REVIEW ARTICLE
SKELETAL MUSCLE ADAPTABILITY.
I : REVIEW OF BASIC PROPERTIES
Richard L. Lieber
The muscular system is one of the bodys
most adaptable organs. The clinician is
often in the position of repairing an
adapted, abnormal muscle, e.g. after
limb immobilization, tenotomy, chronic
exercise, chronic electrical stimulation,
mechanical overload, denervation, upper
or lower motoneuron lesion, myopathy,
cross-innervation and reinnervation. All
I- of these conditions can cause muscle
s adaptation as a result of change in (1) the
0
s amount and type of nervous-system
00- input; (2) the degree of muscle use; (3) the
c1
stress on the muscle; and/or (4) the
\o-
m muscle length. Treatment of an affected
2
muscle can be improved by understanding
$
c the nature of muscular adaptation.
In order to understand the adaptation
2 of muscle to a particular environmental
2 change, an understanding of normal
0
ir: muscle structure and function is
d necessary. Skeletal muscle is a highly
p1 organized tissue at the structural and
.s
g functional level. The purpose of the first
s2
c
paper in this series is to review basic
skeletal muscle structure and function.
z
22
Subsequent reports will provide examples
of the ways in which skeletal muscle
adapts to specific environmental changes
4 such as limb immobilization, upper
or lower motoneuron lesion, and
390 functional electrical stimulation (FES).
Skeletal muscle structural hierarchy
Skeletal muscle represents a classic
biological example of the relationship
between structure and function. There-
fore discussions of muscle function are
intimately tied to discussions of muscle
structure (Huxley 1974, Squire 1981). The
elegant structural hierarchy of skeletal
muscle is presented in Figure 1. (Note that
a given muscle may take on almost any
size or shape.) Variability in this gross
structure does not result from different
muscles being constructed of different
components: in fact all muscles are
composed of single muscle fibers which
are almost identical in diameter. While
the length and arrangement of these
individual fibers (i.e. the architecture, see
below) may vary between muscles, the
fibers themselves have a diameter of 20 to
80pm in mammalian muscle.
A whole muscle can thus be viewed as
a number of muscle fibers arranged in
series and in parallel (Fig. 1A). The
muscle fiber itself is composed of 500 to
10,ooO myofibrils arranged in parallel,
with diameters of about lpm (Fig. 1B).
The myofibril is composed of 1000 to
2,000,000 sarcomeres arranged in series,
which are 2 to 3pm in length and about
1pm in diameter (Fig. 1C). The sarcomere
is composed of a number of inter-
 m
N

Fig. 1. Structural hierarchy in skeletal muscle. (A) Whole muscles

are composed of single muscle fibers arranged in series and in
parallel. (B)Single muscle fibers are composed of myofibrils
arranged in parallel. (C) Myofibrils are composed of sarcomeres
arranged in series. (D)Sarcomeres are composed of interdigitating
thick and thin filaments, which in turn are composed of contractile
and regulatory proteins.

Fig. 2. Schematic example of the

functional significance of muscle
architecture: (A) a single sarcomere
which generates one unit of force and
one unit of ,velocity; (B)two sarcomeres
arranged in series to yield a muscle,
which contracts with two units of
velocity and one unit of force; and (C)
two sarcomeres arranged in parallel to
yield a muscle, which contracts with
two units of force and one unit of
velocity. 391
kj
9 digitating protein filaments (the so-called
;3" 'thick' and 'thin' filaments) which are
4 responsible for the regulation and
'D generation of muscle force (Fig. 1D).
a
-c Thus the sarcomere (actually the half-
z sarcomere due to sarcomere symmetry) is
.-*c the functional unit of force generation in
skeletal muscle. Structural hierarchy is
a
e also demonstrated at the molecular level,
.-0 as the individual thick filaments are
-
m3 composed of a number of myosin
molecules arranged in an antiparallel
-B
.->;
4- fashion. The thin filament is composed of
a series of actin monomers arranged in a
a
I helical pattern. One regulatory protein
a (tropomyosin) is inserted along the groove
-cY
e,
of the helix and the other (troponin) at
discrete locations along the actin
-E filament. These regulatory proteins
m
-
4-
2
modulate the interaction between the
actin and myosin filaments and thus also
vl
modulate the force generated by the
muscle (Squire 1981).
The process by which the myofilaments
are activated is known as excitation-
contraction coupling (Ebashi 1976).
Following nerve depolarization, an action
potential propagates down the moto-
neuron and is transmitted to the muscle
fiber at the neuromuscular junction by
diffusion of the neurotransmitter
acetylcholine. The resulting depolar-
ization is conducted across the muscle via
the surface membrane (sarcolemma),
which is itself an excitable membrane.
The action potential is then conducted
deep within the muscle fibers via the
transverse tubular system, or T-SyStem,
which runs perpendicular (or transverse)
to the myofilaments. The action potential
from the T-SyStem then signals the
sarcoplasmic reticulum (SR) membrane
system by an unknown mechanism to
release calcium, causing force generation.
Relaxation occurs as the calcium is
pumped back into the SR by the calcium
pump localized within the SR membrane.
Skeletal muscle architecture
As presented above, the sarcomere can be
considered the functional unit of force
generation in muscle. In order to
understand the contractile properties of
whole muscle it is necessary to appreciate
the significance of the three-dimensional
392 arrangement of these sarcomeres. The
crux of this concept is that whole-muscle
force is proportional to the number of
sarcomeres acting in parallel, while
whole-muscle velocity is proportional to
the number of sarcomeres acting in series
(Gans and Bock 1965, Gans 1982, Sacks
and Roy 1982). As an illustration, let us
assume (as shown in Figure 2A) that when
a single sarcomere is activated it generates
one unit of force and one unit of velocity.
Thus if a muscle were composed of two
sarcomeres arranged in series (Fig. 2B) it
would contract with two units of velocity
(two sarcomeres in series) and one unit of
force (one sarcomere in parallel).
Conversely, if a muscle were composed of
two sarcomeres in parallel (Fig. 2C) it
would contract with one unit of velocity
(one sarcomere in series) and two units of
force (two sarcomeres in parallel).
This concept is easily generalized to
the case of the whole muscle, where we
usually consider muscle fiber length and
muscle fiber arrangement instead of
sarcomere arrangement (Fig. 3). Obvi-
ously the longer the fibers, the more
sarcomeres in series, and therefore the
greater the contraction velocity. Simi-
larly, the more fibers in parallel, the
greater the muscle force because of the
increased number of sarcomeres in
parallel. Thus many muscles can be
assigned into one of two classes: those
designed for high-force output (e.g.
vastus muscles, deltoid and gas-
trocnemius) and those designed for high-
velocity output (e.g. sartorius, orbicularis
oris and brachialis) (Brand et al. 1981,
Wickiewicz et al. 1983, Powell et al.
1984). For high-velocity output the
muscle fibers are aligned along the axis of
force generation and are very long
(thereby increasing the number of
sarcomeres in series (Fig. 3A)). For high-
force output, in order to increase muscle-
fiber cross-sectional area while conserving
space, the muscle fibers may take on a
pinnated arrangement at a fixed angle
relative to the axis of force generation
(thereby increasing the number of
sarcomeres in parallel (Fig. 3B)), or the
muscle may exhibit pinnation over a range
of angles relative to the axis of force
generation (Fig. 3C).
Based on this discussion, it should be
apparent that differences in muscle
 A. C. F
m
rn

- Bone d
m
Q\

W-
-Tendon N

2,
4
e
Muscle
-Tendon z

Fig. 3. Example of whole-muscle architecture demonstrating different fiber arrangements. Average fiber
length is shown by the line labelled FL. Average fiber cross-sectional area is shown by the line labelled
CSA. Muscle contraction velocity is proportional to the length of line FL, whereas contraction force is
proportional to the length of line CSA. ( A )Longitudinal architecture designed to provide high contraction
velocify. (B)Pinnated architecture with the fibers oriented at a fixed angle relative to the axis of force
generation, in order to provide high contractile force while conserving space. Note that this muscle would
generate more force and contract at a slower velocity than the muscle shown in A. ( C ) Pinnated
architecture with the fibers oriented at varying angles relative to the axis of force-generation. This muscle
would generate more force than the muscles in either A or B, but would contract at a slower velocity than
either muscle. On inspection, all three muscles would appear to be of similar size, but normalization of
force or velocity to gross muscle size would be misleading.

architecture are not always obvious on
gross inspection. Therefore in muscle
transplant or transfer operations, when
the transferred muscle is to be used as a
synergist to existing muscles, efforts
should be made to match architectural
properties in terms of fiber length and
total fiber cross-sectional area. Without
consideration of these architectural
properties the resultant muscle action
(force or velocity) may be inappropriate.
Skeletal muscle functional hierarchy
To this point, it is implied that all muscle
fibers are created equal. Obviously this is
a gross oversimplification. While muscle
architecture has the greatest influence on
whole-muscle contractile properties, there
is also a hierarchy of functional muscle-
fiber properties which permits fine
tuning of muscle function to a particular
task. This hierarchy is manifested by the
distribution of fiber types and motor units
within a muscle.
The motor unit is defined as the alpha-
motoneuron and the muscle fibers which
itinnervates. While controversy still exists
as to the precise properties of the motor
unit. eeneralizations can be made. Motor
units can be distributed into three main
classes: s (which stands for slow), FF
(which stands for fast fatigable), and FR
(which stands for fast fatigue-resistant)
(Burke et al. 1973). It is apparent,
therefore, that motor units are classified
according to the muscle-fiber properties
and not the alpha-motoneuron properties.
Several workers have shown that the
muscle fibers within the motor units can
also be grouped into three main classes
(McDonagh et al. 1980a, b; Burke 1981).
The classification scheme implemented
depends on the species of animal (Brooke
and Kasier 1970, Peter et al. 1972,
Dubowitz and Brooke 1973) but for
humans these three main classes are
type 1 (the physiologically slow muscle
fibers); type 2A (the physiologically fast
fibers with high oxidative and glycolytic
capacity); and type 2B (the physio-
logically fast fibers with low oxidative
capacity and high glycolytic capacity).
Fiber types (1, 2A and 2B) generally are
determined histochemically, while motor-
unit types (s, FF and FR) are determined
according to the contractile properties
resulting from intracellular stimulation of
a single ahha-motoneuron. (Therefore 393
ki
9 one should not refer to fast muscle
5 fibers from a histochemical study or
4 type 1 fibers from a physiological
h study.) It is generally assumed that there
s is a correspondence between motor units
g
-c
and muscle fibers, i.e. s motor units are
8
.c composed of type 1 fibers, FR motor
ii
2
units are composed of type 2A fibers, and
FF motor units are composed of type 2B
a animals and to manipulate, has been
.-
0 muscle fibers.
-.-
3
a
.->;
Motoneuron innervation to muscle
fibers is also highly organized. Small
alpha-motoneurons innervate slow
.n.
.- (type 1) muscle fibers, while the very large
a
Y motoneurons innervate fast (type 2)
id
a muscle fibers. Activation threshold is
4
-::
6J lowest for small motoneurons. Thus at
low exertion levels (e.g. during the early
5... phase of a movement), slow muscle fibers
-
m
c
24
are activated first. Type 2A fibers are
activated next, and type 2B fibers are
v)
activated only in the maximal efforts due
to the large caliber of their alpha-
motoneurons.
The precise co-ordination which exists
between motoneuron and muscle-fiber
properties is not coincidental, but results
from the significant influence that
motoneurons have on the muscle fibers
they innvervate (Pette and Vrbova 1985).
Slow muscle fibers (type 1) are regularly
activated at relatively low levels (i.e. tonic
activation), while fast muscle fibers (type
2) are intermittently activated at relatively
high levels ( i e . phasic activation) (Freund
1983). The muscle-fiber properties
become tailored to the activation,
resulting in a relatively homogeneous
population of muscle fibers within a
single motor unit (Burke et al. 1973,
Burke 1981). Therefore, slow muscle
fibers, which are adapted to tonic
activation, demonstrate a Iarge capacity
for oxidative metabolism, are generally
smaller in diameter, facilitating oxygen
diffusion from the many capillaries
surrounding them, and have a poorly
developed SR and T-system. Conversely,
fast fibers, which are adapted to phasic
activation, are rich in glycolytic enzymes,
and have a highly developed SR and T-
system, enabling the rapid release and
sequestration of calcium needed for rapid
contraction and relaxation. Measurement
of skeletal-muscle contractile, histo-
394 chemical and mornhological Dronerties
thus provides insights into muscle-
activation history.
Skeletal muscle contractile properties
The purpose of skeletal muscle is to
generate force and/or movement, so
often it is useful to measure these
quantities experimentally. Muscle, being
one of the easiest tissues to remove from
studied for over 100 years (Hill 1953,
1970). Two relationships have been
defined which characterize a muscles
ability to generate force and/or
movement: these are (1) the length-
tension relationship and (2) the force-
velocity relationship.
Length-tension relationship
The length-tension relationship states that
isometric muscle force (i.e. force
developed at constant length) varies as a
function of muscle length. The length-
tension curve is obtained by stimulating a
muscle maximally and measuring the
force developed at discrete muscle
lengths. This muscle property can be
explained by the structural hierarchy
presented above. A whole muscle is
composed of billions of sarcomeres in
series and in parallel, so whole-muscle
properties are amplified (and smoothed)
sarcomere properties. The classic length-
tension relationship of a single muscle
fiber which relates force to sarcomere
length is shown in Figure 4 (Gordon et al.
1966a, b). This curve demonstrates that
the force developed is directly pro-
portional to the number of available
interaction sites between the actin and
myosin filaments (Huxley and Hanson
1954, Huxley and Niedergerke 1954).
While controversy still exists regarding
the exact form of this relationship
(Pollock and Sugi 1984) it is agreed that
force varies as a function of muscle
length. This fact obviously has profound
implications for the surgeon who must
reattach a muscle at the optimum length
secondary to trauma or tendon transfer.
Two measures characterize the muscles
ability to generate force. The first, LO, is
the length at which the muscle generates
the maximum force. (It is important to
compare forces between muscles with the
muscle length set to L n to ensure that the
muscle is generating maximum force.)
The second, PO, is the maximum force
developed by the muscle at LO. p0 can be
expressed in absolute units (e.g. Newtons
or grams) or normalized units (e.g.
Newtons/cm2 or grams/gram weight of
muscles). Obviously p0 expressed in
absolute units will vary with muscle size,
and therefore animal size. To make
effective comparisons between individuals
or between muscles, the force must be
normalized. Based on the discussion of
architecture, it is clear that normalization
to muscle weight alone will not permit
effective comparison between muscles of
different architecture. It is possible to
have two muscles which have identical
weights but generate dramatically
different forces because of their different
fiber arrangements. The best way to
normalize muscle force is to express the
force relative to the total muscle-fiber
(not total muscle) cross-sectional area
(Gans 1982, Sacks and Roy 1982, Powell
et al. 1984). Several investigators have
presented indirect methods for calculating
this so-called physiological cross-
sectional area.
Force-velocity relationship
The force-velocity relationship states that
steady-state isotonic velocity (i.e.
contraction velocity against a constant
force) varies as a function of muscle
force. The force-velocity curve is obtained
by stimulating a muscle maximally,
allowing it to experience a constant force,
and measuring the initial contraction
velocity at each force level. This
relationship takes the form of a
rectangular hyperbola (Hill 1953). The
force-velocity relationship not only
characterizes normal muscle but also
differentiates between muscles with
different fiber-type distributions (Close
1972). The force-velocity property of
muscle results from the physical and
biochemical properties of the muscle
force-generators (i.e. the cross-bridges on
the myosin filaments). Several models
have been proposed which define cross-
bridge properties such that the whole-
muscle force-velocity relationship is
obtained (Huxley 1957, 1974). Present
mechanical studies of muscle are
modifying the description of these cross-
8 5 4 3 2 1
80
I
Striation spacing (3
-4 3.65p (a+b) -------+
1 d - A
42.20-2.25p (b+c)+
2- ti
c- 2 . 0 5 ~(b) --b
3- R R 4
4.85-1.QOP(b-ch
- 4
Fig. 4. Sarcomere length-tension curve. (Upper)
Curve representation of relativeforce generated as
function of sarcomere length. (Middle) Schematic
representation of relevant dimensions of
sarcomere: a = length of thick myosin filament
( I .60pn); b = length of thin actin filament
(2.05pm); c=length of bare zone of myosin
filament (0-25/tnt), which contains no cross-
bridges; z = width of z-band (0.05pn). (Lower)
Schematic representation of appearance of
sarcomere at various sarcomere lengths, which
explains length-tension shown above. 1 =No force
is generated because there are no cross-bridges
connected between actin and myosin;
2 = maximum force is generated because all
possible cross-bridges are connected between actin
and myosin; 3 = tips of opposing actin filaments
are juxtaposed without interfering with one
another; 4 and 5 =force decreases because actin
filaments begin to interfere with formation of
cross-bridge connections on opposite side of
sarcomere; 6 =force drops to near zero because
myosin filaments have perforated z-band or
broken off as result of compression by z-band.
Steps I to 5 represent reversible stages of
contraction; step 6 represents irreversible stage of
supercontraction. (From Gordon et al. 1966b;
reproduced by permission.)
395
 bridge kinetic properties in order to
explain the transient mechanical
4 phenomena in muscle which are observed
on a very fast time-scale (Huxley and
Simmons 1971, Ford et al. 1977, 1980).
As with the length-tension relationship,
a factor can be extracted from the force-
velocity relationship which characterizes
the muscle's ability to produce
.-u movement. This is v,,,, the maximum
1 muscle contraction velocity. As with
m molecule.
length-tension properties, it is often
useful to compare Vmax between muscles.
v,,, can be expressed in absolute units
(e.g. mm/sec) or normalized units (e.g.
muscle lengthdsec, fiber lengthslsec, or
sarcomere lengthdsec). Vmax, expressed in
absolute units, will vary with muscle
length, and therefore animal size (Hill
1970). To compare muscles or individuals
effectively, the velocity must be
normalized. Based on the discussion of
architecture, normalization to muscle
length alone will not permit effective
comparison between muscles of different
architecture, as muscle fibers do not
necessarily run the entire muscle length. It
is possible, therefore, to have two muscles
which have the same length, but because
of architectural difference, will contract
at dramatically different velocities. The
best way to normalize Vmax is to express
the velocity relative to total muscle-fiber
(not total muscle) length. Muscle-fiber
length is generally determined by
microdissection of individual fibers from
a formalin-fixed muscle (Gans 1982,
Sacks and Roy 1982).
It is technically difficult to perform
force-velocity experiments which yield the
useful measure Vmax. In normal muscle, it
has been demonstrated that the time-
course of an isometric twitch (a
contraction resulting from a single
stimulus delivered to the muscle at
constant length) is proportional to Vmax
(Barany 1967). Thus by measuring the
time required to reach the peak of an
isometric twitch i.e. time-to-peak tension,
or TPT, and/or the time required for
relaxation during an isometric twitch i.e.
half-relaxation time, or HRT,it is possible
to estimate dynamic muscle properties. It
should be noted, however, that the time-
course of an isometric twitch is related to
396 the quantity of T-system and the quantity
and activity of SR calcium-transport
proteins, while vmaxis related to the
ATPase activity of the myosin molecule
itself (Barany 1967). While TPT, HRT
and v,,, are correlated, they are
manifestations of different muscle
properties, so it is possible to demonstrate
dramatic changes in TPT and HRT,
without any change in Vmax. This is
because the SR and T-SyStem adapt much
more readily than does the myosin
Contractile properties of muscles can
be compared according to experimentally
obtained values for Lo, PO,TPT, HRT and
vmaX,which are sensitive to changes in
architecture, fiber-type distribution and
fiber-size distribution. For example, given
two muscles of roughly equal size and
architecture but different fiber-type
distributions, values for LO would be
identical, as fast and slow muscle fibers
apparently have identical length-tension
properties. However, the force generated
by a slow muscle fiber is probably less
than the force generated by a fast muscle
fiber (McDonagh et al. 1980b, Burke
1981, Powell et al. 1984). Therefore POof
a muscle composed mainly of slow fibers
will be less than PO measured from a
muscle of equal size but composed mainly
of fast fibers. In addition, Vmax will be
greater and TPT and HRT will be smaller
for the predominantly fast muscle relative
to the predominantly slow muscle.
Conclusions
Whole skeletal muscles and individual
muscle-fibers differ in their physiological,
biochemical and structural properties. In
order to characterize fully the muscle
fiber, structural, biochemical and physio-
logical properties must be measured.
This review may give the erroneous
impression that the structural and
functional properties of skeletal muscle
are static, whereas in fact each property
discussed above (architecture, fiber-type
distribution, motor-unit distribution,
length-tension properties and force-
velocity properties) is subject to
significant change, given the appropriate
external stimulus (Edgerton 1978, Pette
1980, Salmons and Henriksson 1981,
Jolesz and Sreter 1981, Pette and Vrbova
1985). Examples of these types of stimuli
and the resulting changes will be the
subject of the two succeeding papers in
this series.
Accepted for publication 17th March 1986.
Acknowledgements
The author would like to thank Professor Alan R.
Hargens and Dr. Jennette L. Boakes (University of
California, San Diego) and Dr. Jan Friden
(University of Umea, Sweden) for their helpful
comments on this manuscript. Thanks also to
Professor V. Reggie Edgerton and Dr. Roland R.
Roy (University of California, Los Angeles) for
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