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Cellular Engineering

of RBC Membrane

Micropipette Aspiration

and Shear Flow

Scanning Electron Micrograph

of a Human Red Blood Cell

The Red Blood Cell Membrane

Approaches for Measuring

Mechanical Properties of Cells

Apply known force, measure deformation

Indentation experiment

Micropipette aspiration

Flow chamber

Magnetic twisting cytometry

Red blood cell membrane flickering

Thermal fluctuations of actin filaments and microtubules

Approaches for Measuring

Mechanical Properties of Cells

Apply known force, measure deformation

Indentation experiment

Micropipette aspiration

Flow chamber

Magnetic twisting cytometry

Red blood cell membrane flickering

Thermal fluctuations of actin filaments and microtubules

Approaches for Measuring

Mechanical Properties of Cells

Apply known force, measure deformation

Indentation experiment

Micropipette aspiration

Flow chamber

Magnetic twisting cytometry

Red blood cell membrane flickering

Thermal fluctuations of actin filaments and microtubules

Measuring Shear Modulus

of RBC Membrane

Difficulties

Small size and complex geometry

Non-uniform and time dependent stress/strain

Reversed Problem Approach

Assume a stress-strain relationship with unknown coefficients

Solve the boundary value problem

Fit the solution to the data to test the validity of the model

Start over with a new model if the test result is negative

Evaluate the parameter values if the test result is positive

Problem

Parameter values may be model- and technique-dependent

Solution

Conduct separate experiments using distinct techniques

Mechanical Properties of

Human Red Blood Cells

Mechanical model for the human red blood cell

A liquid bag enclosed by a membrane

The liquid can be modeled as a Newtonian viscous fluid

The membrane can be modeled as an elastic sheet

Membrane shear modulus: = 6-9 pN/m = 6-9 10-6 N/m

Membrane area modulus: K = 4.5 105 pN/m = 0.45 N/m

Membrane bending modulus: B = 44 5 kBT

Hemoglobin solution viscosity : = 6 10-3 N-s/m2

Force balance (Law of LaPlace)

Micropipette p0 p = 2Tm/Rp

Aspiration

Neglecting friction at the pipette wall

Experiment

T = Tm

T RBC membrane

Model as an

Neglect infinite sheet

constraint of elastic

membrane

of the p0

Tm r

2Rp p

RBC

Closed

Polar coordinates

shape

T

Stress Analysis of an Elementary Membrane Segment

Tr + dTr dr Re-arranging terms:

d/2 d/2 - dTr = (Tr - T)dr/r

Axisymmetric Integrating:

Problem T + dT rd T - Tr|r + Tr|r = Rp

Tr

= Rp (Tr - T)dr/r

Tr and T are 0

functions of r r LaPlace law:

only Tr|r = Rp = pRp/2

d

Re-arranging terms:

p = (2/Rp) Rp (Tr - T)dr/r

(Tr + dTr)(r + dr)d - Trrd sin(d/2)Tdr - sin(d/2)(T + dT)dr = 0

rTr/r + Tr - T = 0

Force balance in the circumferential direction:

cos(d/2)(T + dT)dr - cos(d/2)Tdr = 0

T/ = 0

Analysis of Deformation

r0

Area prior to Hemi-spherical cap:

deformation r

1 = 2Rp2

0 = r02 Cylindrical segment:

mapping 2 = 2Rp(L Rp)

Area remains Circular ring:

r0

constant 3 = (r2 Rp2)

r

during

deformation Total area:

0 = = 1 + 2 + 3

Area modulus K = 4.5 105 pN/m >> shear modulus = 6-9 pN/m

Constant area: r02 = 2Rp2 + 2Rp(L Rp) + (r2 Rp2)

Radial stretch ratio: r2 r02/r2 = 1 + (Rp2/r2)(2L/Rp - 1)

Differentiation: dr2/(1 - r2) = 2dr/r

Boundary conditions: r2|r = 1 r2|r = Rp = 2L/Rp

Stress-Strain Relation

r, r

Stresses or

, ,

Strains or

( r)/2 (2 r2)/2

Stretch ratios

Assume linear elasticity:

Tr - T = (r2 2) = (r2 r-2)

Re-arranging terms:

Tr = (r2 2)/2 T = (2 r2)/2

Solution

p = (2/Rp) Rp (Tr - T)dr/r

= (2/Rp) Rp (r2 r-2)dr/r

1 p

= (/Rp) 2l/Rp (r2 r-2)dr2/(1 - r2)

Slope = 2/Rp

1

= - (/Rp) 2l/Rp (1 + r-2)dr2

= (/Rp)[2L/Rp - 1 + ln(2L/Rp)] L/Rp

Why or why not?

Measuring the BRC Shear Modulus

by the Flow Chamber

dx

Equilibrium:

2d(Txy) = dA

v 2yTx + Integration:

2y 2yTx

d(2yTx) T = A/y

x 0

0dA

dA0 dA

y0 y

0

x0 dx0 x dx

mapping

(x0,y0) (x,y)

Analysis of Deformation

x0 = R0 (1 - cos0) Asector = R020

R0

y0 = R0sin0 Atriangle = R0sin0 R0cos0

0

A0 A0 = R02(0 - sin0 cos0)

cos0 = 1 - x0/R0

0 = cos-1(1 - x0/R0) Constant area:

R0cos0

sin0 = [1 (1 - x0/R0)2]1/2 A = A0 = R02{cos-1(1 - x0/R0)

(1 - x0/R0)[1 (1 - x0/R0)2]1/2}

Stretch ratios: x = dx/dx0 y = y/y0

Constant area: xy = 1 y = x-1

Expressing y as a function of x0:

y = y0y = R0sin0/x = R0sin[cos-1(1 - x0/R0)](dx0/dx)

Stress-strain relation: Tx = (/2)(x2 x-2)

Re-arranging: (/2)[(dx/dx0)2 (dx/dx0)-2] = 0A/y

= 0R02{cos-1(1 - x0/R0) (1 - x0/R0)[1 (1 - x0/R0)2]1/2}/{R0sin[cos-1(1 - x0/R0)]}(dx/dx0)

(dx/dx0)4 (0R0/)(dx/dx0)3{cos-1(1 - x0/R0)[1 (1 - x0/R0)2]-1/2 - (1 - x0/R0)} = 1

Comparison to Experiment

Calculated shapes resemble those observed

10 m 10 m

Comparison to Experiment

The predicted 0R0/ vs. L/L0 curve is nearly linear

0R0

Slope = Stheor

Slope = Sexp

L/L0

= Cexp/Ctheor

Summary of Steps

Deformation mapping from the undeformed to the

deformed shape, analysis of strains or stretch ratios

Stress-strain relation constant area, linear elasticity

Solving the math problem

Comparison to experiment

Red Blood Cell Membrane Undulations

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