J. Anat. (2008) 212, pp144–152 doi: 10.1111/j.1469-7580.2007.00848.

Comparative anatomy and muscle architecture of selected

Blackwell Publishing Ltd

hind limb muscles in the Quarter Horse and Arab

T. C. Crook,1 S. E. Cruickshank,1 C. M. McGowan,2 N. Stubbs,2 J. M. Wakeling,1 A. M. Wilson1
and R. C. Payne1
1
Structure and Motion Lab, Royal Veterinary College, London, UK
2
Animal Sciences Division, University of Queensland, Queensland, Australia

Abstract
The Quarter Horse (bred for acceleration) and the Arab (bred for endurance) are situated at either end of the
equine athletic spectrum. Studies into the form and function of the leg muscles in human sprint and endurance
runners have demonstrated that differences exist in their muscle architecture. It is not known whether similar
differences exist in the horse. Six Quarter Horse and six Arab fresh hind limb cadavers were dissected to gain
information on the muscle mass and architecture of the following muscles: gluteus medius; biceps femoris;
semitendinosus; vastus lateralis; gastrocnemius; tibialis cranialis and extensor digitorum longus. Specifically, muscle
mass, fascicle length and pennation angle were quantified and physiological cross-sectional area (PCSA) and
maximum isometric force were estimated. The hind limb muscles of the Quarter Horse were of a significantly
greater mass, but had similar fascicle lengths and pennation angles when compared with those of the Arab; this
resulted in the Quarter Horse hind limb muscles having greater PCSAs and hence greater isometric force potential.
This study suggests that Quarter Horses as a breed inherently possess large strong hind limb muscles, with the
potential to accelerate their body mass more rapidly than those of the Arab.
Key words Arab; architecture; biomechanics; equine; locomotion; muscle; Quarter Horse.

Research has suggested that the anatomy, and in

Introduction
The horse is an exceptional athlete, capable of sprinting
rapidly over short distances whilst maintaining the ability
to travel great distances at slower speeds. Selective breed-
ing for performance has resulted in distinct breeds of
horse which excel at different equine disciplines, such as
the Quarter Horse (bred for acceleration) and the Arab
(bred for endurance).
In Quarter Horse racing, race times of 21 s for a quarter
of a mile (402 m) have been recorded. The horse, acceler-
ating from a standing start, achieved a mean speed of
approximately 70 km h–1 (‘A Long Goodbye’; AQHA, 2005),
whereas Arab horses in endurance racing travel at a
much slower speed of 17 km h–1 covering 160 km in 9.5 h
(‘Xandu Haji Buba’; UAE Equestrian & Racing Federation,
2006). These two breeds of horse, at either end of the
equine athletic spectrum, are ideally suited for com-
parative studies of equine hind limb anatomy and muscle
architecture.
Correspondence
Tracy Crook, Structure and Motion Laboratory, The Royal Veterinary
College, Hawkshead Lane, Hatfield, Herts, AL9 7TA, UK. T: + 44 (0)1707
666333; E: tcrook@rvc.ac.uk
Accepted for publication 21 November 2007
particular the muscle architecture of the fore and hind
limbs of the horse, are optimized for biomechanically
distinct functions (Payne et al. 2004, 2005). The proximal
muscles of the horse’s hind limb provide the power
required for locomotion (Merkens et al. 1993; Dutto et al.
2004b), whilst the forelimbs act as stiff spring-like struts
(McGuigan & Wilson, 2003) and support a greater proportion
of the body mass (Witte et al. 2004).
Equine hind limb anatomy
The proximal hind limb muscles, many of which attach
directly to the skeleton without a tendon (Payne et al.
2005), account for the majority of the horse’s locomotor
mass (Gunn, 1987; Payne et al. 2005). Anatomical, bio-
mechanical and electromyographic studies indicate that
these muscles provide the mechanical power output for
activities which require net external work, such as accelera-
tion, running uphill and jumping (Robert et al. 2000;
Clayton et al. 2002; Dutto et al. 2004a).
The distal hind limb muscles, small in comparison, attach
to the skeleton via long thin spring-like tendons, an adapta-
tion in cursorial animals which reduces the need for
energetically expensive muscle length change (Dimery
et al. 1986). Further information regarding the locomotor

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Journal compilation © 2008 Anatomical Society of Great Britain and Ireland

Fig. 1 Locomotor anatomy of the equine hind limb. (A) Lateral superficial and (B) lateral deep views. *Gracilis and sartorius are transparent so that
deeper structures can be seen. Figure taken directly from Payne et al. (2005), by permission.
anatomy, origin, insertion and action of the muscles that
were studied can be found in Fig. 1 and Table 1.
The role of an individual muscle during locomotion is
influenced by several features: muscle architecture; fibre
type; moment-generating capacity (muscle force multiplied
by moment arm) (Biewener & Roberts, 2000); and activity
patterns. In this study we focus on breed differences in
muscle mass and architecture in the equine hind limb;
future studies will address differences in fibre type and
moment arms.
Muscle architecture (the arrangement of the muscle
fibres within the muscle, relative to the axis of muscle
force generation) is often described in terms of the follow-
ing five parameters: muscle belly and tendon length;
muscle fascicle length; muscle fascicle pennation angle;
and physiological cross-sectional area (PCSA) (Zajac, 1992).
Physiological cross-sectional area (PCSA) is defined as the
sum of the cross-sectional area of the muscle fibres within
the muscle belly. It can be estimated using the following
equation:
2 muscle mass (g)
PCSA (cm ) = −3
muscle density (g cm ) × fascicle length (cm)
where muscle density is taken as 1.06 g cm–3 (Mendez,
1960).
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Journal compilation © 2008 Anatomical Society of Great Britain and Ireland
Locomotor anatomy of the equine hind limb, T. C. Crook et al. 145
PCSA is related to the muscle’s maximum isometric force
potential (Fmax) (Powell et al. 1984), because Fmax is a product of
PCSA and the maximum isometric stress of vertebrate skeletal
muscle. However, owing to pennation of the fascicles, only
a proportion of the fascicle force acts in the direction of
the muscle belly and tendon. Thus Fmax is estimated by:
Fmax = PCSA · maximum isometric stress · cos θ
where the maximum isometric stress of vertebrate skeletal
muscle is assumed to be 0.3 MPa (Wells, 1965; Woledge
et al. 1985; Zajac, 1989; Medler, 2002), and θ is the pennation
angle of the muscle fascicles.
Fascicle length (which is indicative of the number of
sarcomeres arranged in series within the fascicle) relates
to the range of motion over which a muscle can exert its
force and its velocity of contraction (Gordon et al. 1966;
Wickiewicz et al. 1984).
Muscle architecture has been shown to vary between
muscles within the same individual (Brown et al. 2003;
Payne et al. 2005), and between the same muscle in different
individuals (Abe et al. 2000; Blazevich et al. 2006). Exercise
(Blazevich et al. 2003) and genetic make-up, in terms of gender
(Abe et al. 1998) are known to influence muscle architecture,
but the extent to which differences in training and genetic
profile relate to athletic performance remains unclear.
146 Locomotor anatomy of the equine hind limb, T. C. Crook et al.

Table 1 Origin, insertion and action of muscles of the equine hind limb

Muscle Origin Insertion Action

Gluteus medius Gluteal fascia, sacrum Greater trochanter Extension and

and sacro-iliac ligament of femur abduction of hip
Biceps femoris
Vertebral head Spinous and transverse Blend with femoral and crural Extension and abduction
processes of last three fascia and insert onto fascia of hip during stance, flexion
sacral vertebrae, caudal onto patella ligament, patella, of stifle and extension of
fascia, broad pelvic ligament, tibial crest and lastly onto hock during swing
ischiadic tuber calcaneal tuber via calcaneal tendon
Intermediate head Ischial tuber and ischium As above As above
Caudal head As above As above As above
Semitendinosus
Vertebral head Last spinous and transverse Medial tibial crest with aponeurosis Extension of hip during stance,
processes of sacrum, caudal of gracilis and sartorius and calcaneal flexion of stifle and extension
fascia, first 3–4 caudal vertebrae tuber via calcaneal tendon which of hock during swing
and broad pelvic ligament unites with that of biceps femoris
to become accessory tendon
Pelvic head Ischial tuber As above As above
Vastus lateralis Craniolateral femoral shaft Middle patella ligament to Extension of stifle
tibial tuberosity and tibial crest
Gastrocnemius
Lateralis Distal femoral shaft Via common calcaneal tendon Extension of hock and
onto tuber calcanei flexion of stifle
Gastrocnemius
Medialis As above As above As above
Tibialis cranialis Shaft, lateral condyle Medial and intermediate Flexion of hock
and crest of tibia, cuneiforms and metatarsal III
proximal fibula
Extensor Digitorum Extensor fossa Extensor process of Extension of digits
longus of femur distal phalanx and flexion of hock

Primary action of each joint is listed first, followed by secondary/auxillary actions (Nickel et al. 1986). The hamstring muscles flex the stifle
during swing but constrain extension during the first half of stance due to the antagonistic action of rectus femoris and the cranial position
of the vertical ground reaction force vector (Clayton et al. 2001).

When compared to elite distance runners, human elite
sprint runners have been observed to have longer muscle
fascicle lengths, arranged at lesser pennation angles in
their leg muscles (Abe et al. 2000), which has been linked
to running speed (Kumagai et al. 2000).
Research into the effect of differences in muscle
architecture and performance in the horse is sparse;
instead researchers have tended to focus on the effects of
different muscle fibre types (Rivero & Henckel, 1996;
Rivero et al. 2001) and metabolic influences such as the
maximum capacity for oxygen consumption by the body
during maximum exertion (VO2max) (Langsetmo et al. 1997;
Prince et al. 2002). This paper attempts to address this gap
in the literature by comparing the hind limb muscle
architecture in two breeds of horse – the Quarter Horse,
which has been selectively bred for sprinting, and the
Arab, which has been selectively bred for endurance – to
see whether differences in breed athleticism are reflected
in differences in muscle architecture. Whilst all animals
were being regularly exercised, none was being trained
for either sprint or endurance activities; as such it is
expected that any observed differences in muscle
architecture will be as a result of breed differences rather
than training effect.
It was hypothesized that, when comparing horses of
similar height and body mass, the Quarter Horse hind limb
muscles would, firstly, have a greater mass and, secondly,
have longer fascicle lengths with lesser pennation angles
when compared to the same muscles in the Arab.
Materials and methods
Horses
The study was conducted at the University of Queensland
and approved by the University’s Animal Ethics Committee.
Twelve adult horses (minimum age 3.5 years) were eutha-
nased for reasons unrelated to musculoskeletal disease
(Table 2): six Quarter Horses, consisting of four geldings
and two mares (13 ± 8 years); and six Arab horses, consisting
of five geldings and one mare (13 ± 7 years). All horses had
previously been in light ridden work.

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 Locomotor anatomy of the equine hind limb, T. C. Crook et al. 147

Table 2 Subjects

Age Mass Height Femur Tibia MT3

(years) Sex (kg) (m) Length (cm) Length (cm) Length (cm) Breed

Limb 1 20 Gelding * 1.51 46.0 38.5 44.5 Quarter Horse

Limb 2 12 Mare * 1.49 45.0 40.0 47.0 Quarter Horse
Limb 3 14 Gelding 504 1.51 46.0 41.0 45.5 Quarter Horse
Limb 4 25 Mare 461 1.52 44.0 37.0 42.5 Quarter Horse
Limb 5 3.5 Gelding 445 1.49 45.0 40.0 44.0 Quarter Horse
Limb 6 4.5 Gelding 419 1.48 44.0 38.5 43.0 Quarter Horse
Limb 7 4 Gelding * 1.50 44.0 37.5 45.0 Arab
Limb 8 8 Gelding * 1.46 43.0 36.5 45.0 Arab
Limb 9 20 Mare 388 1.41 45.0 37.0 42.5 Arab
Limb 10 20 Gelding 370 1.44 41.5 33.5 41.5 Arab
Limb 11 20 Gelding 327 1.48 44.0 37.0 43.0 Arab
Limb 12 20 Gelding 448 1.51 44.5 37.5 46.0 Arab
Mean QH 13 457 1.50 45.0 39.0 44.4
Mean Arab 15 383 1.47 44.0 36.5 43.8

MT3 = third metatarsal bone, *It was not possible to determine the mass of these animals because scales were not available prior to
euthanasia.

The height (m) of each animal was determined by the
use of a tape measure, measuring from the ground up to
the highest point of the withers whilst the horse was
standing square, sedated in stocks. All of the subjects were
unshod. The mass (kg) of each subject was determined
when feasible by standing the animal on an electronic
weigh scale (‘Tru Test Easy Weigh II’ Sunbeam, Sydney,
Australia). Limb length was determined by measuring and
combining femur, tibia and the third metatarsal bone
(MT3) length once the muscles had been removed. Bone
lengths were measured using standard anatomical land-
marks. Femur length was measured from the most proxi-
mal part of the greater trochanter to the most distal point
on the lateral femoral condyle; tibia length was measured
from the inter-condylar eminence to the most distal point
on the medial malleolus; and MT3 length was measured
from the most proximal lateral part of the head to the
most distal part of the lateral condyle. It was not possible
to measure the bone lengths of the individual phalanges
as these distal bones were used in a separate study and
hence were not available.
Muscle architecture
Immediately following euthanasia the hind limb was
skinned and the overlying fascia was removed to expose
the superficial muscles. The superficial gluteal and tensor
fascia lata muscles were dissected away from their fascial
and bony attachments to expose gluteus medius. The
following muscles, together with their tendons, were
identified and removed for further analysis: gluteus medius;
biceps femoris; semitendinosus; vastus lateralis; gastrocne-
mius; extensor digitorum longus; and tibialis cranialis.
The three heads of biceps femoris and the two heads of
gastrocnemius were separated from each other by
carefully dividing adjoining fascia.
Muscle belly length (mm) was determined by measuring
the distance from the origin of the most proximal muscle
fibres to the insertion of the most distal fibres with a
flexible tape measure. Muscle mass (g) to the nearest 0.1 g
was recorded using electronic scales (EKS, Hereford, UK)
and reported to the nearest gram. Muscles over 2000 g
were weighed in several pieces.
A minimum of three (central, medial and lateral) full
thickness cuts were made in the muscle belly, in a plane
following the direction of the muscle fascicles. Five
measurements of fascicle length were obtained from each
section, at varying depths, by laying a flexible measuring
tape along the length of each fascicle. Five resting fascicle
pennation angles (Gans & de Vree, 1987; Payne et al. 2005)
were determined by measuring the angles between the
muscle fascicle and internal tendon and/or aponeurosis
using a clear plastic protractor (see plate 1). Multiple
measurements within the same muscle and, where
appropriate, within muscle compartments, were averaged.
If resting pennation angles were less than 5°, they were
given a value of zero and were not considered in calculations
of Fmax.
Statistical analysis
The Mann–Whitney U-test, at a significance level of
P < 0.05, was used to compare the height, body mass, hind
limb bone length, individual muscle mass and muscle belly
lengths between the two subject groups. A two-way
hierarchical ANOVA at a significance level of P < 0.05 was
used to determine whether statistical differences in
fascicle length and pennation angles were present between

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148 Locomotor anatomy of the equine hind limb, T. C. Crook et al.
the two subject groups. Data were tabulated using MICROSOFT
OFFICE EXCEL 2003 (Microsoft Corporation, Redmond, WA,
USA). Statistical analysis was performed using SPSS,
version 14 (SPSS Inc., Chicago, IL, USA).
Results
Height and body mass
It was not possible to detect a statistical difference in the
height (m) (P = 0.09, n = 12) or body mass (kg) (P = 0.33,
n = 8) between the two groups (Table 2).
Hind limb bone length
No statistical difference was detected in femoral (P = 0.07,
n = 12) or third metatarsal bone lengths (P = 0.62, n = 12),
but tibial bone length was slightly greater in the Quarter
Horse group (P = 0.015, n = 12) (Table 2).
Muscle mass
The Quarter Horses exhibited larger muscle masses
(Table 3), and hence muscle volumes, in both the proximal
and distal muscles (P ≤ 0.01, n = 12). Gluteus medius,
which had the largest mass in both breeds, represented
2.0% of body mass in the Quarter Horse and 1.7% in the
Arab. Gastrocnemius had the largest mass of the distal leg
muscles, the lateral head having a greater mass than the
medial. Tibialis cranialis, with the smallest muscle mass,
represented 0.08% and 0.05% body mass in the Quarter
Horse and Arab respectively.
Muscle belly length
The muscle belly lengths of gluteus medius, semitendinosus
(vertebral and pelvic heads), biceps femoris (vertebral
head), and extensor digitorum longus muscles were
statistically longer (P < 0.05, n = 12) in the Quarter Horse
group (Table 3).
Fascicle length
No statistical difference (P = 0.20, n = 12) in fascicle length
was observed when comparing the same muscle in either
breed of horse (Table 3). The proximal muscles in both
groups, with the exception of the proximal part of
tibialis cranialis, had longer fascicle lengths than distal
leg muscles.
Regional differences were observed in fascicle length
within individual muscle bellies. Specifically, the medial
head of gastrocnemius had longer fascicle lengths than
the lateral head, and the proximal part of tibialis cranialis
had longer fascicle lengths when compared to the
distal part.
Journal compilation © 2008 Anatomical Society of Great Britain and Ireland
Pennation angle
There was no significant difference (P = 0.61, n = 12) in the
fascicle pennation angles when comparing the same
muscle in either breed (Table 3).
Biceps femoris could be divided into three distinct
compartments, named according to their anatomical
positioning: the vertebral, intermediate and caudal heads
(Payne et al. 2005).
In both breeds, the muscle fascicles were parallel in the
middle and caudal heads and had a similar pennation
angle in the vertebral heads. Similarly, semitendinosus
could be divided into two distinct sections: the pelvic head,
which consisted of parallel muscle fascicles (with a pennation
angle of less than 5°); and a pennate vertebral head. Tibialis
cranialis was composed of parallel fascicles proximally
and pennate fascicles in the distal section. There was no
statistical difference in the pennation angles of gastrocne-
mius medialis or lateralis in either breed or between the
two breeds.
Estimated parameters
PCSA and isometric force potential
All the Quarter Horse muscles had larger PCSAs and hence
greater isometric force potential (Fmax) (Table 4). Gluteus
medius had the largest PCSA and force potential in both
groups. PCSA and isometric force potential ( Fmax)
decreased in magnitude as follows: biceps femoris;
semitendinosus; gastrocnemius; vastus lateralis; extensor
digitorum longus; tibialis cranialis. The mean PCSA of
semitendinosus in the Quarter Horse was approximately
double that of the Arab.
Discussion
The mass and architecture of selected hind limb muscles
of the Quarter Horse and Arab were investigated via
cadaveric dissection. This study found that the hind limb
muscles of the Quarter Horses had a greater muscle mass
and PCSA when compared to the Arabs, which were of
a similar height and body mass overall. Based on these
findings, calculations of F max confirmed that the hind
limb muscles of the Quarter Horse had the capacity to
generate higher isometric forces than those of the
Arab. This was primarily due to their larger volumes,
rather than differences in muscle fascicle length or
pennation angle.
Proximal-to-distal muscle architecture
The proximal muscles in both breeds of horse had larger
volumes (Table 4) and fascicle lengths when compared to
the more distal pennate muscles (with the exception of the
© 2008 The Authors
Journal compilation © 2008 Anatomical Society of Great Britain and Ireland
© 2008 The Authors

Table 3 Muscle data. The mean ± standard deviation of muscle mass (g), belly length (mm), fascicle length (mm) and pennation angles (°) of individual muscles in the Quarter Horse (n = 6) and Arab (n = 6)
hind limb

Muscle mass (g) Muscle belly length (mm) Fascicle length (mm) Pennation Angle (°)

Muscle QH Arab QH Arab QH Arab QH Arab

Gluteus Medius *10010 ± 559 6789 ± 772 *577 ± 50 540 ± 32 155 ± 15 140 ± 23 44 ± 3 46 ± 5
Biceps femoris (vertebral head) *6557 ± 750 4715 ± 646 *684 ± 33 618 ± 42 191 ± 20 183 ± 23 52 ± 3 52 ± 9
Biceps femoris (intermediate head) *702 ± 81 571 ± 132 296 ± 26 267 ± 37 188 ± 17 152 ± 19 ≤5 ≤5
Biceps femoris (caudal head) *1178 ± 358 618 ± 284 343 ± 46 316 ± 46 189 ± 20 199 ± 47 ≤5 ≤5
Semitendinosus (vertebral & pelvic head) *4360 ± 1196 2426 ± 279 ж ж ж ж ≤5 ≤5
Semitendinosus vertebral head ж ж *709 ± 36 624 ± 49 143 ± 9 137 ± 19 39 ± 5 23 ± 20
Semitendinosus pelvic head ж ж *408 ± 22 358 ± 38 149 ± 28 153 ± 11 ≤5 ≤5
Vastus lateralis *2134 ± 311 1559 ± 209 303 ± 24 303 ± 24 117 ± 19 104 ± 12 51 ± 6 41 ± 7
Gastrocnemius (lateralis & medialis) *1673 ± 129 1179 ± 108 ж ж 42 ± 3 38 ± 6 ж ж
Gastrocnemius lateralis *962 ± 86 684 ± 83 267 ± 8 282 ± 83 36 ± 4 33 ± 6 48 ± 4 44 ± 3

Locomotor anatomy of the equine hind limb, T. C. Crook et al. 149

Gastrocnemius medialis *711 ± 47 495 ± 32 249 ± 17 272 ± 83 47 ± 4 44 ± 7 46 ± 3 46 ± 3
Extensor digitorum longus *431 ± 70 310 ± 28 *252 ± 19 231 ± 14 51 ± 5 54 ± 13 36 ± 6 35 ± 5
Tibialis cranialis (proximal & distal) *353 ± 30 262 ± 41 320 ± 22 318 ± 13 68 ± 7 70 ± 5 ж ж
Tibialis cranialis (proximal compartment) ж ж ж ж 117 ± 14 126 ± 9 ≤5 ≤5
Tibialis cranialis (distal compartment) ж ж ж ж 20 ± 5 15 ± 2 44 ± 9 49 ± 4

QH = Quarter Horse; *A significant difference (P < 0.05) in value between the two breeds; ≤ 5 = pennation angles of 5º or less; ж = value not reported as data are presented for the entire
muscle or for separate compartments/heads.
150 Locomotor anatomy of the equine hind limb, T. C. Crook et al.

Table 4 Estimates of volume (cm3), physiological cross-sectional area (cm2) and maximum isometric force (N), of named hind limb muscles in the
Quarter Horse (n = 6) and Arab (n = 6). Note: Semitendinosus pelvic head, and the intermediate and caudal heads of biceps femoris had relatively short
muscle fascicles (which were arranged both in series and in parallel to each other), relative to muscle belly length. The estimated values of Fmax given
are therefore an overestimation of between 200 and 300% of the actual maximum isometric force potential of the muscles.

Volume (cm3) PCSA (cm2) Force (N)

Muscle QH Arab QH Arab QH Arab

Gluteus medius 9443 6405 609 457 13152 9531

Biceps femoris (vertebral head) 6186 4449 324 243 5978 4493
Biceps femoris (intermediate head) 663 539 35 35 1060 1065
Biceps femoris (caudal head) 1111 583 59 29 1759 877
Semitendinosus (vertebral & pelvic head) 4114 2289 282 158 7950 4636
Vastus lateralis 2013 1471 172 141 3252 3201
Gastrocnemius lateralis 907 645 250 197 5009 4250
Gastrocnemius medialis 671 467 141 106 2944 2208
Extensor digitorum longus 406 292 79 54 1918 1333
Tibialis cranialis 333 247 49 35 1357 960

QH = Quarter Horse.

proximal part of tibialis cranialis muscle). These findings enable the muscles to generate force throughout a greater
are in agreement with previous studies of equine hind range of motion than those of the Arab, facilitating
limb muscles (Payne et al. 2005). increased stride length (i.e. speed = stride length × stride
frequency).

Muscle mass
In this study, the Quarter Horse hind limb muscles con-
tributed a higher percentage to total body mass when
compared to those of the Arab, but the relatively small
sample (n = 4) makes it difficult to draw any firm conclusions.
However, a similar study by Gunn (1987) found the hind
limb muscle mass to body mass ratio to be greater in
thoroughbred horses trained for sprinting when compared
to hurdling, suggesting this attribute may well be related
to the ability to accelerate rapidly.
Muscle belly length
The Quarter Horse hip extensor (gluteus medius, semi-
tendinosus and the vertebral head of biceps femoris) and
extensor digitorum longus muscles had significantly longer
muscle bellies when compared to the same muscles in the
Arab (Table 3). This may be owing to the fact that the
Quarter Horse group had slightly longer limb lengths
(afforded by a longer tibia), although the remaining
muscles in both groups were of a similar belly length. It
may alternatively/additionally be due to conformational
differences between the breeds: for example, differences
in muscle–tendon length (which were not determined),
or differences in muscle moment arms (which will be
the subject of a future study). It is suggested that the
longer muscle belly lengths of the Quarter Horse hind
limb retractors may be a reflection of the breed’s ability
to accelerate rapidly, as the longer muscle belly lengths
Fascicle length/pennation angle
As it was not possible to detect a significant difference
between the two breeds of horse in muscle fascicle length
or pennation angle in either the proximal or distal leg
muscles, we rejected the hypothesis that the Quarter
Horse hind limb muscles would be composed of longer fas-
cicle lengths with lesser pennation angles when compared
to the same muscles in the Arab. This hypothesis was
proposed based on the fact that longer muscle fascicle
lengths (associated with a greater velocity of contraction),
arranged at lesser pennation angles, have been observed
in the calf muscles of human elite sprint runners when
compared to elite distance runners (Abe et al. 2000) and
that fascicle length appears to be linked to running speed
(Kumagai et al. 2000). Our study, however, suggests that
the observed differences in muscle fascicle length and
pennation angle in the leg muscles of human elite sprint
and endurance runners may have occurred as an adaptive
response to a specific training programme, in the same
way that muscle mass increases in response to resistance
training (Kearns et al. 2000) and pennation angle and
PCSA increases in human weight lifters (Aagaard et al.
2001).
Pennation of the muscle fascicles influences the
amount of force that is transmitted through the attaching
tendon (Zajac, 1992). Greater pennation angles result in
a significant force reduction, whilst smaller pennation
angles have minimal effect [because in muscles with

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Journal compilation © 2008 Anatomical Society of Great Britain and Ireland

pennation angles of 20° and below, cos(θ) is close to 1.0].
Although increased pennation decreases force due to
the direction of the force application, it also allows
muscles to have a larger PCSA for a given volume and
therefore results in an increased capacity of the muscle to
exert force. In this study, the intermediate and caudal
heads of biceps femoris, the pelvic head of semitendinosus,
and the proximal part of tibialis cranialis had muscle
fascicle pennation angles of less than 5°. The remainder
of the muscles were pennate and many exhibited pen-
nation angles in excess of 40°. The pennation angles of
all the muscles, in both breeds, are greater than those
recorded in the Thoroughbred by (Payne et al. 2005).
These differences may be breed-dependent, or because
of differences in sampling technique: for example, a
different location/compartment within the muscle may
have been sampled in the two studies. This is especially
relevant considering that pennation angle can vary
dramatically throughout the same muscle, a feature
particularly evident in the tibialis cranialis muscle. This
variation in pennation angle coupled with differences in
fascicle length suggests that functional specialization
exists within the same muscle, a finding that has been
mirrored by other researchers investigating human
muscle architecture (Aagaard et al. 2001; Blazevich et al.
2006). More detailed studies of individual muscles
would provide important information on muscle mor-
phological compartmentalization and hence functional
compartmentalization.
It is possible that differences in study design make it
inappropriate to compare the findings of our research to
published human studies which have determined differences
in muscle architecture in vivo using B-mode ultrasound
(Kumagai et al. 2000; Kearns et al. 2000; Blazevich et al.
2006). Undoubtedly, ultrasound allows the dynamics of
muscle architecture to be examined in vivo, enabling the
recordings of fascicle lengths and pennation angles at
specific muscle lengths (joint angles) and levels of muscle
tension (Narici et al. 1996; Maganaris & Paul, 2000);
however, it is constrained to providing information on
superficial musculature only and hence would not have
been a viable method of obtaining architectural data in
this study. We determined fascicle length and pennation
angle after the muscles had been removed from the limb.
It is possible that subsequent changes in fascicle length
occurred, especially following rigor mortis and potential
dehydration of any of the muscles, which can result in
tissue shrinkage (Cutts et al. 1991). However, as all muscles
in this study were examined and compared within 48 h of
euthanasia, under the same conditions, breed differences
in resting fascicle length and pennation angle should
not have been affected. It is appreciated that under ideal
circumstances muscle fascicle length should have been
normalized to sarcomere ‘resting’ length, but this was not
possible in this study.
© 2008 The Authors
Journal compilation © 2008 Anatomical Society of Great Britain and Ireland
Locomotor anatomy of the equine hind limb, T. C. Crook et al. 151
Calculated parameters
Estimates of maximum isometric force potential
The proximal limb muscles in both breeds of horse tended
to have the greater isometric force potential ( Fmax) than
the distal leg muscles; these findings are in agreement
with those of Payne et al. (2005).
All the Quarter Horse muscles had the ability to generate
greater isometric forces when compared to the same
muscles in the Arab. This was owing to differences in muscle
mass rather than differences in muscle fascicle length or
pennation angle (Table 3). The Quarter Horse semitendinosus
muscle had almost double the physiological cross-sectional
area when compared to that of the Arab, which resulted
in the Quarter Horse group having a much greater isometric
force potential. Studies comparing cross-sectional area of
semitendinosus in Thoroughbred horses (selectively bred
for racing) and other breeds of horse have similarly found
it to be greater in the Thoroughbred (Gunn, 1979), which
suggests it is an important hind limb retractor, capable of
generating large forces required for acceleration.
Semitendinosus pelvic head, and the intermediate and
caudal heads of biceps femoris had relatively short muscle
fascicles (which were arranged both in series and in parallel
to each other), relative to muscle belly length. The estimated
values of Fmax given in Table 4 are therefore an overestima-
tion of 200 –300% of the actual maximum isometric force
potential of the muscles. There is not a simple way of
compensating for this effect, although an alternative
method of calculating Fmax when muscle fascicles are
arranged in series, may be to use the following equation:
muscle mass (g)
Fmax =
·
maximum isometric stress
−3
muscle density (g cm ) ×
muscle belly length (cm)
However, the methodology used in this paper permits
comparison with previously published data and does not
affect the comparisons made between muscles in the two
breeds of horse studied.
Conclusion
The hind limb muscles of the Quarter Horse were of a
significantly greater mass, but had similar fascicle lengths
and pennation angles when compared to those of the
Arab. This resulted in the Quarter Horse hind limb muscles
having greater physiological cross-sectional areas and
greater isometric force potential. As both groups of
animals were undertaking similar exercise regimes,
this study suggests that the Quarter Horse as a breed
inherently possesses large strong hind limb muscles with
the potential to accelerate its body mass more rapidly
than those of the Arab. It is hoped that this study will
152 Locomotor anatomy of the equine hind limb, T. C. Crook et al.
complement future studies of muscle architecture in elite
Quarter Horse sprinters and elite Arab endurance horses
and will help to differentiate further between the effects
of training and genetic predisposition.
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