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Biomedical Engineering – From Theory to Applications

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472<br />

Nb<br />

b b l<br />

l1<br />

<strong>Biomedical</strong> <strong>Engineering</strong> <strong>–</strong> <strong>From</strong> <strong>Theory</strong> <strong>to</strong> <strong>Applications</strong><br />

1 1 nl1 nl2<br />

W k L<br />

2 1n<br />

n<br />

(3)<br />

l1 l2<br />

The resistances <strong>to</strong> changes in the surface area of the whole membrane and <strong>to</strong> an area change<br />

of a local element are both modelled. The former corresponds <strong>to</strong> the situation whereby lipid<br />

molecules can move freely over the cy<strong>to</strong>skeletal network, while the latter corresponds <strong>to</strong> the<br />

situation where movement of the lipid molecules is confined <strong>to</strong> a local element. The area<br />

expansion energy WA is thus formulated as a summation of the energy due <strong>to</strong> a change in<br />

the whole membrane area and due <strong>to</strong> a change in the local area:<br />

2<br />

N<br />

2<br />

1 e<br />

A A0 1 Ae A0e<br />

<br />

A A 0 a 0e<br />

2 A0 2 e1<br />

A0e<br />

W k A k A<br />

(4)<br />

<br />

where A is the area of the whole membrane, subscript 0 denotes the natural state, and kA is a<br />

coefficient for the whole area constraint, Ae is the area of the element, ka is a coefficient for<br />

the local area constraint, and Ne is the number of elements. The <strong>to</strong>tal elastic energy s<strong>to</strong>red is<br />

thus expressed as:<br />

where j denotes CM (j = c) and NE (j = n).<br />

j j j j<br />

s b A<br />

(a)<br />

(b) n l1<br />

W W W W<br />

(5)<br />

Fig. 3. (a) Mesh of the cellular membrane and nuclear envelope and (b) mechanical model of<br />

the cell membrane.<br />

2.3 Modelling of CSK<br />

As demonstrated in various studies (Wang, 1998; Nagayama et al., 2006), CSKs play a<br />

pivotal role in cellular mechanics. The CSK consists primarily of actin filaments,<br />

microtubules, and intermediate filaments (see Fig. 4). Here, these were modelled as CSK<br />

regardless of the type of cy<strong>to</strong>skeletal filament. For simplicity, a CSK is expressed as a<br />

straight spring that generates a force as a function of its extension. The energy WCSK<br />

generated is thus modelled as<br />

k s<br />

NCSK<br />

1<br />

2<br />

CSK CSK ( i 0i<br />

)<br />

2 i1<br />

i r i<br />

θ l<br />

l<br />

k b<br />

e<br />

n l2<br />

W k l l<br />

(6)<br />

where kCSK is the spring constant of the CSK, l0i and li are the length of CSKi at the natural<br />

state and after deformation, and NCSK is the <strong>to</strong>tal number of CSKs.

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