Frasch, Martin G. (2025) Causal Thinking in Physiology: A Search for Vertically Organizing Principles. [Preprint]
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Abstract
Physiology has excelled at elucidating biological mechanisms at specific scales of organization, yet it lacks a robust framework for understanding causality across these scales. This paper argues for a paradigm shift, moving from a primary focus on scale-specific, or "horizontal," causality to a search for "vertically organizing" principles that are invariant across biological levels. Drawing inspiration from concepts in physics such as scaling laws, emergence, coarse-graining, and action principles, we explore the limitations of current causal thinking in physiology, particularly the prevailing genome-centric bias.
This paper is the first in a series exploring vertically organizing principles in biology. Here we establish the philosophical and theoretical foundation, examining existing frameworks and their limitations. We review information-theoretic approaches including the Free Energy Principle and dissipative adaptation, acknowledging both their insights and their challenges in achieving scale invariance. We introduce the concept of network-weighted action principles (minAction.net) as a promising candidate for scale-invariant organization that may complement these existing frameworks.
Building on Schrödinger's two secrets of life — heredity and self-organization — we argue that a third secret lies in the scale-invariant action principles that give rise to biological design itself. We present evidence from immunometabolism, developmental biology, complex systems physiology, and computational evolution studies supporting multi-scale, information-centric views of biological organization. Notably, recent computational evidence demonstrates that minimizing connection costs—a key component of action functionals—spontaneously generates the modular, evolvable architectures ubiquitous in biological systems. Finally, we posit that meaning—operationally defined as the successful reduction of uncertainty through predictive modeling and efficient action—can be conceptualized as a candidate for a high-level, vertically organizing principle that operates from the cellular to the cognitive level. This manuscript lays the conceptual foundation for a more unified and scalable science of life.
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