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Mendelian population as a model, intended as a
“stable target of explanation”
Emanuele Serrelli
emanuele.serrelli@unimib.it




                                                 1
2
OUTLINE
• Some super-simple examples of usually-called “population genetics models”.

• Some ideas about mathematical models that were expressed in a 1989 book on modeling (Casti &
  Karlqvist 1989), because we may still hold such ideas when we think to mathematical models.


• General pluralistic position on model notions (Leonelli 2003, 2007, Serrelli 2010, 2011).

• Report of the demarcation and defense of the notion of “model organism” (Ankeny & Leonelli 2011) as
  opposed to “experimental organism”.


• Statement of a more general notion of a model as a “stable target of explanation” (Keller 2002) =>
  several (though not all) requirements of “model organisms” end by falling into this notion.

• STE model notion describes also the essential part of population genetics, i.e. the Mendelian
  population => proposal of revising the semantic extension of “model”; revisiting standard distinctions
  like formal vs. material.


• Sketch of some interesting epistemological features and issues that pertain STE models are shared
  by model organisms and Mendelian population.


                                                                                                           3
OUTLINE
• Some super-simple examples of usually-called “population genetics models”.

• Some ideas about mathematical models that were expressed in a 1989 book on modeling (Casti &
  Karlqvist 1989), because we may still hold such ideas when we think to mathematical models.


• General pluralistic position on model notions (Leonelli 2003, 2007, Serrelli 2010, 2011).

• Report of the demarcation and defense of the notion of “model organism” (Ankeny & Leonelli 2011) as
  opposed to “experimental organism”.


• Statement of a more general notion of a model as a “stable target of explanation” (Keller 2002) =>
  several (though not all) requirements of “model organisms” end by falling into this notion.

• STE model notion describes also the essential part of population genetics, i.e. the Mendelian
  population => proposal of revising the semantic extension of “model”; revisiting standard distinctions
  like formal vs. material.


• Sketch of some interesting epistemological features and issues that pertain STE models are shared
  by model organisms and Mendelian population.


                                                                                                           4
OUTLINE
• Some super-simple examples of usually-called “population genetics models”.

• Some ideas about mathematical models that were expressed in a 1989 book on modeling (Casti &
  Karlqvist 1989), because we may still hold such ideas when we think to mathematical models.


• General pluralistic position on model notions (Leonelli 2003, 2007, Serrelli 2010, 2011).

• Report of the demarcation and defense of the notion of “model organism” (Ankeny & Leonelli 2011) as
  opposed to “experimental organism”.


• Statement of a more general notion of a model as a “stable target of explanation” (Keller 2002) =>
  several (though not all) requirements of “model organisms” end by falling into this notion.

• STE model notion describes also the essential part of population genetics, i.e. the Mendelian
  population => proposal of revising the semantic extension of “model”; revisiting standard distinctions
  like formal vs. material.


• Sketch of some interesting epistemological features and issues that pertain STE models are shared
  by model organisms and Mendelian population.


                                                                                                           5
OUTLINE
• Some super-simple examples of usually-called “population genetics models”.

• Some ideas about mathematical models that were expressed in a 1989 book on modeling (Casti &
  Karlqvist 1989), because we may still hold such ideas when we think to mathematical models.


• General pluralistic position on model notions (Leonelli 2003, 2007, Serrelli 2010, 2011).

• Report of the demarcation and defense of the notion of “model organism” (Ankeny & Leonelli 2011) as
  opposed to “experimental organism”.


• Statement of a more general notion of a model as a “stable target of explanation” (Keller 2002) =>
  several (though not all) requirements of “model organisms” end by falling into this notion.

• STE model notion describes also the essential part of population genetics, i.e. the Mendelian
  population => proposal of revising the semantic extension of “model”; revisiting standard distinctions
  like formal vs. material.


• Sketch of some interesting epistemological features and issues that pertain STE models are shared
  by model organisms and Mendelian population.


                                                                                                           6
OUTLINE
• Some super-simple examples of usually-called “population genetics models”.

• Some ideas about mathematical models that were expressed in a 1989 book on modeling (Casti &
  Karlqvist 1989), because we may still hold such ideas when we think to mathematical models.


• General pluralistic position on model notions (Leonelli 2003, 2007, Serrelli 2010, 2011).

• Report of the demarcation and defense of the notion of “model organism” (Ankeny & Leonelli 2011) as
  opposed to “experimental organism”.


• Statement of a more general notion of a model as a “stable target of explanation” (Keller 2002) =>
  several (though not all) requirements of “model organisms” end by falling into this notion.

• STE model notion describes also the essential part of population genetics, i.e. the Mendelian
  population => proposal of revising the semantic extension of “model”; revisiting standard distinctions
  like formal vs. material.


• Sketch of some interesting epistemological features and issues that pertain STE models are shared
  by model organisms and Mendelian population.


                                                                                                           7
OUTLINE
• Some super-simple examples of usually-called “population genetics models”.

• Some ideas about mathematical models that were expressed in a 1989 book on modeling (Casti &
  Karlqvist 1989), because we may still hold such ideas when we think to mathematical models.


• General pluralistic position on model notions (Leonelli 2003, 2007, Serrelli 2010, 2011).

• Report of the demarcation and defense of the notion of “model organism” (Ankeny & Leonelli 2011) as
  opposed to “experimental organism”.


• Statement of a more general notion of a model as a “stable target of explanation” (Keller 2002) =>
  several (though not all) requirements of “model organisms” end by falling into this notion.

• STE model notion describes also the essential part of population genetics, i.e. the Mendelian
  population => proposal of revising the semantic extension of “model”; revisiting standard distinctions
  like formal vs. material.


• Sketch of some interesting epistemological features and issues that pertain STE models are shared
  by model organisms and Mendelian population.


                                                                                                           8
OUTLINE
• Some super-simple examples of usually-called “population genetics models”.

• Some ideas about mathematical models that were expressed in a 1989 book on modeling (Casti &
  Karlqvist 1989), because we may still hold such ideas when we think to mathematical models.


• General pluralistic position on model notions (Leonelli 2003, 2007, Serrelli 2010, 2011).

• Report of the demarcation and defense of the notion of “model organism” (Ankeny & Leonelli 2011) as
  opposed to “experimental organism”.


• Statement of a more general notion of a model as a “stable target of explanation” (Keller 2002) =>
  several (though not all) requirements of “model organisms” end by falling into this notion.

• STE model notion describes also the essential part of population genetics, i.e. the Mendelian
  population => proposal of revising the semantic extension of “model”; revisiting standard distinctions
  like formal vs. material.


• Sketch of some interesting epistemological features and issues that pertain STE models are shared
  by model organisms and Mendelian population.


                                                                                                           9
OUTLINE
• Some super-simple examples of usually-called “population genetics models”.

• Some ideas about mathematical models that were expressed in a 1989 book on modeling (Casti &
  Karlqvist 1989), because we may still hold such ideas when we think to mathematical models.


• General pluralistic position on model notions (Leonelli 2003, 2007, Serrelli 2010, 2011).

• Report of the demarcation and defense of the notion of “model organism” (Ankeny & Leonelli 2011) as
  opposed to “experimental organism”.


• Statement of a more general notion of a model as a “stable target of explanation” (Keller 2002)
  several (though not all) requirements of “model organisms” end by falling into this notion.

• STE model notion describes also the essential part of population genetics, i.e. the Mendelian
  population => proposal of revising the semantic extension of “model”; revisiting standard distinctions
  like formal vs. material.


• Sketch of some interesting epistemological features and issues that pertain STE models are shared
  by model organisms and Mendelian population.


                                                                                                           10
“Population genetics models”

• a, A = two alleles in a diallelic locus, in an indefinitely large population




                                                                                 11
“Population genetics models”

• a, A = two alleles in a diallelic locus, in an indefinitely large population


• q = the frequency of allele A in the population




                                                                                 11
“Population genetics models”

• a, A = two alleles in a diallelic locus, in an indefinitely large population


• q = the frequency of allele A in the population


• [(1-q)a+qA] = 1 relative frequencies of the two alleles




                                                                                 11
“Population genetics models”

• a, A = two alleles in a diallelic locus, in an indefinitely large population


• q = the frequency of allele A in the population


• [(1-q)a+qA] = 1 relative frequencies of the two alleles


• aa, Aa, AA zygotes -> what frequencies?




                                                                                 11
“Population genetics models”

• a, A = two alleles in a diallelic locus, in an indefinitely large population


• q = the frequency of allele A in the population


• [(1-q)a+qA] = 1 relative frequencies of the two alleles


• aa, Aa, AA zygotes -> what frequencies?


• Hardy-Weinberg equilibrium
  (expansion of [(1-q)a+qA]2)




                                                                                 11
“Population genetics models”

• a, A = two alleles in a diallelic locus, in an indefinitely large population


• q = the frequency of allele A in the population


• [(1-q)a+qA] = 1 relative frequencies of the two alleles


• aa, Aa, AA zygotes -> what frequencies?


• Hardy-Weinberg equilibrium
  (expansion of [(1-q)a+qA]2)




                                                                                 11
“Population genetics models”

• MUTATION


    • Δq = -uq + v(1 - q) allele A’s frequency as a function of mutation
      rates




                                                                           12
“Population genetics models”

• MUTATION


    • Δq = -uq + v(1 - q) allele A’s frequency as a function of mutation
      rates


• SELECTION


    • [(1-s)(1-q)a+qA]/[1-s(1-q)] = 1 relative frequencies in presence of
      negative selection s on a


    • Δq = [sq(1-q)]/[1-s(1-q)] change in the frequency of A




                                                                            12
“Population genetics models”

• MUTATION


     • Δq = -uq + v(1 - q) allele A’s frequency as a function of mutation
       rates


• SELECTION


     • [(1-s)(1-q)a+qA]/[1-s(1-q)] = 1 relative frequencies in presence of
       negative selection s on a


     • Δq = [sq(1-q)]/[1-s(1-q)] change in the frequency of A


• More and more complicated equations can be built...

                                                                             12
“Population genetics models”

• Work in population genetics goes on and on still today, developing those
  earlier ideas, as in the following example (Hartl & Clark 2007, pp. 97-98):




                                                                                13
“Population genetics models”

• Work in population genetics goes on and on still today, developing those
  earlier ideas, as in the following example (Hartl & Clark 2007, pp. 97-98):




• All this – basically, equations – is commonly referred to as “population
  genetics models”

                                                                                13
Ideas on mathematical models




• Every model, mathematical or otherwise, is a way of representing some aspects of the real world in an
  abbreviated, or encapsulated, form. Mathematical models translate certain features of a natural system
  N into the elements of a mathematical system M, with the goal being to mirror whatever is relevant
  about N in the properties of M.


• The […] diagram shows the two essential aspects of a mathematical model: (i) An encoding
  operation by which the explanatory scheme for the real-world system N is translated into the
  language of the formal system M, and (ii) a decoding process whereby the logical inferences in M
  are translated back into predictions about the temporal behavior in N.

  (Casti & Karlqvist 1989 p. 3).

                                                                                                           14
Ideas on mathematical models




• Every model, mathematical or otherwise, is a way of representing some aspects of the real world in an
  abbreviated, or encapsulated, form. Mathematical models translate certain features of a natural system
  N into the elements of a mathematical system M, with the goal being to mirror whatever is relevant
  about N in the properties of M.


• The […] diagram shows the two essential aspects of a mathematical model: (i) An encoding
  operation by which the explanatory scheme for the real-world system N is translated into the
  language of the formal system M, and (ii) a decoding process whereby the logical inferences in M
  are translated back into predictions about the temporal behavior in N.

  (Casti & Karlqvist 1989 p. 3).

                                                                                                           14
Ideas on mathematical models




• Every model, mathematical or otherwise, is a way of representing some aspects of the real world in an
  abbreviated, or encapsulated, form. Mathematical models translate certain features of a natural system
  N into the elements of a mathematical system M, with the goal being to mirror whatever is relevant
  about N in the properties of M.


• The […] diagram shows the two essential aspects of a mathematical model: (i) An encoding
  operation by which the explanatory scheme for the real-world system N is translated into the
  language of the formal system M, and (ii) a decoding process whereby the logical inferences in M
  are translated back into predictions about the temporal behavior in N.

  (Casti & Karlqvist 1989 p. 3).

                                                                                                           14
Ideas on mathematical models




• Every model, mathematical or otherwise, is a way of representing some aspects of the real world in an
  abbreviated, or encapsulated, form. Mathematical models translate certain features of a natural system N
  into the elements of a mathematical system M, with the goal being to mirror whatever is relevant about N
  in the properties of M.


• The […] diagram shows the two essential aspects of a mathematical model: (i) An encoding operation by
  which the explanatory scheme for the real-world system N is translated into the language of the formal
  system M, and (ii) a decoding process whereby the logical inferences in M are translated back into
  predictions about the temporal behavior in N.

  (Casti & Karlqvist 1989 p. 3).


                                                                                                             15
Ideas on mathematical models




• Every model, mathematical or otherwise, is a way of representing some aspects of the real world in an
  abbreviated, or encapsulated, form. Mathematical models translate certain features of a natural system N
  into the elements of a mathematical system M, with the goal being to mirror whatever is relevant about N
  in the properties of M.


• The […] diagram shows the two essential aspects of a mathematical model: (i) An encoding operation by
  which the explanatory scheme for the real-world system N is translated into the language of the formal
  system M, and (ii) a decoding process whereby the logical inferences in M are translated back into
  predictions about the temporal behavior in N.

  (Casti & Karlqvist 1989 p. 3).


                                                                                                             15
Ideas on mathematical models




• An “axiom of modeling faith” holds that it is possible to bring into harmony
  the two worlds, i.e. the causal structure of the external world, and the
  inferential structure of the internal world. Moreover, such harmony is seen
  as the condition for a modeling relation to subsist between M and N
  (Rosen, pp. 16-17).


                                                                                 16
Ideas on mathematical models




                               17
Ideas on mathematical models




                               17
Pluralism of model notions

• “Every model, mathematical or otherwise...” ???




                                                    18
Pluralism of model notions

• “Every model, mathematical or otherwise...” ???


• Leonelli (2007) defined the “single model approach” as “the tendency to
  explain away, rather than value and analyse, the diversity among models”.




                                                                              18
Pluralism of model notions

• “Every model, mathematical or otherwise...” ???


• Leonelli (2007) defined the “single model approach” as “the tendency to
  explain away, rather than value and analyse, the diversity among models”.


• For Leonelli, the diversity of models is scientifically important: it secures
  “several epistemic goals of potential interest to practicing scientists”, and
  it allows biologists to combine them in order to pursue their research
  outcomes.




                                                                                  18
Pluralism of model notions

• “Every model, mathematical or otherwise...” ???


• Leonelli (2007) defined the “single model approach” as “the tendency to
  explain away, rather than value and analyse, the diversity among models”.


• For Leonelli, the diversity of models is scientifically important: it secures
  “several epistemic goals of potential interest to practicing scientists”, and
  it allows biologists to combine them in order to pursue their research
  outcomes.


• Should we be permanently content of grouping heterogeneous activities
  under the single term “modeling”?




                                                                                  18
Pluralism of model notions

• “Every model, mathematical or otherwise...” ???


• Leonelli (2007) defined the “single model approach” as “the tendency to
  explain away, rather than value and analyse, the diversity among models”.


• For Leonelli, the diversity of models is scientifically important: it secures
  “several epistemic goals of potential interest to practicing scientists”, and
  it allows biologists to combine them in order to pursue their research
  outcomes.


• Should we be permanently content of grouping heterogeneous activities
  under the single term “modeling”?


• Surely a pluralistic account is the best thing we can do for now.

                                                                                  18
Model organisms




• Demarcating the concept “model organisms” vs. “experimental
  organisms” (Ankeny & Leonelli 2011).




                                                                19
Model organisms




• Demarcating the concept “model organisms” vs. “experimental
  organisms” (Ankeny & Leonelli 2011).


     • Model organisms are non-human species that are extensively studied in
       order to understand a range of biological phenomena, with the hope that
       data and theories generated through use of the model will be applicable to
       other organisms, particularly those that are in some way more complex than
       the original model (p. 313).




                                                                                    19
Model organisms




• Demarcating the concept “model organisms” vs. “experimental
  organisms” (Ankeny & Leonelli 2011).


     • Model organisms are non-human species that are extensively studied in
       order to understand a range of biological phenomena, with the hope that
       data and theories generated through use of the model will be applicable to
       other organisms, particularly those that are in some way more complex than
       the original model (p. 313).


• Model organisms can be clearly distinguished from the broader class of
  experimental organisms by several features.

                                                                                    19
Model organisms




• My view: several features identified by Ankeny & Leonelli fall into a more
  general category - not experimental organism, but “stable target of
  explanation”.




                                                                               20
Model organisms




• My view: several features identified by Ankeny & Leonelli fall into a more
  general category - not experimental organism, but “stable target of
  explanation”.


• Two exclusive features of model organisms:


      • Material features


      • Representational target

                                                                               20
Model organisms




• My view: several features identified by Ankeny & Leonelli fall into a more
  general category - not experimental organism, but “stable target of
  explanation”.


• Two exclusive features of model organisms:
                                     to become (and remain) model, organisms
                                     have to be suitable to be brood and tamed
                                     cost-effectively; the “wild type strain” has to
      • Material features
                                     be isolated and standardized, so to assure
                                      the comparability of results across a large
                                              research community, etc.
      • Representational target

                                                                                       20
Model organisms




• My view: several features identified by Ankeny & Leonelli fall into a more
  general category - not experimental organism, but “stable target of
  explanation”.


• Two exclusive features of model organisms:


      • Material features


      • Representational target

                                                                               21
Model organisms




• My view: several features identified by Ankeny & Leonelli fall into a more
                             differs from representational
  general category - not experimental organism, but “stable target of
                                         scope
  explanation”.
                         describes the conceptual reasons
• Two exclusive features of model organisms:
                          why researchers are studying a


      • Material features


      • Representational target

                                                                               21
Model organisms




• My view: several features identified by Ankeny & Leonelli fall into a more
                                                whole, intact organisms
  general category - not experimental organism, but “stable target of
  explanation”.                               …model organisms […] involve
                                            attempts to generate complete
                                            knowledge of the fundamental
• Two exclusive features of model organisms:processes at work […] including
                                                the molecular, cellular, and
                                           developmental processes; in this
     • Material features                      sense the model organism is
                                              understood as a test tube for
                                            achieving a full understanding of
     • Representational target              all biological processes (p. 317).


                                                                                 22
Stable target of explanation (STE)




                                     23
Stable target of explanation (STE)




• ...[model in experimental biology] is an organism, an organism that can be
  taken to represent (that is, stand in for) a class of organisms. A model in this
  sense is not expected to serve an explanatory function in itself, nor is it a
  simplified representation of a more complex phenomenon for which we
  already have explanatory handles. Rather, its primary function is to provide
  simply a stable target of explanation (Keller 2002, p. 115).

                                                                                     24
Stable target of explanation (STE)
1. targets of explanation: not immediately tools for explaining




{
                                                                  25
Stable target of explanation (STE)
1. targets of explanation: not immediately tools for explaining


2. autonomous: from theory and from data (Morgan & Morrison 1999)




{
                                                                    25
Stable target of explanation (STE)
1. targets of explanation: not immediately tools for explaining


2. autonomous: from theory and from data (Morgan & Morrison 1999)


3. stable: answers to the challenge of producing lawlike knowledge in fields such as experimental biology (Creager et
   al. 2007, p. 5)




{
                                                                                                                        25
Stable target of explanation (STE)
1. targets of explanation: not immediately tools for explaining


2. autonomous: from theory and from data (Morgan & Morrison 1999)


3. stable: answers to the challenge of producing lawlike knowledge in fields such as experimental biology (Creager et
   al. 2007, p. 5)




{
4. representational scope




                                                                                                                        25
Stable target of explanation (STE)
1. targets of explanation: not immediately tools for explaining


2. autonomous: from theory and from data (Morgan & Morrison 1999)


3. stable: answers to the challenge of producing lawlike knowledge in fields such as experimental biology (Creager et
   al. 2007, p. 5)




{
4. representational scope




                                      how extensively the results […] can be
                                   projected onto a wider group of organisms

                                    ...the extent to which researchers see their
                                     findings as applicable across organisms
                                             (Ankeny & Leonelli, p. 315).


                                                                                                                        25
Stable target of explanation (STE)
1. targets of explanation: not immediately tools for explaining


2. autonomous: from theory and from data (Morgan & Morrison 1999)


3. stable: answers to the challenge of producing lawlike knowledge in fields such as experimental biology (Creager et
   al. 2007, p. 5)




{
4. representational scope is tendentially wide and changeable




                                                                                                                        26
Stable target of explanation (STE)
1. targets of explanation: not immediately tools for explaining


2. autonomous: from theory and from data (Morgan & Morrison 1999)


3. stable: answers to the challenge of producing lawlike knowledge in fields such as experimental biology (Creager et
   al. 2007, p. 5)




{
4. representational scope is tendentially wide and changeable


5. unified research community: ethos of sharing reinforces stability




                                                                                                                        26
Stable target of explanation (STE)
1. targets of explanation: not immediately tools for explaining


2. autonomous: from theory and from data (Morgan & Morrison 1999)


3. stable: answers to the challenge of producing lawlike knowledge in fields such as experimental biology (Creager et
   al. 2007, p. 5)




{
4. representational scope is tendentially wide and changeable


5. unified research community: ethos of sharing reinforces stability


6. socio-technical features: associated “experimental resources”, standardization, comparability, cumulative
   establishment of techniques, practices, and results




                                                                                                                        26
Stable target of explanation (STE)
1. targets of explanation: not immediately tools for explaining


2. autonomous: from theory and from data (Morgan & Morrison 1999)


3. stable: answers to the challenge of producing lawlike knowledge in fields such as experimental biology (Creager et
   al. 2007, p. 5)




{
4. representational scope is tendentially wide and changeable


5. unified research community: ethos of sharing reinforces stability


6. socio-technical features: associated “experimental resources”, standardization, comparability, cumulative
   establishment of techniques, practices, and results


7. artificiality: even model organisms “have been developed using complex processes of standardization that allow the
   establishment of a standard strain which then serves as the basis for future research” (Ankeny & Leonelli, p. 316, cf.
   e.g. Clarke & Fujimura 1992)




                                                                                                                        26
Stable target of explanation (STE)
1. targets of explanation: not immediately tools for explaining


2. autonomous: from theory and from data (Morgan & Morrison 1999)


3. stable: answers to the challenge of producing lawlike knowledge in fields such as experimental biology (Creager et
   al. 2007, p. 5)




{
4. representational scope is tendentially wide and changeable


5. unified research community: ethos of sharing reinforces stability


6. socio-technical features: associated “experimental resources”, standardization, comparability, cumulative
   establishment of techniques, practices, and results


7. artificiality: even model organisms “have been developed using complex processes of standardization that allow the
   establishment of a standard strain which then serves as the basis for future research” (Ankeny & Leonelli, p. 316, cf.
   e.g. Clarke & Fujimura 1992)


8. inexhaustedness: “…although model organisms are standardized in order to facilitate highly controlled biological
   experimentation, their inherent complexity means that the systems are never fully understood and can continue to
   generate surprising results” (Creager et al. 2007, p. 7)


                                                                                                                        26
How STE applies to population genetics

• To understand how the STE model notion
  applies to mathematical population genetics,
  we have to move our focus away from
  equations, and recognize that there is
  another, more fundamental object: the
  Mendelian population.




• i.e., the large-scale derivation of Mendel’s
  rules of inheritance.



                                                 27
How STE applies to population genetics

• To understand how the STE model notion
  applies to mathematical population genetics,
  we have to move our focus away from
  equations, and recognize that there is
  another, more fundamental object: the
  Mendelian population.




• i.e., the large-scale derivation of Mendel’s
  rules of inheritance.



                                                 27
How STE applies to population genetics

• To understand how the STE model notion
  applies to mathematical population genetics,
  we have to move our focus away from
  equations, and recognize that there is
  another, more fundamental object: the
  Mendelian population.




• i.e., the large-scale derivation of Mendel’s
  rules of inheritance.



                                                 27
How STE applies to population genetics

• Mendelian population is the space of all
  possible individual combinations given a
  number of loci and a number of alleles => a
  combination space


     • alleles
                 :     :     :
     • loci


     • individual combinations




                                                28
How STE applies to population genetics

• Mendelian population is the space of all
  possible individual combinations given a
  number of loci and a number of alleles => a
  combination space


     • alleles
                 :     :     :
     • loci


     • individual combinations


• What is the relationship between this space
  and population genetics equations?
  An epistemological gap!

                                                28
How STE applies to population genetics

• Only some positions in the combination space are
  actually occupied at a certain time. Which
  combinations are realized?




                                                     29
How STE applies to population genetics

• Only some positions in the combination space are
  actually occupied at a certain time. Which
  combinations are realized?


• With a minimally realistic number of loci and
  alleles, the dimensionality of this space is so high
  that no equation or algorithm can be developed.




                                                         29
How STE applies to population genetics

• Only some positions in the combination space are
  actually occupied at a certain time. Which
  combinations are realized?


• With a minimally realistic number of loci and
  alleles, the dimensionality of this space is so high
  that no equation or algorithm can be developed.


• Statistical equations address what happens to the
  allele frequencies in one or two loci in a
  population inhabiting an oversimplified di-allelic
  Mendelian population space.




                                                         29
How STE applies to population genetics

• Only some positions in the combination space are
  actually occupied at a certain time. Which
  combinations are realized?


• With a minimally realistic number of loci and
  alleles, the dimensionality of this space is so high
  that no equation or algorithm can be developed.


• Statistical equations address what happens to the
  allele frequencies in one or two loci in a
  population inhabiting an oversimplified di-allelic
  Mendelian population space.


• Even the more complicated population genetics
  equations are incredibly partial statistical studies
  of the Mendelian population space.

                                                         29
How STE applies to population genetics




• The Mendelian population (a combination
  space) is, in my view, it is the model of
  population genetics (at least of Mendelian
  population genetics), it is the space equations
  are about.




                                                    30
How STE applies to population genetics
• Review and apply STE features:

          1. target of explanation


          2. autonomous

          3. stable


          4. representational scope

          5. unified research community


          6. socio-technical features

          7. artificiality


          8. inexhaustedness

                                          31
How STE applies to population genetics
• Review and apply STE features:

          1. target of explanation


          2. autonomous

          3. stable


          4. representational scope

          5. unified research community


          6. socio-technical features

          7. artificiality                the “connectedness” structure of
                                             Mendelian population is very
          8. inexhaustedness                different from what we always
                                                   imagined intuitively
                                                                             32
How STE applies to population genetics
• Review and apply STE features:           this discovery enlarges the
                                            representational scope of
          1. target of explanation        Mendelian population: from
                                          adaptation to speciation too
          2. autonomous

          3. stable


          4. representational scope

          5. unified research community


          6. socio-technical features

          7. artificiality                   the “connectedness” among
                                          genotypes in Mendelian population
          8. inexhaustedness                is very different from what we
                                              always imagined intuitively
                                                                              33
Issues about STE model notion

• Epistemological questions (dilemmas?).
  If, as several authors point out (e.g., Creager et al. 2007), models are not
  chosen because they are typical of a certain set of systems, nor they are
  built to represent some other system by reduction, deduction, encoding
  (Casti & Karlqvist 1989, Rosen 1989) or the like, how can they...


   • REPRESENT?


   • EXPLAIN?


   • PREDICT?




                                                                                 34
Issues about STE model notion


• Discussing such relationships is not
  essential within a notion of a model as a
  stable target of explanation. That is, if we
  choose this notion of model we can
  provisionally remain silent on how and
  what the model represents and explains.




                                                 35
Issues about STE model notion


• Discussing such relationships is not
  essential within a notion of a model as a
  stable target of explanation. That is, if we
  choose this notion of model we can
  provisionally remain silent on how and
  what the model represents and explains.


• The most notable fact is that all the issues   Formal   Material
  are shared between a formal and a material
  models.




                                                                     35
Issues about STE model notion


• Discussing such relationships is not
  essential within a notion of a model as a
  stable target of explanation. That is, if we
  choose this notion of model we can
  provisionally remain silent on how and
  what the model represents and explains.


• The most notable fact is that all the issues   Formal   Material
  are shared between a formal and a material
  models.




                                                                     35
THANK
 YOU!




        36

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Models in Population Genetics and Organisms

  • 1. Mendelian population as a model, intended as a “stable target of explanation” Emanuele Serrelli emanuele.serrelli@unimib.it 1
  • 2. 2
  • 3. OUTLINE • Some super-simple examples of usually-called “population genetics models”. • Some ideas about mathematical models that were expressed in a 1989 book on modeling (Casti & Karlqvist 1989), because we may still hold such ideas when we think to mathematical models. • General pluralistic position on model notions (Leonelli 2003, 2007, Serrelli 2010, 2011). • Report of the demarcation and defense of the notion of “model organism” (Ankeny & Leonelli 2011) as opposed to “experimental organism”. • Statement of a more general notion of a model as a “stable target of explanation” (Keller 2002) => several (though not all) requirements of “model organisms” end by falling into this notion. • STE model notion describes also the essential part of population genetics, i.e. the Mendelian population => proposal of revising the semantic extension of “model”; revisiting standard distinctions like formal vs. material. • Sketch of some interesting epistemological features and issues that pertain STE models are shared by model organisms and Mendelian population. 3
  • 4. OUTLINE • Some super-simple examples of usually-called “population genetics models”. • Some ideas about mathematical models that were expressed in a 1989 book on modeling (Casti & Karlqvist 1989), because we may still hold such ideas when we think to mathematical models. • General pluralistic position on model notions (Leonelli 2003, 2007, Serrelli 2010, 2011). • Report of the demarcation and defense of the notion of “model organism” (Ankeny & Leonelli 2011) as opposed to “experimental organism”. • Statement of a more general notion of a model as a “stable target of explanation” (Keller 2002) => several (though not all) requirements of “model organisms” end by falling into this notion. • STE model notion describes also the essential part of population genetics, i.e. the Mendelian population => proposal of revising the semantic extension of “model”; revisiting standard distinctions like formal vs. material. • Sketch of some interesting epistemological features and issues that pertain STE models are shared by model organisms and Mendelian population. 4
  • 5. OUTLINE • Some super-simple examples of usually-called “population genetics models”. • Some ideas about mathematical models that were expressed in a 1989 book on modeling (Casti & Karlqvist 1989), because we may still hold such ideas when we think to mathematical models. • General pluralistic position on model notions (Leonelli 2003, 2007, Serrelli 2010, 2011). • Report of the demarcation and defense of the notion of “model organism” (Ankeny & Leonelli 2011) as opposed to “experimental organism”. • Statement of a more general notion of a model as a “stable target of explanation” (Keller 2002) => several (though not all) requirements of “model organisms” end by falling into this notion. • STE model notion describes also the essential part of population genetics, i.e. the Mendelian population => proposal of revising the semantic extension of “model”; revisiting standard distinctions like formal vs. material. • Sketch of some interesting epistemological features and issues that pertain STE models are shared by model organisms and Mendelian population. 5
  • 6. OUTLINE • Some super-simple examples of usually-called “population genetics models”. • Some ideas about mathematical models that were expressed in a 1989 book on modeling (Casti & Karlqvist 1989), because we may still hold such ideas when we think to mathematical models. • General pluralistic position on model notions (Leonelli 2003, 2007, Serrelli 2010, 2011). • Report of the demarcation and defense of the notion of “model organism” (Ankeny & Leonelli 2011) as opposed to “experimental organism”. • Statement of a more general notion of a model as a “stable target of explanation” (Keller 2002) => several (though not all) requirements of “model organisms” end by falling into this notion. • STE model notion describes also the essential part of population genetics, i.e. the Mendelian population => proposal of revising the semantic extension of “model”; revisiting standard distinctions like formal vs. material. • Sketch of some interesting epistemological features and issues that pertain STE models are shared by model organisms and Mendelian population. 6
  • 7. OUTLINE • Some super-simple examples of usually-called “population genetics models”. • Some ideas about mathematical models that were expressed in a 1989 book on modeling (Casti & Karlqvist 1989), because we may still hold such ideas when we think to mathematical models. • General pluralistic position on model notions (Leonelli 2003, 2007, Serrelli 2010, 2011). • Report of the demarcation and defense of the notion of “model organism” (Ankeny & Leonelli 2011) as opposed to “experimental organism”. • Statement of a more general notion of a model as a “stable target of explanation” (Keller 2002) => several (though not all) requirements of “model organisms” end by falling into this notion. • STE model notion describes also the essential part of population genetics, i.e. the Mendelian population => proposal of revising the semantic extension of “model”; revisiting standard distinctions like formal vs. material. • Sketch of some interesting epistemological features and issues that pertain STE models are shared by model organisms and Mendelian population. 7
  • 8. OUTLINE • Some super-simple examples of usually-called “population genetics models”. • Some ideas about mathematical models that were expressed in a 1989 book on modeling (Casti & Karlqvist 1989), because we may still hold such ideas when we think to mathematical models. • General pluralistic position on model notions (Leonelli 2003, 2007, Serrelli 2010, 2011). • Report of the demarcation and defense of the notion of “model organism” (Ankeny & Leonelli 2011) as opposed to “experimental organism”. • Statement of a more general notion of a model as a “stable target of explanation” (Keller 2002) => several (though not all) requirements of “model organisms” end by falling into this notion. • STE model notion describes also the essential part of population genetics, i.e. the Mendelian population => proposal of revising the semantic extension of “model”; revisiting standard distinctions like formal vs. material. • Sketch of some interesting epistemological features and issues that pertain STE models are shared by model organisms and Mendelian population. 8
  • 9. OUTLINE • Some super-simple examples of usually-called “population genetics models”. • Some ideas about mathematical models that were expressed in a 1989 book on modeling (Casti & Karlqvist 1989), because we may still hold such ideas when we think to mathematical models. • General pluralistic position on model notions (Leonelli 2003, 2007, Serrelli 2010, 2011). • Report of the demarcation and defense of the notion of “model organism” (Ankeny & Leonelli 2011) as opposed to “experimental organism”. • Statement of a more general notion of a model as a “stable target of explanation” (Keller 2002) => several (though not all) requirements of “model organisms” end by falling into this notion. • STE model notion describes also the essential part of population genetics, i.e. the Mendelian population => proposal of revising the semantic extension of “model”; revisiting standard distinctions like formal vs. material. • Sketch of some interesting epistemological features and issues that pertain STE models are shared by model organisms and Mendelian population. 9
  • 10. OUTLINE • Some super-simple examples of usually-called “population genetics models”. • Some ideas about mathematical models that were expressed in a 1989 book on modeling (Casti & Karlqvist 1989), because we may still hold such ideas when we think to mathematical models. • General pluralistic position on model notions (Leonelli 2003, 2007, Serrelli 2010, 2011). • Report of the demarcation and defense of the notion of “model organism” (Ankeny & Leonelli 2011) as opposed to “experimental organism”. • Statement of a more general notion of a model as a “stable target of explanation” (Keller 2002) several (though not all) requirements of “model organisms” end by falling into this notion. • STE model notion describes also the essential part of population genetics, i.e. the Mendelian population => proposal of revising the semantic extension of “model”; revisiting standard distinctions like formal vs. material. • Sketch of some interesting epistemological features and issues that pertain STE models are shared by model organisms and Mendelian population. 10
  • 11. “Population genetics models” • a, A = two alleles in a diallelic locus, in an indefinitely large population 11
  • 12. “Population genetics models” • a, A = two alleles in a diallelic locus, in an indefinitely large population • q = the frequency of allele A in the population 11
  • 13. “Population genetics models” • a, A = two alleles in a diallelic locus, in an indefinitely large population • q = the frequency of allele A in the population • [(1-q)a+qA] = 1 relative frequencies of the two alleles 11
  • 14. “Population genetics models” • a, A = two alleles in a diallelic locus, in an indefinitely large population • q = the frequency of allele A in the population • [(1-q)a+qA] = 1 relative frequencies of the two alleles • aa, Aa, AA zygotes -> what frequencies? 11
  • 15. “Population genetics models” • a, A = two alleles in a diallelic locus, in an indefinitely large population • q = the frequency of allele A in the population • [(1-q)a+qA] = 1 relative frequencies of the two alleles • aa, Aa, AA zygotes -> what frequencies? • Hardy-Weinberg equilibrium (expansion of [(1-q)a+qA]2) 11
  • 16. “Population genetics models” • a, A = two alleles in a diallelic locus, in an indefinitely large population • q = the frequency of allele A in the population • [(1-q)a+qA] = 1 relative frequencies of the two alleles • aa, Aa, AA zygotes -> what frequencies? • Hardy-Weinberg equilibrium (expansion of [(1-q)a+qA]2) 11
  • 17. “Population genetics models” • MUTATION • Δq = -uq + v(1 - q) allele A’s frequency as a function of mutation rates 12
  • 18. “Population genetics models” • MUTATION • Δq = -uq + v(1 - q) allele A’s frequency as a function of mutation rates • SELECTION • [(1-s)(1-q)a+qA]/[1-s(1-q)] = 1 relative frequencies in presence of negative selection s on a • Δq = [sq(1-q)]/[1-s(1-q)] change in the frequency of A 12
  • 19. “Population genetics models” • MUTATION • Δq = -uq + v(1 - q) allele A’s frequency as a function of mutation rates • SELECTION • [(1-s)(1-q)a+qA]/[1-s(1-q)] = 1 relative frequencies in presence of negative selection s on a • Δq = [sq(1-q)]/[1-s(1-q)] change in the frequency of A • More and more complicated equations can be built... 12
  • 20. “Population genetics models” • Work in population genetics goes on and on still today, developing those earlier ideas, as in the following example (Hartl & Clark 2007, pp. 97-98): 13
  • 21. “Population genetics models” • Work in population genetics goes on and on still today, developing those earlier ideas, as in the following example (Hartl & Clark 2007, pp. 97-98): • All this – basically, equations – is commonly referred to as “population genetics models” 13
  • 22. Ideas on mathematical models • Every model, mathematical or otherwise, is a way of representing some aspects of the real world in an abbreviated, or encapsulated, form. Mathematical models translate certain features of a natural system N into the elements of a mathematical system M, with the goal being to mirror whatever is relevant about N in the properties of M. • The […] diagram shows the two essential aspects of a mathematical model: (i) An encoding operation by which the explanatory scheme for the real-world system N is translated into the language of the formal system M, and (ii) a decoding process whereby the logical inferences in M are translated back into predictions about the temporal behavior in N. (Casti & Karlqvist 1989 p. 3). 14
  • 23. Ideas on mathematical models • Every model, mathematical or otherwise, is a way of representing some aspects of the real world in an abbreviated, or encapsulated, form. Mathematical models translate certain features of a natural system N into the elements of a mathematical system M, with the goal being to mirror whatever is relevant about N in the properties of M. • The […] diagram shows the two essential aspects of a mathematical model: (i) An encoding operation by which the explanatory scheme for the real-world system N is translated into the language of the formal system M, and (ii) a decoding process whereby the logical inferences in M are translated back into predictions about the temporal behavior in N. (Casti & Karlqvist 1989 p. 3). 14
  • 24. Ideas on mathematical models • Every model, mathematical or otherwise, is a way of representing some aspects of the real world in an abbreviated, or encapsulated, form. Mathematical models translate certain features of a natural system N into the elements of a mathematical system M, with the goal being to mirror whatever is relevant about N in the properties of M. • The […] diagram shows the two essential aspects of a mathematical model: (i) An encoding operation by which the explanatory scheme for the real-world system N is translated into the language of the formal system M, and (ii) a decoding process whereby the logical inferences in M are translated back into predictions about the temporal behavior in N. (Casti & Karlqvist 1989 p. 3). 14
  • 25. Ideas on mathematical models • Every model, mathematical or otherwise, is a way of representing some aspects of the real world in an abbreviated, or encapsulated, form. Mathematical models translate certain features of a natural system N into the elements of a mathematical system M, with the goal being to mirror whatever is relevant about N in the properties of M. • The […] diagram shows the two essential aspects of a mathematical model: (i) An encoding operation by which the explanatory scheme for the real-world system N is translated into the language of the formal system M, and (ii) a decoding process whereby the logical inferences in M are translated back into predictions about the temporal behavior in N. (Casti & Karlqvist 1989 p. 3). 15
  • 26. Ideas on mathematical models • Every model, mathematical or otherwise, is a way of representing some aspects of the real world in an abbreviated, or encapsulated, form. Mathematical models translate certain features of a natural system N into the elements of a mathematical system M, with the goal being to mirror whatever is relevant about N in the properties of M. • The […] diagram shows the two essential aspects of a mathematical model: (i) An encoding operation by which the explanatory scheme for the real-world system N is translated into the language of the formal system M, and (ii) a decoding process whereby the logical inferences in M are translated back into predictions about the temporal behavior in N. (Casti & Karlqvist 1989 p. 3). 15
  • 27. Ideas on mathematical models • An “axiom of modeling faith” holds that it is possible to bring into harmony the two worlds, i.e. the causal structure of the external world, and the inferential structure of the internal world. Moreover, such harmony is seen as the condition for a modeling relation to subsist between M and N (Rosen, pp. 16-17). 16
  • 30. Pluralism of model notions • “Every model, mathematical or otherwise...” ??? 18
  • 31. Pluralism of model notions • “Every model, mathematical or otherwise...” ??? • Leonelli (2007) defined the “single model approach” as “the tendency to explain away, rather than value and analyse, the diversity among models”. 18
  • 32. Pluralism of model notions • “Every model, mathematical or otherwise...” ??? • Leonelli (2007) defined the “single model approach” as “the tendency to explain away, rather than value and analyse, the diversity among models”. • For Leonelli, the diversity of models is scientifically important: it secures “several epistemic goals of potential interest to practicing scientists”, and it allows biologists to combine them in order to pursue their research outcomes. 18
  • 33. Pluralism of model notions • “Every model, mathematical or otherwise...” ??? • Leonelli (2007) defined the “single model approach” as “the tendency to explain away, rather than value and analyse, the diversity among models”. • For Leonelli, the diversity of models is scientifically important: it secures “several epistemic goals of potential interest to practicing scientists”, and it allows biologists to combine them in order to pursue their research outcomes. • Should we be permanently content of grouping heterogeneous activities under the single term “modeling”? 18
  • 34. Pluralism of model notions • “Every model, mathematical or otherwise...” ??? • Leonelli (2007) defined the “single model approach” as “the tendency to explain away, rather than value and analyse, the diversity among models”. • For Leonelli, the diversity of models is scientifically important: it secures “several epistemic goals of potential interest to practicing scientists”, and it allows biologists to combine them in order to pursue their research outcomes. • Should we be permanently content of grouping heterogeneous activities under the single term “modeling”? • Surely a pluralistic account is the best thing we can do for now. 18
  • 35. Model organisms • Demarcating the concept “model organisms” vs. “experimental organisms” (Ankeny & Leonelli 2011). 19
  • 36. Model organisms • Demarcating the concept “model organisms” vs. “experimental organisms” (Ankeny & Leonelli 2011). • Model organisms are non-human species that are extensively studied in order to understand a range of biological phenomena, with the hope that data and theories generated through use of the model will be applicable to other organisms, particularly those that are in some way more complex than the original model (p. 313). 19
  • 37. Model organisms • Demarcating the concept “model organisms” vs. “experimental organisms” (Ankeny & Leonelli 2011). • Model organisms are non-human species that are extensively studied in order to understand a range of biological phenomena, with the hope that data and theories generated through use of the model will be applicable to other organisms, particularly those that are in some way more complex than the original model (p. 313). • Model organisms can be clearly distinguished from the broader class of experimental organisms by several features. 19
  • 38. Model organisms • My view: several features identified by Ankeny & Leonelli fall into a more general category - not experimental organism, but “stable target of explanation”. 20
  • 39. Model organisms • My view: several features identified by Ankeny & Leonelli fall into a more general category - not experimental organism, but “stable target of explanation”. • Two exclusive features of model organisms: • Material features • Representational target 20
  • 40. Model organisms • My view: several features identified by Ankeny & Leonelli fall into a more general category - not experimental organism, but “stable target of explanation”. • Two exclusive features of model organisms: to become (and remain) model, organisms have to be suitable to be brood and tamed cost-effectively; the “wild type strain” has to • Material features be isolated and standardized, so to assure the comparability of results across a large research community, etc. • Representational target 20
  • 41. Model organisms • My view: several features identified by Ankeny & Leonelli fall into a more general category - not experimental organism, but “stable target of explanation”. • Two exclusive features of model organisms: • Material features • Representational target 21
  • 42. Model organisms • My view: several features identified by Ankeny & Leonelli fall into a more differs from representational general category - not experimental organism, but “stable target of scope explanation”. describes the conceptual reasons • Two exclusive features of model organisms: why researchers are studying a • Material features • Representational target 21
  • 43. Model organisms • My view: several features identified by Ankeny & Leonelli fall into a more whole, intact organisms general category - not experimental organism, but “stable target of explanation”. …model organisms […] involve attempts to generate complete knowledge of the fundamental • Two exclusive features of model organisms:processes at work […] including the molecular, cellular, and developmental processes; in this • Material features sense the model organism is understood as a test tube for achieving a full understanding of • Representational target all biological processes (p. 317). 22
  • 44. Stable target of explanation (STE) 23
  • 45. Stable target of explanation (STE) • ...[model in experimental biology] is an organism, an organism that can be taken to represent (that is, stand in for) a class of organisms. A model in this sense is not expected to serve an explanatory function in itself, nor is it a simplified representation of a more complex phenomenon for which we already have explanatory handles. Rather, its primary function is to provide simply a stable target of explanation (Keller 2002, p. 115). 24
  • 46. Stable target of explanation (STE) 1. targets of explanation: not immediately tools for explaining { 25
  • 47. Stable target of explanation (STE) 1. targets of explanation: not immediately tools for explaining 2. autonomous: from theory and from data (Morgan & Morrison 1999) { 25
  • 48. Stable target of explanation (STE) 1. targets of explanation: not immediately tools for explaining 2. autonomous: from theory and from data (Morgan & Morrison 1999) 3. stable: answers to the challenge of producing lawlike knowledge in fields such as experimental biology (Creager et al. 2007, p. 5) { 25
  • 49. Stable target of explanation (STE) 1. targets of explanation: not immediately tools for explaining 2. autonomous: from theory and from data (Morgan & Morrison 1999) 3. stable: answers to the challenge of producing lawlike knowledge in fields such as experimental biology (Creager et al. 2007, p. 5) { 4. representational scope 25
  • 50. Stable target of explanation (STE) 1. targets of explanation: not immediately tools for explaining 2. autonomous: from theory and from data (Morgan & Morrison 1999) 3. stable: answers to the challenge of producing lawlike knowledge in fields such as experimental biology (Creager et al. 2007, p. 5) { 4. representational scope how extensively the results […] can be projected onto a wider group of organisms ...the extent to which researchers see their findings as applicable across organisms (Ankeny & Leonelli, p. 315). 25
  • 51. Stable target of explanation (STE) 1. targets of explanation: not immediately tools for explaining 2. autonomous: from theory and from data (Morgan & Morrison 1999) 3. stable: answers to the challenge of producing lawlike knowledge in fields such as experimental biology (Creager et al. 2007, p. 5) { 4. representational scope is tendentially wide and changeable 26
  • 52. Stable target of explanation (STE) 1. targets of explanation: not immediately tools for explaining 2. autonomous: from theory and from data (Morgan & Morrison 1999) 3. stable: answers to the challenge of producing lawlike knowledge in fields such as experimental biology (Creager et al. 2007, p. 5) { 4. representational scope is tendentially wide and changeable 5. unified research community: ethos of sharing reinforces stability 26
  • 53. Stable target of explanation (STE) 1. targets of explanation: not immediately tools for explaining 2. autonomous: from theory and from data (Morgan & Morrison 1999) 3. stable: answers to the challenge of producing lawlike knowledge in fields such as experimental biology (Creager et al. 2007, p. 5) { 4. representational scope is tendentially wide and changeable 5. unified research community: ethos of sharing reinforces stability 6. socio-technical features: associated “experimental resources”, standardization, comparability, cumulative establishment of techniques, practices, and results 26
  • 54. Stable target of explanation (STE) 1. targets of explanation: not immediately tools for explaining 2. autonomous: from theory and from data (Morgan & Morrison 1999) 3. stable: answers to the challenge of producing lawlike knowledge in fields such as experimental biology (Creager et al. 2007, p. 5) { 4. representational scope is tendentially wide and changeable 5. unified research community: ethos of sharing reinforces stability 6. socio-technical features: associated “experimental resources”, standardization, comparability, cumulative establishment of techniques, practices, and results 7. artificiality: even model organisms “have been developed using complex processes of standardization that allow the establishment of a standard strain which then serves as the basis for future research” (Ankeny & Leonelli, p. 316, cf. e.g. Clarke & Fujimura 1992) 26
  • 55. Stable target of explanation (STE) 1. targets of explanation: not immediately tools for explaining 2. autonomous: from theory and from data (Morgan & Morrison 1999) 3. stable: answers to the challenge of producing lawlike knowledge in fields such as experimental biology (Creager et al. 2007, p. 5) { 4. representational scope is tendentially wide and changeable 5. unified research community: ethos of sharing reinforces stability 6. socio-technical features: associated “experimental resources”, standardization, comparability, cumulative establishment of techniques, practices, and results 7. artificiality: even model organisms “have been developed using complex processes of standardization that allow the establishment of a standard strain which then serves as the basis for future research” (Ankeny & Leonelli, p. 316, cf. e.g. Clarke & Fujimura 1992) 8. inexhaustedness: “…although model organisms are standardized in order to facilitate highly controlled biological experimentation, their inherent complexity means that the systems are never fully understood and can continue to generate surprising results” (Creager et al. 2007, p. 7) 26
  • 56. How STE applies to population genetics • To understand how the STE model notion applies to mathematical population genetics, we have to move our focus away from equations, and recognize that there is another, more fundamental object: the Mendelian population. • i.e., the large-scale derivation of Mendel’s rules of inheritance. 27
  • 57. How STE applies to population genetics • To understand how the STE model notion applies to mathematical population genetics, we have to move our focus away from equations, and recognize that there is another, more fundamental object: the Mendelian population. • i.e., the large-scale derivation of Mendel’s rules of inheritance. 27
  • 58. How STE applies to population genetics • To understand how the STE model notion applies to mathematical population genetics, we have to move our focus away from equations, and recognize that there is another, more fundamental object: the Mendelian population. • i.e., the large-scale derivation of Mendel’s rules of inheritance. 27
  • 59. How STE applies to population genetics • Mendelian population is the space of all possible individual combinations given a number of loci and a number of alleles => a combination space • alleles : : : • loci • individual combinations 28
  • 60. How STE applies to population genetics • Mendelian population is the space of all possible individual combinations given a number of loci and a number of alleles => a combination space • alleles : : : • loci • individual combinations • What is the relationship between this space and population genetics equations? An epistemological gap! 28
  • 61. How STE applies to population genetics • Only some positions in the combination space are actually occupied at a certain time. Which combinations are realized? 29
  • 62. How STE applies to population genetics • Only some positions in the combination space are actually occupied at a certain time. Which combinations are realized? • With a minimally realistic number of loci and alleles, the dimensionality of this space is so high that no equation or algorithm can be developed. 29
  • 63. How STE applies to population genetics • Only some positions in the combination space are actually occupied at a certain time. Which combinations are realized? • With a minimally realistic number of loci and alleles, the dimensionality of this space is so high that no equation or algorithm can be developed. • Statistical equations address what happens to the allele frequencies in one or two loci in a population inhabiting an oversimplified di-allelic Mendelian population space. 29
  • 64. How STE applies to population genetics • Only some positions in the combination space are actually occupied at a certain time. Which combinations are realized? • With a minimally realistic number of loci and alleles, the dimensionality of this space is so high that no equation or algorithm can be developed. • Statistical equations address what happens to the allele frequencies in one or two loci in a population inhabiting an oversimplified di-allelic Mendelian population space. • Even the more complicated population genetics equations are incredibly partial statistical studies of the Mendelian population space. 29
  • 65. How STE applies to population genetics • The Mendelian population (a combination space) is, in my view, it is the model of population genetics (at least of Mendelian population genetics), it is the space equations are about. 30
  • 66. How STE applies to population genetics • Review and apply STE features: 1. target of explanation 2. autonomous 3. stable 4. representational scope 5. unified research community 6. socio-technical features 7. artificiality 8. inexhaustedness 31
  • 67. How STE applies to population genetics • Review and apply STE features: 1. target of explanation 2. autonomous 3. stable 4. representational scope 5. unified research community 6. socio-technical features 7. artificiality the “connectedness” structure of Mendelian population is very 8. inexhaustedness different from what we always imagined intuitively 32
  • 68. How STE applies to population genetics • Review and apply STE features: this discovery enlarges the representational scope of 1. target of explanation Mendelian population: from adaptation to speciation too 2. autonomous 3. stable 4. representational scope 5. unified research community 6. socio-technical features 7. artificiality the “connectedness” among genotypes in Mendelian population 8. inexhaustedness is very different from what we always imagined intuitively 33
  • 69. Issues about STE model notion • Epistemological questions (dilemmas?). If, as several authors point out (e.g., Creager et al. 2007), models are not chosen because they are typical of a certain set of systems, nor they are built to represent some other system by reduction, deduction, encoding (Casti & Karlqvist 1989, Rosen 1989) or the like, how can they... • REPRESENT? • EXPLAIN? • PREDICT? 34
  • 70. Issues about STE model notion • Discussing such relationships is not essential within a notion of a model as a stable target of explanation. That is, if we choose this notion of model we can provisionally remain silent on how and what the model represents and explains. 35
  • 71. Issues about STE model notion • Discussing such relationships is not essential within a notion of a model as a stable target of explanation. That is, if we choose this notion of model we can provisionally remain silent on how and what the model represents and explains. • The most notable fact is that all the issues Formal Material are shared between a formal and a material models. 35
  • 72. Issues about STE model notion • Discussing such relationships is not essential within a notion of a model as a stable target of explanation. That is, if we choose this notion of model we can provisionally remain silent on how and what the model represents and explains. • The most notable fact is that all the issues Formal Material are shared between a formal and a material models. 35

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