W. Fontana et Lw. Buss, THE ARRIVAL OF THE FITTEST - TOWARD A THEORY OF BIOLOGICAL ORGANIZATION, Bulletin of mathematical biology, 56(1), 1994, pp. 1-64
The formal structure of evolutionary theory is based upon the dynamics
of alleles, individuals and populations. As such, the theory must ass
ume the prior existence of these entities. This existence problem was
recognized nearly a century ago, when DeVries (1904, Species and Varie
ties: Their Origin by Mutation) stated, ''Natural selection may explai
n the survival of the fittest, but it cannot explain the arrival of th
e fittest.'' At the heart of the existence problem is determining how
biological organizations arise in ontogeny and in phylogeny. We develo
p a minimal theory of biological organization based on two abstraction
s from chemistry. The theory is formulated using lambda-calculus, whic
h provides a natural framework capturing (i) the constructive feature
of chemistry, that the collision of molecules generates specific new m
olecules, and (ii) chemistry's diversity of equivalence classes, that
many different reactants can yield the same stable product. We employ
a well-stirred and constrained stochastic flow reactor to explore the
generic behavior of large numbers of applicatively interacting lambda-
expressions. This constructive dynamical system generates fixed system
s of transformation characterized by syntactical and functional invari
ances. Organizations are recognized and defined by these syntactical a
nd functional regularities. Objects retained within an organization re
alize an algebraic structure and possess a grammar which is invariant
under the interaction between objects. An organization is self-maintai
ning, and is characterized by (i) boundaries established by the invari
ances, (ii) strong self-repair capabilities responsible for a robustne
ss to perturbation, and (iii) a center, defined as the smallest kineti
cally persistent and self-maintaining generator set of the algebra. Im
position of different boundary conditions on the stochastic flow react
or generates different levels of organization, and a diversity of orga
nizations within each level. Level 0 is defined by self-copying object
s or simple ensembles of copying objects. Level 1 denotes a new object
class, whose objects are self-maintaining organizations made of Level
0 objects, and Level 2 is defined by self-maintaining metaorganizatio
ns composed of Level 1 organizations. These results invite analogy to
the history of life, that is, to the progression from self-replication
to self-maintaining procaryotic organizations to ultimately yield sel
f-maintaining eucaryotic organizations. In our system self-maintaining
organizations arise as a generic consequence of two features of chemi
stry, without appeal to natural selection. We hold these findings as c
alling for increased attention to the structural basis of biological o
rder.