’s Fitness Landscapes and the Origin of Species, 2004) shows: • advancement of modeling is not trivially a better and more complete description of the world; • in population genetics there is a fundamental model, and a “patchwork” of interrelated tools thatcf. III.3.2 - III.3.3 22
  76. 76. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• The landscape example (and particularly the analysis of Gavrilets’s Fitness Landscapes and the Origin of Species, 2004) shows: • advancement of modeling is not trivially a better and more complete description of the world; • in population genetics there is a fundamental model, and a “patchwork” of interrelated tools that ‣ increase our knowledge of the model;cf. III.3.2 - III.3.3 22
  77. 77. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• The landscape example (and particularly the analysis of Gavrilets’s Fitness Landscapes and the Origin of Species, 2004) shows: • advancement of modeling is not trivially a better and more complete description of the world; • in population genetics there is a fundamental model, and a “patchwork” of interrelated tools that ‣ increase our knowledge of the model; ‣ make it suitable to be a model for phenomena of interest (adaptation? speciation?), and perhaps not others.cf. III.3.2 - III.3.3 22
  78. 78. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• The landscape example (and particularly the analysis of Gavrilets’s Fitness Landscapes and the Origin of Species, 2004) shows: • advancement of modeling is not trivially a better and more complete description of the world; • in population genetics there is a fundamental model, and a “patchwork” of interrelated tools that ‣ increase our knowledge of the model; ‣ make it suitable to be a model for phenomena of interest (adaptation? speciation?), and perhaps not others. • interests vary and change through time;cf. III.3.2 - III.3.3 22
  79. 79. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• The landscape example (and particularly the analysis of Gavrilets’s Fitness Landscapes and the Origin of Species, 2004) shows: • advancement of modeling is not trivially a better and more complete description of the world; • in population genetics there is a fundamental model, and a “patchwork” of interrelated tools that ‣ increase our knowledge of the model; ‣ make it suitable to be a model for phenomena of interest (adaptation? speciation?), and perhaps not others. • interests vary and change through time; • relationships in the patchwork (e.g. between “holey and rugged landscapes”) are to be considered carefully.cf. III.3.2 - III.3.3 22
  80. 80. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology Massimo Pigliucci (2008, 2010) friendlyatheist.comcf. IV.2.1, IV.2.2 - IV.1 23
  81. 81. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Pigliucci’s contemporary interests (worries): Massimo Pigliucci (2008, 2010) friendlyatheist.comcf. IV.2.1, IV.2.2 - IV.1 23
  82. 82. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Pigliucci’s contemporary interests (worries): • modeling phenotypes together with genotypes (cf. G➝P map); Massimo Pigliucci (2008, 2010) friendlyatheist.comcf. IV.2.1, IV.2.2 - IV.1 23
  83. 83. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Pigliucci’s contemporary interests (worries): • modeling phenotypes together with genotypes (cf. G➝P map); • evolvability (sensu Wagner & Altenberg 1996) = the ability of a population to respond to selection; Massimo Pigliucci (2008, 2010) friendlyatheist.comcf. IV.2.1, IV.2.2 - IV.1 23
  84. 84. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Pigliucci’s contemporary interests (worries): • modeling phenotypes together with genotypes (cf. G➝P map); • evolvability (sensu Wagner & Altenberg 1996) = the ability of a population to respond to selection; • general perspective: Extended Evolutionary Synthesis. Massimo Pigliucci (2008, 2010) friendlyatheist.comcf. IV.2.1, IV.2.2 - IV.1 23
  85. 85. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Pigliucci’s contemporary interests (worries): • modeling phenotypes together with genotypes (cf. G➝P map); • evolvability (sensu Wagner & Altenberg 1996) = the ability of a population to respond to selection; • general perspective: Extended Evolutionary Synthesis.• Pitfalls of his analysis of adaptive landscapes (2008): Massimo Pigliucci (2008, 2010) friendlyatheist.comcf. IV.2.1, IV.2.2 - IV.1 23
  86. 86. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Pigliucci’s contemporary interests (worries): • modeling phenotypes together with genotypes (cf. G➝P map); • evolvability (sensu Wagner & Altenberg 1996) = the ability of a population to respond to selection; • general perspective: Extended Evolutionary Synthesis.• Pitfalls of his analysis of adaptive landscapes (2008): Massimo Pigliucci (2008, 2010) friendlyatheist.com • emphasizing too much what is (legitimately) missing;cf. IV.2.1, IV.2.2 - IV.1 23
  87. 87. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Pigliucci’s contemporary interests (worries): • modeling phenotypes together with genotypes (cf. G➝P map); • evolvability (sensu Wagner & Altenberg 1996) = the ability of a population to respond to selection; • general perspective: Extended Evolutionary Synthesis.• Pitfalls of his analysis of adaptive landscapes (2008): Massimo Pigliucci (2008, 2010) friendlyatheist.com • emphasizing too much what is (legitimately) missing; • forced interpretations of past research questions as forerunners;cf. IV.2.1, IV.2.2 - IV.1 23
  88. 88. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Pigliucci’s contemporary interests (worries): • modeling phenotypes together with genotypes (cf. G➝P map); • evolvability (sensu Wagner & Altenberg 1996) = the ability of a population to respond to selection; • general perspective: Extended Evolutionary Synthesis.• Pitfalls of his analysis of adaptive landscapes (2008): Massimo Pigliucci (2008, 2010) friendlyatheist.com • emphasizing too much what is (legitimately) missing; • forced interpretations of past research questions as forerunners; • historical-epistemological conflations.cf. IV.2.1, IV.2.2 - IV.1 23
  89. 89. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology Sidlauskas et al. 2010 (NESCENT)cf. IV.1, IV.2.3 - Conclusions pp. 213 sgg. 24
  90. 90. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Challenges for the Extended Synthesis (synthesis seen as an object): Sidlauskas et al. 2010 (NESCENT)cf. IV.1, IV.2.3 - Conclusions pp. 213 sgg. 24
  91. 91. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Challenges for the Extended Synthesis (synthesis seen as an object): • having a theoretical structure; Sidlauskas et al. 2010 (NESCENT)cf. IV.1, IV.2.3 - Conclusions pp. 213 sgg. 24
  92. 92. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Challenges for the Extended Synthesis (synthesis seen as an object): • having a theoretical structure; • give a proper position, in that structure, to models and tools. Sidlauskas et al. 2010 (NESCENT)cf. IV.1, IV.2.3 - Conclusions pp. 213 sgg. 24
  93. 93. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Challenges for the Extended Synthesis (synthesis seen as an object): • having a theoretical structure; • give a proper position, in that structure, to models and tools.• Another interpretation (by NESCENT): synthesis as a continuing process. Overcome fragmentation by: Sidlauskas et al. 2010 (NESCENT)cf. IV.1, IV.2.3 - Conclusions pp. 213 sgg. 24
  94. 94. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Challenges for the Extended Synthesis (synthesis seen as an object): • having a theoretical structure; • give a proper position, in that structure, to models and tools.• Another interpretation (by NESCENT): synthesis as a continuing process. Overcome fragmentation by: Sidlauskas et al. 2010 (NESCENT) • innovative (also visual) representations of information;cf. IV.1, IV.2.3 - Conclusions pp. 213 sgg. 24
  95. 95. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Challenges for the Extended Synthesis (synthesis seen as an object): • having a theoretical structure; • give a proper position, in that structure, to models and tools.• Another interpretation (by NESCENT): synthesis as a continuing process. Overcome fragmentation by: Sidlauskas et al. 2010 (NESCENT) • innovative (also visual) representations of information; • common languages.cf. IV.1, IV.2.3 - Conclusions pp. 213 sgg. 24
  96. 96. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biology• Challenges for the Extended Synthesis (synthesis seen as an object): • having a theoretical structure; • give a proper position, in that structure, to models and tools.• Another interpretation (by NESCENT): synthesis as a continuing process. Overcome fragmentation by: Sidlauskas et al. 2010 (NESCENT) • innovative (also visual) representations of information; • common languages.• There’s clearly a role for landscapes, and for philosophy.cf. IV.1, IV.2.3 - Conclusions pp. 213 sgg. 24
  97. 97. Adaptive landscapes: A case study of metaphors,models, and synthesis in evolutionary biologyEmanuele SerrelliFinal discussion, XXIII cycle, January 2011PhD School in Human SciencesUniversity of Milano BicoccaCoordinator: prof. Ottavia AlbaneseAdvisor: prof. Dietelmo Pievani 25

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