Published November 2025 | Published
Journal Article

Approximating the living

  • 1. ROR icon California Institute of Technology

Abstract

Is a herd of wildebeest better thought of as a series of individual animals, each with its own glorious and unmanageable volition, or as a field of moving arrows? Are the morphogen gradients that set up the coordinate systems for embryonic anterior–posterior patterning a smooth and continuous concentration field or instead a chaotic collection of protein molecules each jiggling about in the haphazard way first described by Robert Brown in his microscopical observations of pollen? Is water, the great liquid ether of the living world, a collection of discrete molecules or instead a perfectly continuous medium with a density of 1000 kg/m? In this article, I will argue that these questions pose a false dichotomy since there are many different and powerful representations of the world around us. Different representations suit us differently at different times and it is often useful to be able to hold these seemingly contradictory notions in our heads simultaneously. Indeed, mathematics is not only the language of representation, but often is also the engine of reconciliation of such disparate views. In a letter to Alfred Russel Wallace on 14 April 1869, Charles Darwin noted that Lord Kelvin’s “views on the recent age of the world have been for some time one of my sorest troubles”. Here, I will argue that one of the highest attainments of the scientific enterprise is a coherent picture of the world, a picture in which our stories about the geological age of the Earth are coherent with our stories of how whales populated the oceans, our understanding of the living jibes with our understanding of the inanimate, our insights into the dynamics of genes and molecular structures are consonant with our physical understanding of the laws of statistical physics. The underpinnings of such coherency are often best revealed when viewed through the lens of mathematics.

Copyright and License

© 2025 Elsevier Ltd. All rights are reserved, including those for text and data mining, AI training, and similar technologies.

Acknowledgement

I am so grateful to the NIH for support through award numbers DP1OD000217 (Director’s Pioneer Award) and NIH MIRA 1R35 GM118043-01. The trust and financial support of this great institution make it possible for today’s scientists to grapple with the endless fascination of trying to understand the nature of life. Though my Pioneer Award ended long ago, the foundations laid down by that support will never go away. Participating in the CZI Theory Institute Without Walls which has as one its central themes the question of what is the “right” representation of biological systems has been a huge privilege and deeply inspiring with special thanks to Pankaj Mehta, Vincenzo Vitelli, Madhav Mani, Evelyn Tang and Jane Kondev for really pushing me to broaden my perspective. I am grateful to Gabe Salmon, Jane Kondev, Madhav Mani and Hernan Garcia for comments on the manuscript and related discussions. Willem Kegel many important discussions on input–output functions and data collapse as well as the history of the Hill function. Marc Kirschner continues to inspire me with his expansive and insightful vision of what the study of life can look like.

Conflict of Interest

I was invited by Wallace Marshall to submit an article for the special volume in which the question of when should mathematical models be used in biology is addressed. I declare no competing interests.

Additional details

Created:
September 17, 2025
Modified:
September 17, 2025