Abstract
The mathematical models of polymers and membranes are introduced, to describe the microscopic deformation of biological materials. The mechanical properties at the molecular scale are inherently statistical, and remain hidden below macroscopic averages. But we can now see and manipulate a single molecule, and follow its movements in real time. This is made possible thanks to the advent of revolutionary tools, such as the atomic-force microscope and the optical or magnetic tweezers, which allow to test the mechanics of biological materials with unprecedented quality and accuracy, down to the molecular scale. Today it is possible to measure the elasticity of a single molecule, of a piece of DNA, of a fragment of cell membrane, and from such strictly physical comparisons, a totally new wealth of information has started to invade the already flooded desk of the biologist.
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Notes
- 1.
For the definitions of the stress/strain tensors (\(\sigma , \varepsilon \)) and elastic moduli (\(E,\nu \)), see Appendix H on p. 422.
- 2.
As a curiosity, it may be noted that Helfrich was at that time working with Hoffmann-LaRoche Laboratories in Switzerland, on the fabrication of the first liquid-crystals displays, a subject he had started years before when working at RCA in Princeton and for which he is a credited inventor.
- 3.
- 4.
For the many details of this extraordinary process, the reader can consult any good biology textbook, such as the classical Molecular Biology of the Cell, by Alberts, Bray, Lewis, Raff, Roberts and Watson (Garland Science Pub., New York, 4th ed. 2002).
- 5.
Notably, plant cells operate the membrane division by an entirely different mechanism. After chromosome splitting, a portion of the rigid cell wall that surrounds most vegetable cells is nucleated in a new plaque, around the residual microtubules from the spindle that form a temporary structure called phragmoplast. This plaque starts with a roughly discoidal shape in the cell mid-plane, and grows in size with the help of microtubules, which dynamically reorganise around its perimeter. The process continues until the plaque entirely forms a new wall, dividing in two the old cell.
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Problems
Problems
8.1
Hollow versus filled
Calculate the persistence length and the flexural rigidity, for a hollow microtubule with inner and outer radii 10 nm and 12 nm, and Young’s modulus \(E=150\) MPa. Compare the results with a solid microtubule, with the same amount of mass per unit length.
8.2
Bacterial DNA
The DNA of some bacterium contains 3.45 million base pairs. (a) Calculate the contour length. (b) Given the average DNA Young’s modulus \(E=350\) MPa, calculate its persistence length. How does it compare with the contour length? (c) Given the DNA diameter of 2 nm, what is the smallest volume into which the double strand can be packed? (d) Calculate the end-to-end distance and gyration radius of this DNA. How these compare with the size of a typical bacterium?
8.3
Exocitosis
Find an explicit expression for the surface tension energy, \(E_S = \varSigma A\), during the three steps of the formation of a spherical liposome, illustrated in the figure below. Starting from the membrane at rest (1), a pseudopod is extruded (2), which eventually forms a spherical vesicle of radius R, fully detached from the cell membrane.
8.4
Membranes with an edge
A cell membrane is composed by two different types of phospholipids, the A with tails of length \(n_A=14\) and the B with \(n_B=18\). By assuming that bilayers form only with lipids of the same kind, and for a lipid-water interfacial tension \(\varSigma =30\) mJ/m\(^2\), (a) what would be the energy per unit length of an A/B interface? (b) what would be the lowest-energy state for 10 domains, each with average surface 100 nm\(^2\), dispersed in a large membrane patch? how distant in energy are the two configurations? (c) could a transition between the two configurations occur because of thermal fluctuations?
8.5
Membranes with a dimple
Consider a thermal fluctuation making a dimple of height h in a patch \(S=\pi L^2\) of membrane. (a) What is the deformation energy of such a dimple? (b) For a bending constant \(K_b=15 k_BT\), at what h / L ratio the dimple could appear by thermal fluctuation of the membrane surface?
8.6
Pulling chromosomes
An average chromosome can be modelled as an elongated ellipsoid, with major diameter \(b\sim 15 \,\upmu \)m, and minor diameter \(a\sim 1\,\upmu \)m. Observe that during mitosis, each chromosome is pulled by the microtubules by some 15 \(\upmu \)m in about 10 min (such values can vary quite a lot according to the cell type). (a) What is an upper bound to the pulling force? (b) How much work is spent, and how many molecules of ATP are used?
8.7
Pushing cells with a laser
Consider a bacterial cell as a sphere of 2 \(\upmu \)m diameter, and an eukaryote cell ten times larger. A typical laser beam of power P used in optical traps can produce a force of the order of a few pN, as approximated by the expression \(f=nPQ/c\), with n the refractive index of the medium, \(Q\sim 1\)% the quality factor of the laser, and c the speed of light. Estimate the power necessary to move the two cells. What does such a calculation suggest?
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Cleri, F. (2016). Molecular Mechanics of the Cell. In: The Physics of Living Systems. Undergraduate Lecture Notes in Physics. Springer, Cham. https://doi.org/10.1007/978-3-319-30647-6_8
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