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Adventures in protein world reveal
the Tree of Life and the evolution of
             complexity




Graeme T. Lloyd, Philip C. J. Donoghue and
              Julian Gough
Scala Naturae
Schemes for Increasing Complexity I
          The Basis of Progressive Evolution (1969)

                              Self-replicating molecules
                                     Prokaryotes




                                                                   Increasing complexity
                               Single-celled eukaryotes
                          Multicelled eukaryotes with cellular
                                     differentiation
                       Organisms with differentiated tissues and
                                       organs
                       Organisms with well-developed limbs and
                                 nervous systems
 G. Ledyard Stebbins                Homeotherms
                                    Human beings
Schemes for Increasing Complexity II
           Major Transitions in Evolution (1995)




                                                                       Increasing complexity
  Replicating molecules --> Populations of molecules in compartments
              Independent replicators --> Chromosomes
             RNA as gene and enzyme --> DNA + protein
                      Prokaryotes --> Eurkaryotes
                Asexual clones --> Sexual populations
         Protists --> Animals, plants, fungi (cell differentiation)
       Solitary individuals --> Colonies (non-reproductive castes)
           Primate societies --> Human societies (language)


                     Maynard Smith and Szathmáry
Schemes for Increasing Complexity III
           Megatrajectory Sequence




                                                   Increasing complexity
         Increase in efficency of life processes
               Prokaryote diversification
          Unicellular eukaryote diversification
                Aquatic multicellularity
                  Invasion of the land
                      Intelligence


                Knoll & Bambach 2000
The Problem



“Complexity is hard to define or measure…”
            Maynard Smith and Szathmáry, 1995
Segmentation and Complexity I
   Differentiation of centrum length in vertebrates




                                                          Increasing complexity
                                                      (from McShea 1992)
Segmentation and Complexity II
     Functional differentiation in arthropod limbs




                                                       Increasing complexity
                                                     (from Cisne 1974)
Segmentation and Complexity III
 Increasing complexity   Differentiation of segment length in centipedes




                                                                  (from Fusco and Minelli 2000
Fractals and Complexity
   Fractal dimension in ammonoids




   Increasing complexity




                                    (from Boyiajian & Lutz 1992)
Cell Number and Complexity




                                        Increasing complexity
                     (from Valentine et al. 1994)
Genome Size and the Scala Naturae
   Increasing complexity




                           (from Gregory 2005)
Proteome Size and the Scala Naturae




               ?
Protein Structural Domains



Folding
SCOP Protein Domain Classification
Classes
(7)

Folds
(48)

Superfamily
(1445)

Family
(2598)

Domain
(75930)
Protein Domain Classification
             A   B   C   D


Protein 1:                   Architecture = A,A,C


Protein 2:                   Architecture = D,B


Protein 3:                   Architecture = B


Protein 4:                   Architecture = C,A,A


Protein 5:                   Architecture = D,B,C
http://supfam.org   Protein Domain Database I
Protein Domain Database II



238 Bacteria    26 Basal         53 Fungi        8 Arthropods
               Eukaryotes




27 Archaea      9 Plants     5 Invertebrates     19 Chordates
                            (excl. Arthropods)
Superfamily Number and the Scala
             Naturae
Family Number and the Scala Naturae
Architecture Number and the Scala
              Naturae
The Tree of Life I




                                     Architectures
Superfamilies      Families
The Tree of Life II
The Tree of Life III
The Tree of Life IV
Protein Space
Protein Space
Principal Coordinate Analysis – A Very Brief
Introduction



                                               PC1
   Variable 2




                                 PC
                                     2


                        Variable 1
Protein Space
Principal Coordinate Analysis – A Very Brief
Introduction
Protein Space
Principal Coordinate Analysis – A Very Brief
Introduction
Protein Space - Superfamilies
Protein Space - Families
Protein Space - Architectures
Tempo and Protein Evolution
Tempo and Protein Evolution
 Tempo and mode of evolution in a character complex




           (from Westoll 1949)
Tempo and Protein Evolution
   Tempo and mode of cell type evolution I




                                                       Increasing complexity
                                        (from Valentine et al. 1994)
Tempo and Protein Evolution
                           Tempo and mode of cell type evolution II
Increasing complexity




                                                                  (from Hedges et al. 2004)
Tempo and Protein Evolution -
       Superfamilies




LUCA
Tempo and Protein Evolution -
          Families




LUCA
Tempo and Protein Evolution -
       Architectures




LUCA
Conclusions
• Protein domains can help us reconstruct
  the tree of life and offer a promising new
  metric for biologic complexity
• Novel domain families and superfamilies
  have appeared at a roughly constant rate
  over the history of life and across different
  groups
• By contrast, novel architectures (new
  proteins) are acquired at a faster rate in
  animals, and vertebrates in particular

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