Fabrizio Panebianco and Emanuele Serrelli: A Niche Construction Model with Reaction Norms

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Overview Niche Construction Plasticity Reactions Norm NCRN: Basic and Population NCRN and Economics Formalization A Niche Construction Model with Reaction Norms Fabrizio Panebianco and Emanuele Serrelli University of Milano - Bicocca May 6, 2011 CISEPS ANNUAL REPORT

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Short communication atCISEPS Annual LectureMay 6, 9:30am, University of Milano Bicocca, Milano (Italy), building U12, Also with:- Luca Cavalli Sforza "Critical periods of human evolution: interdisciplinarity helps for understanding"- Marcus Feldman, "On Models of Social Transmission: Rates of Evolution and Patterns of Diversity".

Transcript of Fabrizio Panebianco and Emanuele Serrelli: A Niche Construction Model with Reaction Norms

Page 1: Fabrizio Panebianco and Emanuele Serrelli: A Niche Construction Model with Reaction Norms

Overview Niche Construction Plasticity Reactions Norm NCRN: Basic and Population NCRN and Economics Formalization

A Niche Construction Model with ReactionNorms

Fabrizio Panebianco and Emanuele Serrelli

University of Milano - Bicocca

May 6, 2011

CISEPS ANNUAL REPORT

Page 2: Fabrizio Panebianco and Emanuele Serrelli: A Niche Construction Model with Reaction Norms

Overview Niche Construction Plasticity Reactions Norm NCRN: Basic and Population NCRN and Economics Formalization

Cultural Evolution Project

Page 3: Fabrizio Panebianco and Emanuele Serrelli: A Niche Construction Model with Reaction Norms

Overview Niche Construction Plasticity Reactions Norm NCRN: Basic and Population NCRN and Economics Formalization

Outline

• Niche construction• as a neglected major biological process• as modification and innovation of existing models

• Something missing in NC models: phenotypic plasticity

• NCRN model: a niche construction model with reactionnorm

• Basic and population NCRN Models

• NCRN, cultural evolution, and economical modeling

• First Formalization

Page 4: Fabrizio Panebianco and Emanuele Serrelli: A Niche Construction Model with Reaction Norms

Overview Niche Construction Plasticity Reactions Norm NCRN: Basic and Population NCRN and Economics Formalization

Niche Construction: «the selection pressures to which anorganism is exposed exist partly as a consequence of the nicheconstructing activities of past and present generations oforganisms»(Laland et al. (1996), p.294)

Page 5: Fabrizio Panebianco and Emanuele Serrelli: A Niche Construction Model with Reaction Norms

Overview Niche Construction Plasticity Reactions Norm NCRN: Basic and Population NCRN and Economics Formalization

Page 6: Fabrizio Panebianco and Emanuele Serrelli: A Niche Construction Model with Reaction Norms

Overview Niche Construction Plasticity Reactions Norm NCRN: Basic and Population NCRN and Economics Formalization

Page 7: Fabrizio Panebianco and Emanuele Serrelli: A Niche Construction Model with Reaction Norms

Overview Niche Construction Plasticity Reactions Norm NCRN: Basic and Population NCRN and Economics Formalization

That the activities of organisms lead to environmental modifications, on

both on a local and global scale, can hardly be doubted; so much is

obvious to even the most casual observer of the natural world. . . . if

everyone agrees that niche-construction, i.e. organism-induced

modification of the environment, occurs, why does orthodox evolutionary

theory fail to take account of it? (Okasha 2005, p. 1).

Page 8: Fabrizio Panebianco and Emanuele Serrelli: A Niche Construction Model with Reaction Norms

Overview Niche Construction Plasticity Reactions Norm NCRN: Basic and Population NCRN and Economics Formalization

• Laland KN, Odling-Smee FJ, Feldman MW (1996). On theevolutionary consequences of niche construction. Journal ofEvolutionary Biology 9: 293-316.

• Laland KN, Odling-Smee FJ, Feldman MW (1999). Theevolutionary consequences of niche construction and its implicationsfor ecology. Proceedings of the National Academy of Sciences USA96:10242- 47.

• NC models are modification of multiplicative model (two loci)

• Locus E (alleles E, e), Locus A (alleles A, a), Resources R

• Frequency in E influences R, R influences A’s contribution to fitness

The analysis confirms that niche construction can be a potent

evolutionary agent by generating selection that leads to the fixation of

otherwise deleterious alleles, supporting stable polymorphisms where none

are expected, eliminating what would otherwise be stable polymorphisms,

and generating unusual evolutionary dynamics. Even small amounts of

niche construction, or niche construction that only weakly affects

resource dynamics, can significantly alter both ecological and

evolutionary patterns. (Laland et al. 1999, p. 10242).

Page 9: Fabrizio Panebianco and Emanuele Serrelli: A Niche Construction Model with Reaction Norms

Overview Niche Construction Plasticity Reactions Norm NCRN: Basic and Population NCRN and Economics Formalization

• Laland KN, Odling-Smee FJ, Feldman MW (1996). On theevolutionary consequences of niche construction. Journal ofEvolutionary Biology 9: 293-316.

• Laland KN, Odling-Smee FJ, Feldman MW (1999). Theevolutionary consequences of niche construction and its implicationsfor ecology. Proceedings of the National Academy of Sciences USA96:10242- 47.

• NC models are modification of multiplicative model (two loci)

• Locus E (alleles E, e), Locus A (alleles A, a), Resources R

• Frequency in E influences R, R influences A’s contribution to fitness

The analysis confirms that niche construction can be a potent

evolutionary agent by generating selection that leads to the fixation of

otherwise deleterious alleles, supporting stable polymorphisms where none

are expected, eliminating what would otherwise be stable polymorphisms,

and generating unusual evolutionary dynamics. Even small amounts of

niche construction, or niche construction that only weakly affects

resource dynamics, can significantly alter both ecological and

evolutionary patterns. (Laland et al. 1999, p. 10242).

Page 10: Fabrizio Panebianco and Emanuele Serrelli: A Niche Construction Model with Reaction Norms

Overview Niche Construction Plasticity Reactions Norm NCRN: Basic and Population NCRN and Economics Formalization

• Laland KN, Odling-Smee FJ, Feldman MW (1996). On theevolutionary consequences of niche construction. Journal ofEvolutionary Biology 9: 293-316.

• Laland KN, Odling-Smee FJ, Feldman MW (1999). Theevolutionary consequences of niche construction and its implicationsfor ecology. Proceedings of the National Academy of Sciences USA96:10242- 47.

• NC models are modification of multiplicative model (two loci)

• Locus E (alleles E, e), Locus A (alleles A, a), Resources R

• Frequency in E influences R, R influences A’s contribution to fitness

The analysis confirms that niche construction can be a potent

evolutionary agent by generating selection that leads to the fixation of

otherwise deleterious alleles, supporting stable polymorphisms where none

are expected, eliminating what would otherwise be stable polymorphisms,

and generating unusual evolutionary dynamics. Even small amounts of

niche construction, or niche construction that only weakly affects

resource dynamics, can significantly alter both ecological and

evolutionary patterns. (Laland et al. 1999, p. 10242).

Page 11: Fabrizio Panebianco and Emanuele Serrelli: A Niche Construction Model with Reaction Norms

Overview Niche Construction Plasticity Reactions Norm NCRN: Basic and Population NCRN and Economics Formalization

Phenotypic Plasticity

• The term phenotypic plasticity describes the property of agenotype to generate phenotypic variation in response to agiven range of environmental conditions.

• Various measures of plasticity exist and can be united withinthe framework of a polynomial function: the norm ofreaction.

• NC models do not allow factors other than genotypes toparticipate in the production of phenotypes.

• Niche construction advocates always consider the classic ideaof an influence on the population’s gene pool (frequency ofalleles that correspond to determinate phenotypic traits orbehaviours).

Page 12: Fabrizio Panebianco and Emanuele Serrelli: A Niche Construction Model with Reaction Norms

Overview Niche Construction Plasticity Reactions Norm NCRN: Basic and Population NCRN and Economics Formalization

Phenotypic Plasticity

• The term phenotypic plasticity describes the property of agenotype to generate phenotypic variation in response to agiven range of environmental conditions.

• Various measures of plasticity exist and can be united withinthe framework of a polynomial function: the norm ofreaction.

• NC models do not allow factors other than genotypes toparticipate in the production of phenotypes.

• Niche construction advocates always consider the classic ideaof an influence on the population’s gene pool (frequency ofalleles that correspond to determinate phenotypic traits orbehaviours).

Page 13: Fabrizio Panebianco and Emanuele Serrelli: A Niche Construction Model with Reaction Norms

Overview Niche Construction Plasticity Reactions Norm NCRN: Basic and Population NCRN and Economics Formalization

Phenotypic Plasticity

• The term phenotypic plasticity describes the property of agenotype to generate phenotypic variation in response to agiven range of environmental conditions.

• Various measures of plasticity exist and can be united withinthe framework of a polynomial function: the norm ofreaction.

• NC models do not allow factors other than genotypes toparticipate in the production of phenotypes.

• Niche construction advocates always consider the classic ideaof an influence on the population’s gene pool (frequency ofalleles that correspond to determinate phenotypic traits orbehaviours).

Page 14: Fabrizio Panebianco and Emanuele Serrelli: A Niche Construction Model with Reaction Norms

Overview Niche Construction Plasticity Reactions Norm NCRN: Basic and Population NCRN and Economics Formalization

NC vc NCRN• Reaction Norm (G)≡ The rule imposed by the genotype,

that determines the phenotype expressed in the population asa function of the environment and other factors, can beintended in different ways: it can go from simple mechanicalreaction rules up to complex algorithms considering aspects ofthe environment, or from population either seen in the presentor preceding generation.

• Differences: Phenotypic outcome affects the environment.Current phenotype undergoes selection, which affects onpopulation size. Future phenotypic outcomes depend on theenvironment by reaction norm, which takes into account atleast the resource available in the environment and populationsize.

• Common Feature: Aggregate Effect. NC models areconstitutively concerned with selection pressures, and whileselection pressures act on individuals, sorting them throughgenerations, they make sense only at population level.

Page 15: Fabrizio Panebianco and Emanuele Serrelli: A Niche Construction Model with Reaction Norms

Overview Niche Construction Plasticity Reactions Norm NCRN: Basic and Population NCRN and Economics Formalization

NC vc NCRN• Reaction Norm (G)≡ The rule imposed by the genotype,

that determines the phenotype expressed in the population asa function of the environment and other factors, can beintended in different ways: it can go from simple mechanicalreaction rules up to complex algorithms considering aspects ofthe environment, or from population either seen in the presentor preceding generation.

• Differences: Phenotypic outcome affects the environment.Current phenotype undergoes selection, which affects onpopulation size. Future phenotypic outcomes depend on theenvironment by reaction norm, which takes into account atleast the resource available in the environment and populationsize.

• Common Feature: Aggregate Effect. NC models areconstitutively concerned with selection pressures, and whileselection pressures act on individuals, sorting them throughgenerations, they make sense only at population level.

Page 16: Fabrizio Panebianco and Emanuele Serrelli: A Niche Construction Model with Reaction Norms

Overview Niche Construction Plasticity Reactions Norm NCRN: Basic and Population NCRN and Economics Formalization

NC vc NCRN• Reaction Norm (G)≡ The rule imposed by the genotype,

that determines the phenotype expressed in the population asa function of the environment and other factors, can beintended in different ways: it can go from simple mechanicalreaction rules up to complex algorithms considering aspects ofthe environment, or from population either seen in the presentor preceding generation.

• Differences: Phenotypic outcome affects the environment.Current phenotype undergoes selection, which affects onpopulation size. Future phenotypic outcomes depend on theenvironment by reaction norm, which takes into account atleast the resource available in the environment and populationsize.

• Common Feature: Aggregate Effect. NC models areconstitutively concerned with selection pressures, and whileselection pressures act on individuals, sorting them throughgenerations, they make sense only at population level.

Page 17: Fabrizio Panebianco and Emanuele Serrelli: A Niche Construction Model with Reaction Norms

Overview Niche Construction Plasticity Reactions Norm NCRN: Basic and Population NCRN and Economics Formalization

Basic and Population NCRN Models

• The work with a basic NCRN model would be focused on theexploration of the dynamics resulting from different kinds ofreaction norms with no variation of G across the population.

• Although basic NCRN studies seem insensitive to theparticular explanation for the ubiquity of G, such studiesmight yield some results about the kinds of evolutionarynorms that exhibit evolutionary stability. NCRN studies canbe thought as preparatory to population models with multiplereaction norms (e.g. G1 and G2).

Page 18: Fabrizio Panebianco and Emanuele Serrelli: A Niche Construction Model with Reaction Norms

Overview Niche Construction Plasticity Reactions Norm NCRN: Basic and Population NCRN and Economics Formalization

Page 19: Fabrizio Panebianco and Emanuele Serrelli: A Niche Construction Model with Reaction Norms

Overview Niche Construction Plasticity Reactions Norm NCRN: Basic and Population NCRN and Economics Formalization

Page 20: Fabrizio Panebianco and Emanuele Serrelli: A Niche Construction Model with Reaction Norms

Overview Niche Construction Plasticity Reactions Norm NCRN: Basic and Population NCRN and Economics Formalization

NCRN, cultural evolution, and economical modeling

• Reaction norm and Rational Choice are both Rules thatshare the picking of an element from a set of possibilitiesaccording to external constraints.

• This picked up element is respectively the phenotype and thechosen action.

• NCRN models introduce NC in a way that closes a gapbetween biological and economic modeling.

vs

• NC models traits are cultural and genetic, and both kinds aremodeled by means of loci and alleles, where alleles correspondto phenotypic alternatives that are fixed and there is no roomfor plasticity.

Page 21: Fabrizio Panebianco and Emanuele Serrelli: A Niche Construction Model with Reaction Norms

Overview Niche Construction Plasticity Reactions Norm NCRN: Basic and Population NCRN and Economics Formalization

NCRN, cultural evolution, and economical modeling

• Reaction norm and Rational Choice are both Rules thatshare the picking of an element from a set of possibilitiesaccording to external constraints.

• This picked up element is respectively the phenotype and thechosen action.

• NCRN models introduce NC in a way that closes a gapbetween biological and economic modeling.

vs

• NC models traits are cultural and genetic, and both kinds aremodeled by means of loci and alleles, where alleles correspondto phenotypic alternatives that are fixed and there is no roomfor plasticity.

Page 22: Fabrizio Panebianco and Emanuele Serrelli: A Niche Construction Model with Reaction Norms

Overview Niche Construction Plasticity Reactions Norm NCRN: Basic and Population NCRN and Economics Formalization

NCRN, cultural evolution, and economical modeling

• Reaction norm and Rational Choice are both Rules thatshare the picking of an element from a set of possibilitiesaccording to external constraints.

• This picked up element is respectively the phenotype and thechosen action.

• NCRN models introduce NC in a way that closes a gapbetween biological and economic modeling.

vs

• NC models traits are cultural and genetic, and both kinds aremodeled by means of loci and alleles, where alleles correspondto phenotypic alternatives that are fixed and there is no roomfor plasticity.

Page 23: Fabrizio Panebianco and Emanuele Serrelli: A Niche Construction Model with Reaction Norms

Overview Niche Construction Plasticity Reactions Norm NCRN: Basic and Population NCRN and Economics Formalization

First Formalization

• Resources Rt ,

• Population mass mt ,

• Genotype G , (reaction norm)

• Phenotype At ,

At : (Rt , mt , G ) 7→ [0, Rt ]Bt : (mt , At) 7→ <+

Ct : (Rt , At) 7→ <+

Ft : (Bt − Ct) 7→ <+

Rt : (Rt−1, At−1) 7→ <+

mt : (mt−1, Ft−1) 7→ <+

(1)