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It has been hypothesized that one of the main reasons evolution has been able to produce such impressive adaptations is because it has improved its own ability to evolve -- "the evolution of evolvability". Rupert Riedl, for example, an early pioneer of evolutionary developmental biology, suggested that the evolution of complex adaptations is facilitated by a developmental organization that is itself shaped by past selection to facilitate evolutionary innovation. However, selection for characteristics that enable future innovation seems paradoxical: natural selection cannot favor structures for benefits they have not yet produced, and favoring characteristics for benefits that have already been produced does not constitute future innovation. Here we resolve this paradox by exploiting a formal equivalence between the evolution of evolvability and learning systems. We use the conditions that enable simple learning systems to generalize, i.e., to use past experience to improve performance on previously unseen, future test cases, to demonstrate conditions where natural selection can systematically favor developmental organizations that benefit future evolvability. Using numerical simulations of evolution on highly epistatic fitness landscapes, we illustrate how the structure of evolved gene regulation networks can result in increased evolvability capable of avoiding local fitness peaks and discovering higher fitness phenotypes. Our findings support Riedl's intuition: Developmental organizations that "mimic" the organization of constraints on phenotypes can be favored by short-term selection and also facilitate future innovation. Importantly, the conditions that enable the evolution of such surprising evolvability follow from the same non-mysterious conditions that permit generalization in learning systems.

One of the most intriguing questions in evolution is how organisms exhibit suitable phenotypic variation to rapidly adapt in novel selective environments which is crucial for evolvability. Recent work showed that when selective environments vary in a systematic manner, it is possible that development can constrain the phenotypic space in regions that are evolutionarily more advantageous. Yet, the underlying mechanism that enables the spontaneous emergence of such adaptive developmental constraints is poorly understood. How can natural selection, given its myopic and conservative nature, favour developmental organisations that facilitate adaptive evolution in future previously unseen environments? Such capacity suggests a form of \textit{foresight} facilitated by the ability of evolution to accumulate and exploit information not only about the particular phenotypes selected in the past, but regularities in the environment that are also relevant to future environments. Here we argue that the ability of evolution to discover such regularities is analogous to the ability of learning systems to generalise from past experience. Conversely, the canalisation of evolved developmental processes to past selective environments and failure of natural selection to enhance evolvability in future selective environments is directly analogous to the problem of over-fitting and failure to generalise in machine learning. We show that this analogy arises from an underlying mechanistic equivalence by showing that conditions corresponding to those that alleviate over-fitting in machine learning enhance the evolution of generalised developmental organisations under natural selection. This equivalence provides access to a well-developed theoretical framework that enables us to characterise the conditions where natural selection will find general rather than particular solutions to environmental conditions.