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A MECHANICAL
MODEL OF MUSCLE AND ITS
APPLICATION
TO THE INTRAFUSAL
FIBRES OF THE
MAMMALIAN
MUSCLE SPINDLE*
ALAN
CROWE
1. INTRODUCTION
April
2. THE MECHANICALMODEL
The model of the muscle is shown in Fig. 1.
of two elastic components. The
series elastic component has an unstretched
length L, and elasticity E, and the parallel
It consists
1970.
583
A. CROWE
--_
X,
(4)
(5)
= E2X2(s)fI/sX,(s).
(6)
We can eliminate X,(S) and X2(s) from the
above equations and obtain
F(s)=
Fig. 1. Simple three-component
model
of the muscle. The two elastic components
are represented
by springs
having unstretched lengths L, and L,,
moduli of elasticity E, and E,. and are
extended by amounts X, and X,. The
viscous component is represented by a
dashpot having a viscosity V2.
E,tEz+J'z,s,
x(s).
E,+E2+ V,s
The expression
in the { ) brackets is
the transfer function relating a change in
length of the muscle to the resulting tension
change.
The pattern of stretch that is applied to the
model is the ramp shown in Fig. 2. It consists
of two infinite ramps of slope l/T the first
of which starts at time t = 0. From this is
subtracted the second ramp which starts at
1!= T. The Laplace transform of the whole
ramp of unit height is
-icCrt)=&{l-exp
x=x,+x2.
(8)
(2)
F(s)
=E2X,+V,(&).
[-ST]}.
(1)
(7)
(3)
in X2
(9)
is considered
A MECHANICAL
MODEL
Tension ?
Lenqf h
OF
MLISCLE
I/
/Oymmk
: phosei
Static
phose
Laplace
E, 2I:,
F(t)= T(E,ElVz
+E.,)z
[*-eXP[_(5$5)t]]
(10)
In particular, if T is large, the tension at the
end of the dynamic phase is
E,V.,
F(T)=T(E,+E,)+(E,+E~).
(11)
F(t) =
E,Vz
T(E,+E#
x{exp[-(v)(l--T)]
-+E,)
X{exp[-(yqt]]
x (exp [(v)T]_/_
E, E2
T(E,
1)
~5.5
(E,+E,)
12)
= iEEyEi, i .
1 1
(13)
EIV,
T(E, f E.,_ ) .
(14)
586
A. CROWE
to a
3. STIMULATED MUSCLE
V&)
(15)
(16)
= E,"X-X,(E,"+E,"). (17)
(19)
The tension of the system is given by
F(t) = EldX,.
(20)
E,"X,O
> E2*X20.
[-Ct],
(21)
where
(22)
B = E,g(E,dX,o-E~SX20)
(23)
E,"+Efo
and
C=E,*+E~
v*o .
(24)
A MECHANICAL
MODEL
OF MUSCLE
587
parallel elastic component will still be extended, although its unstretched length will
have increased, but it will pull on the viscous
component
until both elastic components
reach their unstretched lengths. The pattern
588
A. CROWE
=A-Bexp
[-Ct]
(25)
where
B,
E,(E,X,O-
E,X,O)
E18+ E,
.
(26)
Comparison
4.1 Tension
4. DISCUSSION
with experimental
development
and single musclejibres
observations.
in whole
muscle
The
isometric
responses
of directly
stimulated single muscle fibres have been
studied by Lannergren
and Smith (1966).
Their records for fast fibres of toad skeletal
muscle are reproduced here (Fig. 5). Quite
clearly in any mechanical system, an instantaneous abrupt change in tension is not
possible and therefore could not be expected
in muscle tissue even if the biochemical
activities which bring about the tension were
A MECHASICAL
MODEL
OF
XIUSCLE
589
stim
Mmq
Length
If
I set
Smm/sec
50 mq
30mmhcc
G_
0200
msec
yG-$i-5
set
5Omg
1
30
5ec
Fig. 5. Isometric
contractile
responses of directlystimulated single muscle fibres.A, D and G responses to
a single stimulus. B, E and H responses to stimulation at
the fusion frequencies indicated. C. F and I responses to
stimulation at 30 set-. Note the rapid rise in tension and,
in the case of records E and H. the rapid fall in tension on
cessation of stimulation.
Records reproduced
from
Lannergren and Smith C1966).
stim
(Fig. 6) is held at zero tension and then stimulated isometrically. Initially there is a sudden
rise in tension and then the rise to the steady
tension is more gradual. This is therefore a
muscle in which E2sX20> E,X,O.
A situation in which the series elastic component does not seem to change its unstretched length is indicated in Fig. 7 which
shows a record for tortoise gastrocnemius
muscle. In this case there is no sudden rise
in tension at the onset of stimulation. The
record also shows the pattern of decay of
tension at the cessation of stimuiation. This
decay follows a roughly exponential course.
It would appear that differences between
fast and slow muscle produce differences in
the value of E,X,O- E,SX,o. The fast muscles
will have a larger value than the slow muscles.
This does not necessarily mean that all fast
muscles will have a positive value. It could be
negative but less negative than for the slow
muscles. For instance, the records of Bulier
Tension _
Length
Fig. 7. The effect on the tension in the tortoise gastrocnemius muscle of stretching while it was contracting (top
record) and in the passive state (middle record). The
isolated nerve-muscle preparation was subjected to a
I mm ramp stretch (bottom record). The tension scale
also applies to the middle record. Record obtained in
collaboration with Dr. A. H. M. F. Ragab.
A. CROWE
590
E*V
=T(E11+&2
{exp
[-(EtzP>t]}
contained in expression (12). If excitation of
the muscle produces a reduction of the term
(E, + E2)/V, then the tension will decay more
A MECHANICAL
MODEL
Fig. 1 to which the elastic region which contains the primary ending is attached in series.
In such a model, changes in unstretched
length at stimulation will cause a previously
passive spindle to start discharging. Also, the
different patterns of response, depending upon
whether a static or dynamic fusimotor fibre is
stimulated when the spindle is held at constant
length, can be explained if it is assumed that
the firing frequency is proportional to the
tension exerted by the muscle. The pattern of
firing will therefore depend upon whether
E1?sXzois greater than or less than E,X,O. As
Crowe and Matthews
(1964a) suggested,
stimulation of a static fusimotor fibre produces
a more rapid contraction than stimulation of
the dynamic fusimotor fibres. If, as has been
suggested above, stimulation of a fast fibre
produces a situation where E,SX,o > Ez8Xzo
then the calculated tension response is as
shown in Fig. 4A, and if stimulation of a slow
fibre produces a situation where E.3sX,o >
E,X,O then the calculated tension response is
as shown in Fig. 4B. If the pattern of firing is
proportional to the tension, then effects similar
to those shown in Fig. 8 will be produced.
5.CONCLUSION
As
pointed
out
by
Pringle
(1960).
the
OF
MUSCLE
591
200
z
<
I00
2
0
Stem
Length
Length
/;--(
Fig. 8. Records showing the discharge of the primary ending of a muscle spindle during a ramp stretch and during
stimulation at 100 set- of a static fusimotor fibre (record A) and at 90 set- of a dynamic fusimotor fibre (record B)
over the periods indicated by the horizontal bars. The records have been chosen to illustrate the patterns of firing
produced at the onset of stimulation of the two types of fusimotor fibre. Records taken from Crowe and Matthews
( 1964a).
8.X-Vol.3.No.6-E
592
A. CROWE