session 14 -...
TRANSCRIPT
Session 14
� Stability of a structure based on a balanced synergy
of continuous tension and discrete compression
components
� Richard Buckminster Fuller coined tensegrity as
“islands of compression inside an ocean of tension”
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“islands of compression inside an ocean of tension”
� Common cellular interpretation
� Actin microfilaments – tension elements
� Microtubules – compression elements
Additional components
3
� Additional components
� ECM – compression element
� Intermediate filaments – tension elements
� Cross-linked actin bundles – compression elements
� Central to tensegrity is the amount of pre-loaded
tension in the structure (prestress).
� Dictates the stiffness and mechanical response
� Cells have prestress that is balanced by ECM
attachments
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attachments
� Detach the cell from culture
dish and it rounds up
� Akin to cortical tension in
lipid drop models
Trypsin added to detach cell
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Here, B-B’ �C-C and C-C’ � B-B
� Element dimensions
� sx sy sz strut-to-strut separation
� l1 l2 l3 tension lengths
� LA LB LC strut lengths (=1)
Recall vector notation:
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� Recall vector notation:
� Vector
� Distance
� Direction
2 2 2
ˆˆ ˆ
ˆ /
i
i
i i i
l xi yj zk
l l x y z
l l l
= + +
= = + +
=
�
�
�
� Based on model geometry
l
( ) ( ) ( )
2 2 2
22 21 1 1 1
1 2 2 2 2
2 212 2
i
x y
l x y z
l s s
l s s s
= + +
= + − +
= + − +
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� Similarly½sx
½½
l12 2
1
12 2
2x y y
l s s s= + − +
½sy
2 2
2
2 2
3
12 2
2
12 2
2
y z z
z x x
l s s s
l s s s
= + − +
= + − +
� Balance of forces on A-A
T
F3
F1
( ) ( )
,
3 1
3 3 1 1
0
ˆ ˆ4 4 0
ˆ ˆˆ ˆ4 4 0
x AAF
T F i F i
T F l i F l i
=
+ ⋅ + ⋅ =
+ ⋅ + ⋅ =
∑� �
ˆi i i
F Fl=�
A
By symmetry,
2 of 8 tension
forces shown
By symmetry,
2 of 8 tension
forces shown
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½sx
½½
l1
½sy� Similarly
( ) ( )1 1 1
2 2 2
3 1
3 1
4 4 0x xs s
T F Fl l
−+ − =
( ) ( )
( ) ( )
1 2
,
1 2
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,
2 3
0 1
0 1
y BB y y
z CC z z
F FF s s
l l
FFF s s
l l
= ⇒ − =
= ⇒ − =
∑
∑
� Struts are slender with zero transverse loading
� Cables are linear elastic springs
where k is spring stiffness and lR is unstressed cable length
( )i RF k l l= −
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R
� Define prestress (strain)
where l0 is uniform initial length of each cable for an initially
isotropic structure
0
0
Rl l
lξ
−=
� At rest, structure is isotropic, therefore
0 1 2 3
1
1/ 2
0.375
A B C
x y z
L L L
s s s
l l l l
= = =
= = =
= = = =
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� For given applied force T and prestress (ξ , k, lR)
� Have 6 unknowns (∆sx ∆sy ∆sz ∆l1 ∆l2 ∆l3)
� Have 3 geometric length equations and 3 force
balance equations
� Can solve…
� Structural stiffness
� High prestress (ξ), stiffness is constant
x
TE
s≡
∆
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constant
� Tensegrity structure is Hookean
� Low prestress, stiffness increases with applied force
� Strain hardening seen by Petersen (1982) and Radmacher (1996)
� Stiffness increases with prestress
� Prestress:� Traction forces for cells on gels
embedded with fluorescent microbeads
� Young’s Modulus� Cell stiffness measured with magnetic
twisting cytometry
� Tensegrity predict linear relationship between prestress and effective
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between prestress and effective stiffness
� At top, slope is 0.4 but tensegrity predicts slope to be 1
� At bottom, dashed-line slope is 1.04.� Accounting for 14% loading in MTs not
measurable by traction forces
Adapted from Wang, N. (2001) PNAS, 98:7765
Adapted from Wang, N. (2002) Am. J. Physiol.
Cell Physiol, 282:C606
� Class Discussion� Mechanotransduction shows that cells adapt to stress
� Tension only, where is compression? Not only MT, could be ECM.
� Cells are fluid filled, how does that relate to tensegrity?
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tensegrity?
� Dampening consideration
� What about AFM? Discrete points have high stiffness at struts, low stiffness at springs
� FEM is more powerful to model cell mechanics
� But disregards the mechanical nature of each protein (filaments)
� Tensegrity: local perturbation results in global rearrangement� Local load should cause distal disturbance
� Continuum models show local response � Dissipate inversely with distance from point
of load
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of load
� Examples� Mantiotis (1997) pulling on ECM coated bead
causes nuclear distortion
� Hu (2003) used MTC and MitoTracker and observed discrete displacements at distal pts
� Both observations were actin-CSK dependent