For over a century, Darwin's theory of natural selection has provided the foundational core of evolutionary biology. This framework, however, is rooted in a reductionist and substance-oriented ontology that focuses on the selection of discrete traits. The integrative, holistic methodologies of systems biology now challenge this view, shifting the focus towards the dynamic processes and relations that constitute living systems, arguing that selection primarily acts on the adaptive potential and r…
Read moreFor over a century, Darwin's theory of natural selection has provided the foundational core of evolutionary biology. This framework, however, is rooted in a reductionist and substance-oriented ontology that focuses on the selection of discrete traits. The integrative, holistic methodologies of systems biology now challenge this view, shifting the focus towards the dynamic processes and relations that constitute living systems, arguing that selection primarily acts on the adaptive potential and regulatory logic of entire developmental systems. This review argues that systems biology does not overturn but fundamentally expands the Darwinian paradigm, catalyzing the development of an Extended Evolutionary Synthesis. By examining concepts such as gene regulatory networks, phenotypic plasticity, and dynamic organism-environment interactions (illustrated through examples like plant photomorphogenesis and the adaptive responses of Gasterosteus aculeatus), we demonstrate that evolution operates as a multi-level process governed by reciprocal causation and feedback loops. Crucially, this scientific shift necessitates a parallel philosophical transition: from a static, substance-based ontology to a processual one. It challenges the primacy of reductionism, linear causality, and the nature-nurture dichotomy, advocating instead for a relational ontology where emergence, agency, and becoming are fundamental. We conclude that systems biology fulfills Darwin's vision by providing a process-oriented framework adequate to the breathtaking complexity of the living world, one where evolution is understood as a dynamic interplay of recursive interactions between organisms and their environments. In the following, we explore four interrelated dimensions of contemporary evolutionary biology: the limitations of reductionism, the nature of dynamic interactions, the regulatory logic encoded in gene networks, and the evolutionary role of phenotypic plasticity.