"Particularly compelling for its smooth integration of biological experiments, physical models, and computational exercises. Readers who complete the text will be well equipped with the computational and mathematical skills needed for a quantitative understanding of a range of biological systems.... Thanks to Nelson's skillful writing and the excellent accompanying online resources, this book will appeal to a broad audience and teach even a beginner how to solve problems numerically." -- Eva-Maria Collins in Physics Today
""Philip Nelson has done a terrific job.... There are numerous traits that make this text unique among the very many books of biological physics.... The presentation of materials is developed in an innovative fashion.... There is a nice balance between conceptual examples and end-of-the-chapter problems.... This book shows a nice intercalation of fundamental laws, brief descriptions of computational strategies for acquiring quantitative information, as well as their implications in biological physics and areas beyond that, including signaling processes, genetic switches, and cellular oscillators.... Physical Models of Living Systems... will benefit undergraduates as well as others with clear interests in genomics, proteomics, cellular signaling, bioengineering, regenerative medicine, and synthetic biology." -- Liviu Movileanu in American Journal of Physics 84 (6), June 2016
"This book inspired me to write [my] book in the first place. Biological Physics is the most interesting and well-written textbook I have ever read." -- Peter M. Hoffman, Life's Ratchet
This 5th printing differs from the 4th by the updating of many of the beautiful drawings by David Goodsell. It also differs from the 3d and earlier printings by the addition of many new homework problems.
Phil NelsonPhysics and Astronomy University of Pennsylvania Philadelphia, PA 19104 USA phone: (215) 898-7001 fax: (215) 898-2010
Sorry, but I get a lot of e-mail. If you are a student currently enrolled in a class that I teach, or a Biophysics major advisee, put that in the subject line. I want to reply, but if I don't --- please come see me. If you wish me to review a grant proposal, article for publication, or promotion case, please don't assume that your e-mail has been read or even seen. It may be necessary to get me on the phone if you need me to do those things.
I am a physicist.
Some of what I know about science is in my first book.. Some more can be found in my second book.
Some of what I know about computers is in my little book. Some more can be found in my little free book.
I teach some courses.
I'm also interested in K-12 education.
See our group's homepage.
I'm a member of Penn's Nano-Bio Interface Center, and the Institute for Medicine and Engineering.
I'm a member of Penn's Applied Math and Computational Science Graduate Group.
I'm a General Member of the Aspen Center for Physics.
Bill Berner's Physics demo shows! Get them via iTunesU or via vimeo.
Spare the (Elastic) Rod, Science 337, 1045 (2012).
Physics in the cell: Spring theory by Brendan Maher (Nature 448, 984-986 (30 August 2007)).
The great hunt for extra compliance by Jan Liphardt (Biophys. J. 2007).
Teaching Biological Physics, Physics Today 58:3 46 (March 2005).
Excerpt from a review of my book, published in The American Journal of Physics.
And another review published in Nature.
Here's one published in Biomedical Computation Review.
Learning physical biology via modeling and simulation, talk at AAPT meeting 7/2016.
Old news and new news about single-photon sensitivity, talk at McGill University, 3/2016.
An intermediate-level course on physical models of living systems, talk at American Assn of Physics Teachers, 1/2015.
Inference in microscopy and biological physics, Talk at Advanced Workshop on Out-of-Equilibrium Matter, San Luis Potosi, 12/2014.
,Simultaneous activity measurements on intact mammalian retina Talk at Advanced Workshop on Out-of-Equilibrium Matter, San Luis Potosi, 12/2014.
Transformation of stimulus correlations by the retina, Talk at American Physical Society, 3/2014.
Mechanical metaphors in biophysics teaching, Talk at American Physical Society, 3/2014.
Evolution--the first single-molecule biophysicist, (Nicholas Cabrera Summer School, Madrid, 2013).
The research in these publications was funded in part by The National Science Foundation and by the Human Frontier Science Program. "Any opinions, findings, confusions, contusions, contortions, and conclusions or recommendations expressed in this material are those of the author(s) and do not necessarily reflect the views of the National Science Foundation."
Note: PDF reprints are provided below within the context of fair use. Please obtain copies from the publisher if appropriate.
|Does the quantum character of light matter for our eyes?
Old and new results about single-photon sensitivity in human vision. Physical Biology 13 025001 (2016).
|Does a molecular motor pause to search for its next stepping spot?
Tilting and wobble of myosin V by high-speed single-molecule polarized fluorescence microscopy. With Beausang, Schroder, and Goldman. Biophysical Journal 104 1263--1273 (2013). (featured in BJ's "Best of 2013"
|How does a wave of cell division spread across an embryo?
Wavefront propagation and mechanical signaling in early Drosophila embryos. With Idema, Dubuis, Kang, Manning, Lubensky, and Liu. PLoS ONE 8 e77216 (2013).
|How much data compression is occurring in your retina?
Retinal adaptation to spatial correlations. With Simmons, Prentice, Tkacik, Homann, Yee, Palmer, and Balasubramanian. PLoS Computational Biology 9 e1003344 (2013).
|What is each retinal ganglion cell looking for?
Neural Spikes, Identification from a Multielectrode Array. With Prentice, Homann, Simmons, Tkacik, Balasubramanian. Encyclopedia of Applied and Computational Mathematics, in press.
|How precisely can you pinpoint a rate change?.
Changepoint analysis for single-molecule polarized total internal reflection fluorescence microscopy experiments. With Beausang and Goldman. Methods Enzymol. 487 431-463 (2011).
|When we eavesdrop on a retina, who said what when?
Fast, scalable, Bayesian spike identification for multi-electrode arrays. With Prentice, Homann, Simmons, Tkacik, and Balasubramanian. PLoS ONE 6(7): e19884.
|How should you design the right DNA to make the best measurements of looping?
Calibration of Tethered Particle Motion Experiments. With Han, Lui, Blumberg, Beausang, and Phillips.In Mathematics of DNA Structure, Function and Interactions, eds. C.J. Benham et al. (Springer, 2009).
|How can we monitor gene-regulatory events without any cells?
Concentration and Length Dependence of DNA Looping. With Han, Garcia, Blumberg, Towles, Beausang, and Phillips. PLoS ONE, 4 e5621 (2009).
|Does myosin twist the `wrong' way around actin as it walks?
Twirling of actin by myosins II and V observed via polarized TIRF in a modified gliding assay. With Beausang, Schroeder, and Goldman. Biophys J. 95 5820 (2008).
|Is there spooky action at a distance in living cells?
The Role of Microtubule Movement in Bidirectional Organelle Transport. With Kulic, Brown, Kim, Kural, Blehm, Selvin, and Gelfand. PNAS 105 10011 (2008). (See also the amazing movies.)
|How does a regulatory protein manage to grab two distant places on a strand of DNA?
First-principles calculation of DNA looping in tethered particle experiments, final version, and its supplement. With Towles, Beausang, Garcia, and Phillips. Physical Biology, 6 025001 (2009).
|When experiments seem to imply that the stiffness of DNA depends on its length, what's really going on?
Elasticity of short DNA molecules: Theory and experiment for contour lengths of 0.6--7 um. With Seol, Li, Perkins, and Betterton. Biophys. J. 93 4360 (2007).
|How accurately can we infer the length of a long molecule from the motion of a bead on its end?
Colloidal Particle Motion as a Diagnostic of DNA Conformational Transitions. Current Opinion in Colloid and Interface Science 12 307 (2007).
|Are there long-lived kinetic substates in DNA looping?
Diffusive hidden Markov model characterization of DNA looping dynamics in tethered particle experiments . With Beausang. Physical Biology 4 205 (2007).
|How can you extract kinetics from noisy data without binning?
DNA looping kinetics analyzed using diffusive hidden Markov model . With Beausang, Zurla, Manzo, Dunlap, and Finzi. Biophys. J. 92, L64 (2007).
|Are there hidden regularities in "random" Brownian motion?
Elementary simulation of tethered Brownian motion. With Beausang, Zurla, Sullivan, and Finzi. Am. J. Phys. 75, 520 (2007).
|What energy source squirts viral DNA into a host cell?
Biological Consequences of Tightly Bent DNA: The Other Life of a Macromolecular Celebrity . With Garcia, Grayson, Han, Inamdar, Kondev, Phillips, Widom, and Wiggins. Biopolymers 85, 115 (2007).
|If DNA is a stiff elastic rod, how can it form those tight regulatory loops?
High flexibility of DNA on short length scales probed by atomic force microscopy. With Wiggins, van der Heijden, Moreno-Herrero, Spakowitz, Phillips, Widom, and Dekker.Nature Nanotechnology 1, 137 (2006) A commentary appears here.
See the Erratum to this article.
|Do cells transport vesicles using active conveyor belts?
Hitchhiking Through the Cytoplasm. With Kulic. Europhys. Lett. 81, 18001-(1--6) (2008).
|How can you see protein binding in real time?
Entropic elasticity of DNA with a permanent kink. With Betterton and Li. Macromolecules 39 8816 (2006).
|When you twist a string, how does that affect loop formation?
Effect of supercoiling on formation of protein mediated DNA loops. With Purohit. Phys. Rev. E74, art. no. 061907 (14 pages) (2006).
See the Erratum to this article.
|How can you detect DNA looping in real time, when you can't even see DNA?
Tethered particle motion as a diagnostic of DNA tether length. With Zurla, Brogioli, Beausang, Finzi, and Dunlap. Journal of Physical Chemistry B110, 17260 (2006).
|How can fluorescence tell us about molecular mechanics?
Generalized theory of semiflexible polymers. With Wiggins. Phys. Rev. E73, art. number 031906 (2006).
|What can a balloon in a hurricane tell us about gene regulation?
Excluded-Volume Effects in Tethered-Particle Experiments: Bead Size Matters. With Segall and Phillips. Phys. Rev. Lett. 96 art. number 088306 (2006).
|How does DNA spontaneously form gene regulatory complexes with
"impossibly" tight loops?
Exact theory of kinkable elastic polymers . With Wiggins and Phillips. Phys. Rev. E71 art. number 021909 (2005).
|Is single-stranded DNA really a freely jointed chain?
Theory of High-Force DNA Stretching and Overstretching. With Storm. Phys. Rev. E67 (2003) art. number 051906 (Erratum ibid. 70, 013902 (2004)).
|What's going on when DNA stretches to 1.6 times its "maximum" length?
The bend stiffness of S-DNA. With Storm. Europhys. Lett. 62 (2003) 760.