- •Contents
- •Contributors
- •Preface
- •1 Introduction, with the biological basis for cell mechanics
- •Introduction
- •The role of cell mechanics in biological function
- •Maintenance of cell shape
- •Cell migration
- •Mechanosensing
- •Stress responses and the role of mechanical forces in disease
- •Active cell contraction
- •Structural anatomy of a cell
- •The extracellular matrix and its attachment to cells
- •Transmission of force to the cytoskeleton and the role of the lipid bilayer
- •Intracellular structures
- •Overview
- •References
- •2 Experimental measurements of intracellular mechanics
- •Introduction
- •Forces to which cells are exposed in a biological context
- •Methods to measure intracellular rheology by macrorheology, diffusion, and sedimentation
- •Whole cell aggregates
- •Sedimentation of particles
- •Diffusion
- •Mechanical indentation of the cell surface
- •Glass microneedles
- •Cell poker
- •Atomic force microscopy
- •Mechanical tension applied to the cell membrane
- •Shearing and compression between microplates
- •Optical traps
- •Magnetic methods
- •Twisting of magnetized particles on the cell surface and interior
- •Passive microrheology
- •Optically detected individual probes
- •One-particle method
- •Two-particle methods
- •Dynamic light scattering and diffusing wave spectroscopy
- •Fluorescence correlation spectroscopy
- •Optical stretcher
- •Acoustic microscopy
- •Outstanding issues and future directions
- •References
- •3 The cytoskeleton as a soft glassy material
- •Introduction
- •Magnetic Twisting Cytometry (MTC)
- •Measurements of cell mechanics
- •The structural damping equation
- •Reduction of variables
- •Universality
- •Scaling the data
- •Collapse onto master curves
- •Theory of soft glassy rheology
- •What are soft glassy materials
- •Sollich’s theory of SGMs
- •Soft glassy rheology and structural damping
- •Open questions
- •Biological insights from SGR theory
- •Malleability of airway smooth muscle
- •Conclusion
- •References
- •4 Continuum elastic or viscoelastic models for the cell
- •Introduction
- •Purpose of continuum models
- •Principles of continuum models
- •Boundary conditions
- •Mechanical and material characteristics
- •Example of studied cell types
- •Blood cells: leukocytes and erythrocytes
- •Limitations of continuum model
- •Conclusion
- •References
- •5 Multiphasic models of cell mechanics
- •Introduction
- •Biphasic poroviscoelastic models of cell mechanics
- •Analysis of cell mechanical tests
- •Micropipette aspiration
- •Cells
- •Biphasic properties of the pericellular matrix
- •Indentation studies of cell multiphasic properties
- •Analysis of cell–matrix interactions using multiphasic models
- •Summary
- •References
- •6 Models of cytoskeletal mechanics based on tensegrity
- •Introduction
- •The cellular tensegrity model
- •The cellular tensegrity model
- •Do living cells behave as predicted by the tensegrity model?
- •Circumstantial evidence
- •Prestress-induced stiffening
- •Action at a distance
- •Do microtubules carry compression?
- •Summary
- •Examples of mathematical models of the cytoskeleton based on tensegrity
- •The cortical membrane model
- •Tensed cable nets
- •Cable-and-strut model
- •Summary
- •Tensegrity and cellular dynamics
- •Conclusion
- •Acknowledgement
- •References
- •7 Cells, gels, and mechanics
- •Introduction
- •Problems with the aqueous-solution-based paradigm
- •Cells as gels
- •Cell dynamics
- •Gels and motion
- •Secretion
- •Muscle contraction
- •Conclusion
- •Acknowledgement
- •References
- •8 Polymer-based models of cytoskeletal networks
- •Introduction
- •The worm-like chain model
- •Force-extension of single chains
- •Dynamics of single chains
- •Network elasticity
- •Nonlinear response
- •Discussion
- •References
- •9 Cell dynamics and the actin cytoskeleton
- •Introduction: The role of actin in the cell
- •Interaction of the cell cytoskeleton with the outside environment
- •The role of cytoskeletal structure
- •Actin mechanics
- •Actin dynamics
- •The emergence of actin dynamics
- •The intrinsic dynamics of actin
- •Regulation of dynamics by actin-binding proteins
- •Capping protein: ‘decommissioning’ the old
- •Gelsolin: rapid remodeling in one or two steps
- •β4-thymosin: accounting (sometimes) for the other half
- •Dynamic actin in crawling cells
- •Actin in the leading edge
- •Monomer recycling: the other ‘actin dynamics’
- •The biophysics of actin-based pushing
- •Conclusion
- •Acknowledgements
- •References
- •10 Active cellular protrusion: continuum theories and models
- •Cellular protrusion: the standard cartoon
- •The RIF formalism
- •Mass conservation
- •Momentum conservation
- •Boundary conditions
- •Cytoskeletal theories of cellular protrusion
- •Network–membrane interactions
- •Network dynamics near the membrane
- •Special cases of network–membrane interaction: polymerization force, brownian and motor ratchets
- •Network–network interactions
- •Network dynamics with swelling
- •Other theories of protrusion
- •Numerical implementation of the RIF formalism
- •An example of cellular protrusion
- •Protrusion driven by membrane–cytoskeleton repulsion
- •Protrusion driven by cytoskeletal swelling
- •Discussion
- •Conclusions
- •References
- •11 Summary
- •References
- •Index
Multiphasic models of cell mechanics |
89 |
To date, there has been only limited application of the triphasic model to cell mechanics (Gu et al., 1997). At mechanochemical equilibrium in vitro, many cells are known to exhibit a passive volumetric response corresponding to an ideal osmometer. For an ideal osmometer, cell volume and inverse osmolality (normalized to their values in the iso-osmotic state) are linearly related via the Boyle van’t Hoff relation. The resulting states of mechanochemical equilibrium of the cell exhibit an internal balance between mechanical and chemical stresses. The triphasic model, in the absence of electrical fields, gives rise to the mixture momentum equation:
· σ = 0, where: σ = − pI + σE |
(5.11) |
where σ is the mixture stress and σE is the extra stress in the solid phase. The balance of electrochemical potentials for intracellular and extracellular water gives rise to the Donnan osmotic pressure relation:
p |
= |
RT (ϕc |
− |
ϕ c ) |
(5.12) |
|
|
|
|
where T is the temperature, ϕ and ϕ are the intracellular and extracellular osmotic activity coefficients, and c and c are the intracellular and extracellular ion concentrations, respectively (R is the universal gas constant). To close the system of governing equations (Eqs. 5.11–5.12), the intracellular extra stress and the intracellular ion concentration need to be characterized. While the former may be postulated via a constitutive description for the subcellular components (such as nucleus, cytoskeleton, membrane), the latter necessitates a detailed analysis of electrochemical ion potentials inside a cell that accounts for intracellular ionic composition and biophysical mechanisms such as the selective permeability of the bilayer lipid membrane, and the nontransient activity of ion pumps and ion channels.
The strength of this type of mixture theory approach was recently illustrated in a study modeling the transient swelling and recovery behavior of a single cell subjected to an osmotic stress with neutrally charged solutes (Ateshian et al., 2006). A generalized “triphasic” formulation and notation (Gu et al., 1998) were used to account for multiple solute species and incorporated partition coefficients for the solutes in the cytoplasm relative to the external solution. Numerical simulations demonstrate that the volume response of the cell to osmotic loading is very sensitive to the partition coefficient of the solute in the cytoplasm, which controls the magnitude of cell volume recovery. Furthermore, incorporation of tension in the cell membrane significantly affected the mechanical response of the cell to an osmotic stress. Of particular interest was the fact that the resulting equations could be reduced to the classical equations of Kedem and Katchalsky (1958) in the limit when the membrane tension is equal to zero and the solute partition coefficient in the cytoplasm is equal to unity. These findings emphasize the strength of using more generalized mixture approaches that can be selectively simplified in their representation of various aspects of cell mechanical behavior.
Analysis of cell mechanical tests
Similar to other experiments of cell mechanics, the analysis of cell multiphasic properties has involved the comparison and matching of different experimental