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Book club


             Andreas Wagner,
   The Origins of Evolutionary Innovations


                 Chapter 1

Book club presented by G. Dall'Olio, IBE-CEXS
Contents of Chapter 1
1)What is an “Evolutionary Innovation”?
2)Definition of “Genotype Space” and “Genotype 
   Network”
Evolutionary Innovations
   An Evolutionary Innovation is a new trait that 
     introduces something “revolutionary” in 
     evolution
   The definition is quite broad... think that any 
     phenotype has been an evolutionary innovation 
     when it first appeared
   Let's see some notable examples 
Examples of Evolutionary
          Innovation
   In Metabolic networks:
          Microbes that evolve the ability to metabolize 
            xenobiotics (waste compounds produced by 
            humans)
          Urea Cycle
          Oxygen as an electron acceptor 
Examples of Evolutionary
          Innovation
   In Regulatory networks:
          Eye­like spots on butterflies' wings (Distal­less gene)
          Lenses of marine animals' eyes
          Plants' leaves (Knox genes)
Examples of Evolutionary
          Innovation
   As novel molecules:
          Enzymes that obtain the ability to catalyze 
            completely different reactions after a mutation
                   E.coli's L­Ru5P, that acquired aldolase activity after a 
                     mutation
                   IDH (citric cycle) and IMDH (leucine synthesis)
          Evolution of anti­freeze proteins
Examples of Evolutionary
          Innovation
   Paper published today (Jan 27th 2012)
   A phage evolved the ability to infect a novel strain 
     of E.coli
Evolutionary Innovations
             resume
   In general, the aim of this book is to describe how 
      novel phenotypes are discovered
Contents of Chapter 1
1)What is an “Evolutionary Innovation”?
2)Definition of “Genotype Space” and “Genotype 
   Network” 
Definition of
              Genotype Space
   The genotype space is the set of all possible 
     genotypes
   Let's represent it as a matrix where two neighbor 
     genotypes differ only for one position:
        AAAAA    AAAAC    AAAAG   AAAAT    AAATT
        AAACA    AAACC    AAACG   AAACT    AAATC
        AACCA    AACCC    AACCG   AACCT    …..
        ACCCA    ACCCC    ACCCG   ACCCT    …..
        CCCCA    CCCCC    CCCCG   CCCCT    …..
        …..      …..      …..     …..      …..
Example of Genotype Space
   The genotype space is the set of all possible 
     genotypes
   Let's represent it as a matrix where two neighbor 
     genotypes differ only for one position:
                                                     Only one
        AAAAA    AAAAC    AAAAG   AAAAT    AAATT     difference
                                                     between
        AAACA    AAACC    AAACG   AAACT    AAATC     neighbor
        AACCA    AACCC    AACCG   AACCT    …..       points

        ACCCA    ACCCC    ACCCG   ACCCT    …..
        CCCCA    CCCCC    CCCCG   CCCCT    …..
        …..      …..      …..     …..      …..
Example of Genotype Space
   The genotype space is the set of all possible 
     genotypes
   Let's represent it as a matrix where two neighbor 
     genotypes differ only for one position:
                                                                      Only one
        AAAAA    AAAAC        AAAAG       AAAAT        AAATT          difference
                                                                      between
        AAACA    AAACC        AAACG       AAACT        AAATC          neighbor
        AACCA    AACCC        AACCG       AACCT        …..            points

        ACCCA    ACCCC        ACCCG       ACCCT        …..
        CCCCA    CCCCC        CCCCG       CCCCT        …..
        …..      …..          …..         …..          …..

                 Here, genotypes are represented as sequences, but they
                 can be other things (will be discussed later)
Genotype network
   A genotype network is a set of genotypes that have the same 
      phenotype, and are connected by single pairwise differences
   Let's assume that the marked genotypes have the same 
      phenotype:
         AAAAA     AAAAC     AAAAG     AAAAT     AAATT
         AAACA     AAACC     AAACG     AAACT     AAATC
         AACCA     AACCC     AACCG     AACCT     …..
         ACCCA     ACCCC     ACCCG     ACCCT     …..
         CCCCA     CCCCC     CCCCG     CCCCT     …..
         …..       …..       …..       …..       …..


   → Yellow genotypes represent a genotype network
Genotype networks
              resume
   The concept of Genotype networks allows us to 
     study how much a genotype can vary, without 
     changing the phenotype
   This is important to get to the final aim of this 
     book: understand how new innovative phenotypes 
     are discovered
Exploring genotype
              networks
        AAAAA   AAAAC   AAAAG   AAAAT    AAATT
        AAACA   AAACC   AAACG   AAACT    AAATC
        AACCA   AACCC   AACCG   AACCT    …..
        ACCCA   ACCCC   ACCCG   ACCCT    …..
        CCCCA   CCCCC   CCCCG   CCCCT    …..
        …..     …..     …..     …..      …..

   How big can a genotype network be?
   How can a population explore a genotype network?
   ….. many answers in the next chapters of the book
Exploring Genotype
               networks
   AAAAA and ACCCT have the same phenotype (they are in the 
     same genotype network)
   Do their neighbors (e.g. AAACA and CCCT) have similar 
      phenotype?
         AAAAA    AAAAC    AAAAG     AAAAT    AAATT
         AAACA    AAACC    AAACG     AAACT    AAATC
         AACCA    AACCC    AACCG     AACCT    …..
         ACCCA    ACCCC    ACCCG     ACCCT    …..
         CCCCA    CCCCC    CCCCG     CCCCT    …..
         …..      …..      …..       …..      …..
Example of Genotype
     network taken from the
         book (fig. 2.6)
   The lines corresponds to genotypes in a genotype network
   G1 and G2 have the same phenotype
   White spaces correspond to genotypes that don't have the 
     phenotype analyzed in this genotype network
Extending the definition of
       “genotype”
   Depending what we want to study, we can use 
     different definitions of “genotype” and 
     “phenotype”
   For example, in a metabolic network, the genotype 
     can be the set of reactions that an organism can 
     catalize  
Example of alternative
  definition of “genotype”
MGAT1 functional       MGAT1 functional       MGAT1 functional       MGAT1 functional
MGAT2 functional       MGAT2 not functional   MGAT2 not functional   MGAT2 not functional
MGAT3 functional       MGAT3 functional       MGAT3 not functional   MGAT3 not functional
MGAT4 functional       MGAT4 functional       MGAT4 functional       MGAT4 not functional
MGAT5 functional       MGAT5 functional       MGAT5 functional       MGAT5 functional
MGAT1 not functional   MGAT1 not functional   MGAT1 not functional   MGAT1 not functional
MGAT2 functional       MGAT2 not functional   MGAT2 not functional   MGAT2 not functional
MGAT3 functional       MGAT3 functional       MGAT3 not functional   MGAT3 not functional
MGAT4 functional       MGAT4 functional       MGAT4 functional       MGAT4 not functional
MGAT5 functional       MGAT5 functional       MGAT5 functional       MGAT5 functional
MGAT1 not functional   MGAT1 not functional   MGAT1 not functional   ….
MGAT2 not functional   MGAT2 not functional   MGAT2 not functional
MGAT3 functional       MGAT3 functional       MGAT3 functional
MGAT4 functional       MGAT4 not functional   MGAT4 not functional
MGAT5 functional       MGAT5 functional       MGAT5 functional
MGAT1 not functional   MGAT1 functional       MGAT1 not functional   …..
MGAT2 not functional   MGAT2 not functional   MGAT2 not functional
MGAT3 not functional   MGAT3 not functional   MGAT3 not functional
MGAT4 functional       MGAT4 functional       MGAT4 functional
MGAT5 functional       MGAT5 functional       MGAT5 not functional
Example of alternative
      definition of “genotype”
    All the yellow cells have the same phenotype (e.g. 
      they can produce glycosylation)
    MGAT1 functional       MGAT1 functional       MGAT1 functional       MGAT1 functional
    MGAT2 functional       MGAT2 not functional   MGAT2 not functional   MGAT2 not functional
    MGAT3 functional       MGAT3 functional       MGAT3 not functional   MGAT3 not functional
    MGAT4 functional       MGAT4 functional       MGAT4 functional       MGAT4 not functional
    MGAT5 functional       MGAT5 functional       MGAT5 functional       MGAT5 functional
    MGAT1 not functional   MGAT1 not functional   MGAT1 not functional   MGAT1 not functional
    MGAT2 functional       MGAT2 not functional   MGAT2 not functional   MGAT2 not functional
    MGAT3 functional       MGAT3 functional       MGAT3 not functional   MGAT3 not functional
    MGAT4 functional       MGAT4 functional       MGAT4 functional       MGAT4 not functional
    MGAT5 functional       MGAT5 functional       MGAT5 functional       MGAT5 functional
    MGAT1 not functional   MGAT1 not functional   MGAT1 not functional   ….
    MGAT2 not functional   MGAT2 not functional   MGAT2 not functional
    MGAT3 functional       MGAT3 functional       MGAT3 functional
    MGAT4 functional       MGAT4 not functional   MGAT4 not functional
    MGAT5 functional       MGAT5 functional       MGAT5 functional
    MGAT1 not functional   MGAT1 functional       MGAT1 not functional   …..
    MGAT2 not functional   MGAT2 not functional   MGAT2 not functional
    MGAT3 not functional   MGAT3 not functional   MGAT3 not functional
    MGAT4 functional       MGAT4 functional       MGAT4 functional
    MGAT5 functional       MGAT5 functional       MGAT5 not functional
Take home messages
   Genotype network != biological pathways
   Genotype network: set of possible genotypes 
     sharing the same phenotype, and connected
   Evolutionary Innovation: any novel phenotype
   Theory of Innovation: studies how populations can 
     explore the genotype space, the properties of 
     genotype networks, and how innovative 
     phenotypes can be found

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Wagner chapter 1

  • 1. Book club Andreas Wagner, The Origins of Evolutionary Innovations Chapter 1 Book club presented by G. Dall'Olio, IBE-CEXS
  • 2. Contents of Chapter 1 1)What is an “Evolutionary Innovation”? 2)Definition of “Genotype Space” and “Genotype  Network”
  • 3. Evolutionary Innovations  An Evolutionary Innovation is a new trait that  introduces something “revolutionary” in  evolution  The definition is quite broad... think that any  phenotype has been an evolutionary innovation  when it first appeared  Let's see some notable examples 
  • 4. Examples of Evolutionary Innovation  In Metabolic networks:  Microbes that evolve the ability to metabolize  xenobiotics (waste compounds produced by  humans)  Urea Cycle  Oxygen as an electron acceptor 
  • 5. Examples of Evolutionary Innovation  In Regulatory networks:  Eye­like spots on butterflies' wings (Distal­less gene)  Lenses of marine animals' eyes  Plants' leaves (Knox genes)
  • 6. Examples of Evolutionary Innovation  As novel molecules:  Enzymes that obtain the ability to catalyze  completely different reactions after a mutation  E.coli's L­Ru5P, that acquired aldolase activity after a  mutation  IDH (citric cycle) and IMDH (leucine synthesis)  Evolution of anti­freeze proteins
  • 7. Examples of Evolutionary Innovation  Paper published today (Jan 27th 2012)  A phage evolved the ability to infect a novel strain  of E.coli
  • 8. Evolutionary Innovations resume  In general, the aim of this book is to describe how  novel phenotypes are discovered
  • 9. Contents of Chapter 1 1)What is an “Evolutionary Innovation”? 2)Definition of “Genotype Space” and “Genotype  Network” 
  • 10. Definition of Genotype Space  The genotype space is the set of all possible  genotypes  Let's represent it as a matrix where two neighbor  genotypes differ only for one position: AAAAA AAAAC AAAAG AAAAT AAATT AAACA AAACC AAACG AAACT AAATC AACCA AACCC AACCG AACCT ….. ACCCA ACCCC ACCCG ACCCT ….. CCCCA CCCCC CCCCG CCCCT ….. ….. ….. ….. ….. …..
  • 11. Example of Genotype Space  The genotype space is the set of all possible  genotypes  Let's represent it as a matrix where two neighbor  genotypes differ only for one position: Only one AAAAA AAAAC AAAAG AAAAT AAATT difference between AAACA AAACC AAACG AAACT AAATC neighbor AACCA AACCC AACCG AACCT ….. points ACCCA ACCCC ACCCG ACCCT ….. CCCCA CCCCC CCCCG CCCCT ….. ….. ….. ….. ….. …..
  • 12. Example of Genotype Space  The genotype space is the set of all possible  genotypes  Let's represent it as a matrix where two neighbor  genotypes differ only for one position: Only one AAAAA AAAAC AAAAG AAAAT AAATT difference between AAACA AAACC AAACG AAACT AAATC neighbor AACCA AACCC AACCG AACCT ….. points ACCCA ACCCC ACCCG ACCCT ….. CCCCA CCCCC CCCCG CCCCT ….. ….. ….. ….. ….. ….. Here, genotypes are represented as sequences, but they can be other things (will be discussed later)
  • 13. Genotype network  A genotype network is a set of genotypes that have the same  phenotype, and are connected by single pairwise differences  Let's assume that the marked genotypes have the same  phenotype: AAAAA AAAAC AAAAG AAAAT AAATT AAACA AAACC AAACG AAACT AAATC AACCA AACCC AACCG AACCT ….. ACCCA ACCCC ACCCG ACCCT ….. CCCCA CCCCC CCCCG CCCCT ….. ….. ….. ….. ….. …..  → Yellow genotypes represent a genotype network
  • 14. Genotype networks resume  The concept of Genotype networks allows us to  study how much a genotype can vary, without  changing the phenotype  This is important to get to the final aim of this  book: understand how new innovative phenotypes  are discovered
  • 15. Exploring genotype networks AAAAA AAAAC AAAAG AAAAT AAATT AAACA AAACC AAACG AAACT AAATC AACCA AACCC AACCG AACCT ….. ACCCA ACCCC ACCCG ACCCT ….. CCCCA CCCCC CCCCG CCCCT ….. ….. ….. ….. ….. …..  How big can a genotype network be?  How can a population explore a genotype network?  ….. many answers in the next chapters of the book
  • 16. Exploring Genotype networks  AAAAA and ACCCT have the same phenotype (they are in the  same genotype network)  Do their neighbors (e.g. AAACA and CCCT) have similar  phenotype? AAAAA AAAAC AAAAG AAAAT AAATT AAACA AAACC AAACG AAACT AAATC AACCA AACCC AACCG AACCT ….. ACCCA ACCCC ACCCG ACCCT ….. CCCCA CCCCC CCCCG CCCCT ….. ….. ….. ….. ….. …..
  • 17. Example of Genotype network taken from the book (fig. 2.6)  The lines corresponds to genotypes in a genotype network  G1 and G2 have the same phenotype  White spaces correspond to genotypes that don't have the  phenotype analyzed in this genotype network
  • 18. Extending the definition of “genotype”  Depending what we want to study, we can use  different definitions of “genotype” and  “phenotype”  For example, in a metabolic network, the genotype  can be the set of reactions that an organism can  catalize  
  • 19. Example of alternative definition of “genotype” MGAT1 functional MGAT1 functional MGAT1 functional MGAT1 functional MGAT2 functional MGAT2 not functional MGAT2 not functional MGAT2 not functional MGAT3 functional MGAT3 functional MGAT3 not functional MGAT3 not functional MGAT4 functional MGAT4 functional MGAT4 functional MGAT4 not functional MGAT5 functional MGAT5 functional MGAT5 functional MGAT5 functional MGAT1 not functional MGAT1 not functional MGAT1 not functional MGAT1 not functional MGAT2 functional MGAT2 not functional MGAT2 not functional MGAT2 not functional MGAT3 functional MGAT3 functional MGAT3 not functional MGAT3 not functional MGAT4 functional MGAT4 functional MGAT4 functional MGAT4 not functional MGAT5 functional MGAT5 functional MGAT5 functional MGAT5 functional MGAT1 not functional MGAT1 not functional MGAT1 not functional …. MGAT2 not functional MGAT2 not functional MGAT2 not functional MGAT3 functional MGAT3 functional MGAT3 functional MGAT4 functional MGAT4 not functional MGAT4 not functional MGAT5 functional MGAT5 functional MGAT5 functional MGAT1 not functional MGAT1 functional MGAT1 not functional ….. MGAT2 not functional MGAT2 not functional MGAT2 not functional MGAT3 not functional MGAT3 not functional MGAT3 not functional MGAT4 functional MGAT4 functional MGAT4 functional MGAT5 functional MGAT5 functional MGAT5 not functional
  • 20. Example of alternative definition of “genotype”  All the yellow cells have the same phenotype (e.g.  they can produce glycosylation) MGAT1 functional MGAT1 functional MGAT1 functional MGAT1 functional MGAT2 functional MGAT2 not functional MGAT2 not functional MGAT2 not functional MGAT3 functional MGAT3 functional MGAT3 not functional MGAT3 not functional MGAT4 functional MGAT4 functional MGAT4 functional MGAT4 not functional MGAT5 functional MGAT5 functional MGAT5 functional MGAT5 functional MGAT1 not functional MGAT1 not functional MGAT1 not functional MGAT1 not functional MGAT2 functional MGAT2 not functional MGAT2 not functional MGAT2 not functional MGAT3 functional MGAT3 functional MGAT3 not functional MGAT3 not functional MGAT4 functional MGAT4 functional MGAT4 functional MGAT4 not functional MGAT5 functional MGAT5 functional MGAT5 functional MGAT5 functional MGAT1 not functional MGAT1 not functional MGAT1 not functional …. MGAT2 not functional MGAT2 not functional MGAT2 not functional MGAT3 functional MGAT3 functional MGAT3 functional MGAT4 functional MGAT4 not functional MGAT4 not functional MGAT5 functional MGAT5 functional MGAT5 functional MGAT1 not functional MGAT1 functional MGAT1 not functional ….. MGAT2 not functional MGAT2 not functional MGAT2 not functional MGAT3 not functional MGAT3 not functional MGAT3 not functional MGAT4 functional MGAT4 functional MGAT4 functional MGAT5 functional MGAT5 functional MGAT5 not functional
  • 21. Take home messages  Genotype network != biological pathways  Genotype network: set of possible genotypes  sharing the same phenotype, and connected  Evolutionary Innovation: any novel phenotype  Theory of Innovation: studies how populations can  explore the genotype space, the properties of  genotype networks, and how innovative  phenotypes can be found