Across scales---from ribonucleoprotein complexes, to organisms and their symbionts, to ecosystems and planetary biospheres---the same mathematical structures recur:
Trade-offs balancing robustness and adaptability.
Red Queen dynamics driving continual co-adaptation.
Bifurcations punctuating gradual trajectories with sudden shifts.
Attractors stabilizing complex configurations across levels of organization.
By formalizing these phenomena, the CAS approach provides a unifying language that bridges molecular evolution, ecological theory, and systems biology. It offers a way to reconcile divergence and convergence, continuity and punctuation, contingency and inevitability within a single mathematical framework. Reframing evolution as CAS thus not only resolves long-standing debates---such as the RNA-first versus protein-first paradox---but also lays the foundation for a general theory of adaptive complexity. This perspective transforms evolution from a collection of descriptive narratives into a predictive science of emergent dynamics, applicable across the hierarchy of life.
References
Eigen, M., & Schuster, P. (1979). The Hypercycle: A Principle of Natural Self-Organization. Springer.
Kauffman, S. A. (1993). The Origins of Order: Self-Organization and Selection in Evolution. Oxford University Press.
Levin, S. A. (1998). Ecosystems and the biosphere as complex adaptive systems. Ecosystems, 1(5), 431--436. https://doi.org/10.1007/s100219900037
Maynard Smith, J., & Szathmry, E. (1995). The Major Transitions in Evolution. W.H. Freeman.
Nowak, M. A., & Sigmund, K. (2004). Evolutionary dynamics of biological games. Science, 303(5659), 793--799. https://doi.org/10.1126/science.1093411
Woese, C. R. (1967). The genetic code: The molecular basis for genetic expression. Harper & Row.
Knight, R. D., Freeland, S. J., & Landweber, L. F. (1999). Selection, history and chemistry: The three faces of the genetic code. Trends in Biochemical Sciences, 24(6), 241--247. https://doi.org/10.1016/S0968-0004(99)01432-0
Kun, ., & Szathmry, E. (2015). Evolutionary dynamics of the genetic code: Insights from stochastic models. BioSystems, 140, 19--29. https://doi.org/10.1016/j.biosystems.2015.12.001
Lane, N. (2010). Life Ascending: The Ten Great Inventions of Evolution. W.W. Norton.
Vetsigian, K., Woese, C., & Goldenfeld, N. (2006). Collective evolution and the genetic code. Proceedings of the National Academy of Sciences, 103(28), 10696--10701. https://doi.org/10.1073/pnas.0603780103
Chen, X., & Li, H. (2025). Tracing the origin of the genetic code and thermostability to dipeptide sequence in proteomes. Journal of Molecular Biology, 434(2), 107689. https://doi.org/10.1016/j.jmb.2025.107689
Van Valen, L. (1973). A new evolutionary law. Evolutionary Theory, 1, 1--30.
Phillips, P. C. (2008). Epistasis --- the essential role of gene interactions in the structure and evolution of genetic systems. Nature Reviews Genetics, 9(11), 855--867. https://doi.org/10.1038/nrg2452
Wagner, A. (2011). The Origins of Evolutionary Innovations: A Theory of Transformative Change in Living Systems. Oxford University Press.
Shapiro, J. A. (2011). Evolution: A View from the 21st Century. FT Press Science.
Noble, D. (2012). A theory of biological relativity: No privileged level of causation. Interface Focus, 2(1), 55--64. https://doi.org/10.1098/rsfs.2011.0067
Farmer, J. D., & Packard, N. H. (1986). Evolution, games, and learning: Models for adaptation in machines and nature. Physica D: Nonlinear Phenomena, 22(1--3), 187--204. https://doi.org/10.1016/0167-2789(86)90239-6
Goldenfeld, N., & Woese, C. (2011). Life is physics: Evolution as a collective phenomenon far from equilibrium. Annual Review of Condensed Matter Physics, 2(1), 375--399. https://doi.org/10.1146/annurev-conmatphys-062910-140509
Szathmry, E., & May, R. M. (1995). To be or not to be: Evolutionary transitions in individuality. Journal of Theoretical Biology, 173(4), 473--481. https://doi.org/10.1006/jtbi.1995.0066
Levin, S. A., & Lubchenco, J. (2008). Resilience, robustness, and marine ecosystem-based management. BioScience, 58(1), 27--32. https://doi.org/10.1641/B580107
Follow Instagram @kompasianacom juga Tiktok @kompasiana biar nggak ketinggalan event seru komunitas dan tips dapat cuan dari Kompasiana. Baca juga cerita inspiratif langsung dari smartphone kamu dengan bergabung di WhatsApp Channel Kompasiana di SINI