thermodynamics
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The Equation of State of a Tissue
An equation of state is something you hear about in introductory thermodynamics, for example, the Ideal gas equation. The ideal gas equation relates the pressure, volume, and the number of particles of the gas, to its temperature, uniquely defining its state. This description is possible in physics when the system under investigation is in equilibrium or near equilibrium. In biology, a tissue is modeled as a fluid composed of cells. These cells are constantly interacting with each other through mechanical and chemical signaling, driving them far from equilibrium. Can an equation of state exist for such a messy interacting system? In this talk, I show that the presence of strong cell-cell interaction in tissues gives rise to a novel non-equilibrium, size-dependent surface tension, something unheard of for classical fluids. This surface tension, in turn, modifies the packing of cells inside the tissue generating a size-dependent density and pressure. Finally, we show that a combination of these non-equilibrium pressure and densities can yield an equation of state for biological tissues arbitrarily far from equilibrium. In the end, I discuss how this new paradigm of size-dependent biological properties gives rise to novel modes of cellular motion in tissues
Is there universality in biology?
It is sometimes said that there are two reasons why physics is so successful as a science. One is that it deals with very simple problems. The other is that it attempts to account only for universal aspects of systems at a desired level of description, with lower level phenomena subsumed into a small number of adjustable parameters. It is a widespread belief that this approach seems unlikely to be useful in biology, which is intimidatingly complex, where “everything has an exception”, and where there are a huge number of undetermined parameters. I will try to argue, nonetheless, that there are important, experimentally-testable aspects of biology that exhibit universality, and should be amenable to being tackled from a physics perspective. My suggestion is that this can lead to useful new insights into the existence and universal characteristics of living systems. I will try to justify this point of view by contrasting the goals and practices of the field of condensed matter physics with materials science, and then by extension, the goals and practices of the newly emerging field of “Physics of Living Systems” with biology. Specific biological examples that I will discuss include the following: Universal patterns of gene expression in cell biology Universal scaling laws in ecosystems, including the species-area law, Kleiber’s law, Paradox of the Plankton Universality of the genetic code Universality of thermodynamic utilization in microbial communities Universal scaling laws in the tree of life The question of what can be learned from studying universal phenomena in biology will also be discussed. Universal phenomena, by their very nature, shed little light on detailed microscopic levels of description. Yet there is no point in seeking idiosyncratic mechanistic explanations for phenomena whose explanation is found in rather general principles, such as the central limit theorem, that every microscopic mechanism is constrained to obey. Thus, physical perspectives may be better suited to answering certain questions such as universality than traditional biological perspectives. Concomitantly, it must be recognized that the identification and understanding of universal phenomena may not be a good answer to questions that have traditionally occupied biological scientists. Lastly, I plan to talk about what is perhaps the central question of universality in biology: why does the phenomenon of life occur at all? Is it an inevitable consequence of the laws of physics or some special geochemical accident? What methodology could even begin to answer this question? I will try to explain why traditional approaches to biology do not aim to answer this question, by comparing with our understanding of superconductivity as a physical phenomenon, and with the theory of universal computation. References Nigel Goldenfeld, Tommaso Biancalani, Farshid Jafarpour. Universal biology and the statistical mechanics of early life. Phil. Trans. R. Soc. A 375, 20160341 (14 pages) (2017). Nigel Goldenfeld and Carl R. Woese. Life is Physics: evolution as a collective phenomenon far from equilibrium. Ann. Rev. Cond. Matt. Phys. 2, 375-399 (2011).
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