  Representations are internal models of the environment that can provide
guidance to a behaving agent, even in the absence of sensory information. It is
not clear how representations are developed and whether or not they are
necessary or even essential for intelligent behavior. We argue here that the
ability to represent relevant features of the environment is the expected
consequence of an adaptive process, give a formal definition of representation
based on information theory, and quantify it with a measure R. To measure how R
changes over time, we evolve two types of networks---an artificial neural
network and a network of hidden Markov gates---to solve a categorization task
using a genetic algorithm. We find that the capacity to represent increases
during evolutionary adaptation, and that agents form representations of their
environment during their lifetime. This ability allows the agents to act on
sensorial inputs in the context of their acquired representations and enables
complex and context-dependent behavior. We examine which concepts (features of
the environment) our networks are representing, how the representations are
logically encoded in the networks, and how they form as an agent behaves to
solve a task. We conclude that R should be able to quantify the representations
within any cognitive system, and should be predictive of an agent's long-term
adaptive success.
