Biomedical Engineering Reference
In-Depth Information
the same d x , the number of cross-bridges in subensembles is the same and constant
for any x due to the lack of register between myosin and actin. Assuming that the
cross-bridge can interact only with the closest actin binding site, the state of the
cross-bridges can be described by fractions n j (x,t) giving the fraction of cross-
bridges in state j ( j is one of W ,or S for two state model) at time t in subensemble
at x to x +
d x . Taking that the distance between actin binding sites is d , fractions
n j (x,t) are defined for x in the interval (
d/ 2 ,d/ 2 ) . At any time t , all cross-
bridges are in one of the two states, j n j (x,t)
1. Changes in cross-bridge states
are induced by chemical transition from one state to another or sliding of actin and
myosin filaments relative to each other with the velocity v of sarcomere lengthening.
For example, for state W , this would result in the following governing equation
=
∂n W
∂t +
∂n W
∂x v(t)
=
k SW n S
k WS n W ,
(7.2)
where k WS and k SW are the first order kinetic rate constants for transition from state
W to state S .
The integral properties of the muscle, such as developed stress and ATPase rate
could be found from integration over subensembles (Hill, 1974 ). The Cauchy stress
σ a developed by the cross-bridges in a half-sarcomere is, according to Zahalak and
Ma ( 1990 )
n S (x,t)F S (x) d x ,
d
2
d
2
ml s
d
σ a =
n W (x,t)F W d x
+
(7.3)
2
2
where m is the number of cross-bridges in the unit volume and l s is the length of the
half-sarcomere. According to our assumptions F W is zero because only the strong
binding state generates force. Assuming that F S is proportional to x with Hooke
constant K , the stress equation will have the following form
2
ml s K
d
σ a =
n S (x,t)x d x.
(7.4)
d
2
The average cross-bridge ATP consumption rate is
d
2
k SW n S (x,t)
k WS n W (x,t) d x,
1
d
V AT P =
(7.5)
2
leading to the total ATP consumption per cross-bridge during a beat
T c
d
2
k SW n S (x,t)
k WS n W (x,t) d x d t,
1
d
V beat
AT P
=
(7.6)
2
0
where T c is the period of a beat.
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