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
APM Welcome, APM North West Network Conference, Synergies Across Sectors
Adaptive landscapes: A case study of metaphors, models, and synthesis in evolutionary biology
1. Adaptive landscapes: A case study of metaphors,
models, and synthesis in evolutionary biology
Emanuele Serrelli
Final discussion, XXIII cycle, January 2011
PhD School in Human Sciences
University of Milano Bicocca
Coordinator: prof. Ottavia Albanese
Advisor: prof. Dietelmo Pievani
1
2. Adaptive landscapes: A case study of metaphors,
models, and synthesis in evolutionary biology
Emanuele Serrelli
Final discussion, XXIII cycle, January 2011
PhD School in Human Sciences
University of Milano Bicocca
Coordinator: prof. Ottavia Albanese
Advisor: prof. Dietelmo Pievani
Introduction 1
3. Adaptive landscapes: A case study of metaphors,
models, and synthesis in evolutionary biology
cf. I.2.2 2
4. Adaptive landscapes: A case study of metaphors,
models, and synthesis in evolutionary biology
• Adaptive landscapes as a case of
scientific metaphor
cf. I.2.2 2
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 science
cf. I.2.2 2
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 Synthesis
cf. I.2.2 2
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. 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 Mayr
sciencemag.org
theoretical genetics evolutionary theory skeptic.com
Sewall Wright Julian Huxley
...
cf. I.3.2 huxley.net 4
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 Mayr
sciencemag.org
theoretical genetics evolutionary theory skeptic.com
Sewall Wright Julian Huxley
...
cf. I.3.2 huxley.net 5
10. Adaptive landscapes: A case study of metaphors,
models, and synthesis in evolutionary biology
cf. I.3 6
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. 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. 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. 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. 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 paleontology
cf. I.2.1, Fig. I.2 p. 9 - I.2.2, Fig. I.3 p. 17 - I.2.4 7
16. Adaptive landscapes: A case study of metaphors,
models, and synthesis in evolutionary biology
cf. I.1.3 - I.1.1 - I.1.2 8
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. 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. 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. 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. 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. 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. Adaptive landscapes: A case study of metaphors,
models, and synthesis in evolutionary biology
cf. I.3.3 9
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. 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. 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. 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. cf. I.2.3,
Fig. I.6 p. 28,
I.2.4
froes.dds.nl
Adaptive landscapes: A case
study of metaphors, models, and Richard Dawkins (1996)
synthesis in evolutionary biology Climbing Mount Improbable
10
29. cf. I.4
nimbios.org
Adaptive landscapes: A case
study of metaphors, models, and Sergey Gavrilets (1997 sgg.)
synthesis in evolutionary biology Holey landscapes 11
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. 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. 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. 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. Adaptive landscapes: A case study of metaphors,
models, and synthesis in evolutionary biology
cf. I.5, Fig. I.8 p. 63 13
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. 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. 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. 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. cf. II.1
• Which population genetic model(s) are adaptive
landscapes metaphors of? Unclear in literature. 14
40. cf. II.1
• Which population genetic model(s) are adaptive
landscapes metaphors of? Unclear in literature. 14
41. Adaptive landscapes: A case study of metaphors,
models, and synthesis in evolutionary biology
Mendelian
population
(high dimensionality)
EPISTEMOLOGICAL
GAP
equations METAPHOR
adaptive
surface
cf. II.2.1-3, Figg. II.1-2 p. 78 - II.2.4 15
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. 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. 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. 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. 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. 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. 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. 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 picture
cf. Table II.1 p. 116 18
Table II.1. Pragmatically-stated vocabulary for the debate on adaptive landscapes.
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 picture
cf. Table II.1 p. 116 18
Table II.1. Pragmatically-stated vocabulary for the debate on adaptive landscapes.
51. Adaptive landscapes: A case study of metaphors,
models, and synthesis in evolutionary biology
cf. III.2 19
52. Adaptive landscapes: A case study of metaphors,
models, and synthesis in evolutionary biology
• Today’s landscape models:
cf. III.2 19
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. 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. 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. 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. 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. 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. 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. 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
sciencemag.org
skeptic.com
theoretical genetics
Sewall Wright
cf. III.1 huxley.net 20
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 inconsistent
sciencemag.org
skeptic.com
theoretical genetics
Sewall Wright
cf. III.1 huxley.net 20
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 inconsistent
sciencemag.org
skeptic.com
theoretical genetics
Sewall Wright
cf. III.1 huxley.net 20
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 inconsistent
sciencemag.org
skeptic.com
theoretical genetics
Sewall Wright
cf. III.1 huxley.net 20
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. Adaptive landscapes: A case study of metaphors,
models, and synthesis in evolutionary biology
Gavrilets (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. 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. 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. Adaptive landscapes: A case study of metaphors,
models, and synthesis in evolutionary biology
“Holey”
combination
A patchwork of formal- spaces
mathematical tools:
Percolation
Gavrilets (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. Adaptive landscapes: A case study of metaphors,
models, and synthesis in evolutionary biology
“Holey”
combination
A patchwork of formal- spaces
mathematical tools:
Percolation
Gavrilets (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. Adaptive landscapes: A case study of metaphors,
models, and synthesis in evolutionary biology
“Holey”
combination
A patchwork of formal- spaces
mathematical tools:
Percolation
Gavrilets (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. Adaptive landscapes: A case study of metaphors,
models, and synthesis in evolutionary biology
“Holey”
combination
A patchwork of formal- spaces
mathematical tools:
Percolation
Gavrilets (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. Adaptive landscapes: A case study of metaphors,
models, and synthesis in evolutionary biology
cf. III.3.2 - III.3.3 22
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. 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. 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
cf. III.3.2 - III.3.3 22
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. 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. 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. 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. Adaptive landscapes: A case study of metaphors,
models, and synthesis in evolutionary biology
Massimo Pigliucci (2008, 2010)
friendlyatheist.com
cf. IV.2.1, IV.2.2 - IV.1 23
81. Adaptive landscapes: A case study of metaphors,
models, and synthesis in evolutionary biology
• Pigliucci’s contemporary interests (worries):
Massimo Pigliucci (2008, 2010)
friendlyatheist.com
cf. IV.2.1, IV.2.2 - IV.1 23
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.com
cf. IV.2.1, IV.2.2 - IV.1 23
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.com
cf. IV.2.1, IV.2.2 - IV.1 23
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.com
cf. IV.2.1, IV.2.2 - IV.1 23
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.com
cf. IV.2.1, IV.2.2 - IV.1 23
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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. Adaptive landscapes: A case study of metaphors,
models, and synthesis in evolutionary biology
Emanuele Serrelli
Final discussion, XXIII cycle, January 2011
PhD School in Human Sciences
University of Milano Bicocca
Coordinator: prof. Ottavia Albanese
Advisor: prof. Dietelmo Pievani
25