Human Locomotion 1 3

TABLE of CONTENTS
Preface v
Acknowledgments vii
Chapter One The Evolution of Bipedality 1

Sahelanthropus tchadensis and Orrorin tugenensis 5
Ardipithecus ramidus 6
Australopithecus anamensis and afarensis 8
Australopithecus garhi and africanus 12
Paranthropus robustus 12
Homo rudolfensis 13
Homo habilis 14
Homo ergaster, georgicus, and erectus 15
Homo floresiensis 16
Homo antecessor and heidelbergensis 17
Homo neanderthalensis 18
Homo sapiens 19
Chapter Two Structural and Functional Anatomy 23
PLANES OF MOTION 24
FUNCTIONAL ANATOMY 27
Interphalangeal Joints 28
Metatarsophalangeal Joints 28
First Ray 30
Second, Third and Fourth Rays 30
ix
x CONTENTS
Fifth Ray 31
Midtarsal Joint 31
Subtalar Joint 35
Ankle Joint 38
Tibiofibular Joints 39
Knee Joint 42
Hip Joint 49
The Pelvis 55
The Spine 60
MUSCULAR EFFICIENCY AND INTERACTION OF FORCES 67
Chapter Three Ideal Motions During the Gait Cycle 87
STANCE PHASE MOTIONS 94

Contact Period 94
Midstance Period 105
Propulsive Period 110
SWING PHASE MOTIONS 117
SUMMARY OF MUSCLE FUNCTION DURING THE GAIT CYCLE 120
GRAPHIC SUMMARY OF THE GAIT CYCLE 124

SUMMARY OF BIOMECHANICAL FACTORS ASSOCIATED
WITH IMPROVED RUNNING ECONOMY 133
Chapter Four Abnormal Motion During the Gait Cycle 139
DEVELOPMENTAL TRENDS IN LOWER EXTREMITY ALIGNMENT 140
Transverse Plane Alignment 140
Frontal Plane Alignment 147
DEVELOPMENT OF THE MEDIAL LONGITUDINAL ARCH 151
ALIGNMENT OF THE METATARSAL HEADS 172
VARIATION IN METATARSAL LENGTH 180
LIMB LENGTH DISCREPANCY 184

CONTENTS xi
EXCESSIVE MOBILITY 189
Tibiofemoral Joint 189
Ankle Joint 190
Subtalar Joint 191
Midtarsal Joint 191

First Ray 193
MINIMUM RANGES of MOTION NECESSARY FOR A
NON-COMPENSATED GAIT 195
Hip Joint 195
Tibiofemoral Joint 197
Ankle Joint 197
Subtalar Joint 199
First Metatarsophalangeal Joint 200
TREATMENT of LIMITED MOTION 201
Restricted Motion Resulting From Muscular Contraction 202
Muscle length assessment techniques 205
Home stretches 212
Restricted Motion Resulting From Osseous Block 219
Restricted Motion Resulting From Joint Stiffness 221
Joint manipulations/mobilizations 223
NEUROMOTOR COORDINATION and STRENGTH 237
Muscle Testing 244
Rehabilitative Exercises 251
Chapter Five Biomechanical Examination 271
SUPINE EXAMINATION 271
PRONE EXAMINATION 277

SEATED EXAMINATION 280
STANDING EXAMINATION 280
DYNAMIC EVALUATION 293

xii CONTENTS
Chapter Six Foot Orthotics 301
MECHANISM of ACTION 301
ORTHOTIC CASTING TECHNIQUES 304
LABORATORY PREPARATION and ORTHOTIC FABRICATION 309
Modification of the Positive Model 309
ORTHOTIC POSTING 311
ORTHOTIC SHELLS 316
ORTHOTIC ADDITIONS 318

Top Covers 318
Heel Lifts 320
Bar Posts 320
Balance for Lesion 320
Morton’s Extension 320
Metatarsal Pads 320
Toe Crests 321
Interdigital Plugs 322
Cuboid Pad 322
Kinetic Wedge 322
IN-OFFICE FABRICATION TECHNIQUES 323
ORTHOTIC DISPENSING 324
ORTHOTIC PROBLEM-SOLVING 324
Chapter Seven Shoe Gear 333
HISTORY OF FOOTWEAR 333
MODERN SHOES 335
SNEAKERS 335
MINIMALIST SNEAKERS 341
Chapter Eight Treatment Protocols 345
ACHILLES TENDINITIS 346

CONTENTS xiii
SESAMOIDITIS 349
METATARSALGIA and METATARSAL STRESS FRACTURES 349
PLANTAR PLATE RUPTURE 350
INTERDIGITAL NEURITIS 350
HALLUX ABDUCTOVALGUS 351
HALLUX LIMITUS and RIGIDUS 353
HEEL PAIN 354
TIBIALIS POSTERIOR TENDINITIS 357
LATERAL ANKLE SPRAIN 359
ANTEROMEDIAL ANKLE IMPINGEMENT SYNDROME 363
COMPARTMENT SYNDROMES of the LEG 364
MEDIAL TIBIAL STRESS SYNDROME 366
STRESS FRACTURES 367
PATELLOFEMORAL PAIN SYNDROME 370
PATELLAR TENDINOPATHY 374
ILIOTIBIAL BAND COMPRESSION SYNDROME 376
MEDIAL KNEE PAIN 378
POPLITEUS TENDINITIS 379
HAMSTRING STRAINS 380
PIRIFORMIS SYNDROME 383
GREATER TROCHANTERIC PAIN SYNDROME 386
ANTERIOR HIP AND GROIN PAIN 386
Snapping Hip Syndrome 387
Osteitis Pubis 388
Hip Osteoarthrosis 389
Adductor Strains 390
Anterior Labral Injury and Femoroacetabular Impingement Syndrome 390
LOW BACK DISORDERS 393
Index 407
Chapter One
The Evolution of Bipedality
The process of walking around on two legs is an converted the once dense forests of eastern Africa into
inherently unstable form of transportation. Watch any the open grasslands of the savanna. Because food sources
toddler attempt a few steps and it is easy to appreciate the became more spread out, our early quadruped ancestors
complexity of the task. It is also an unusual way to get were forced to stand up and walk. This new form of
around: of the more than 4,000 species of mammals on transportation theoretically allowed the early hominids to
earth today, only one is upright when walking (1). Even see over the tall savanna grasses and cover larger distances
Plato commented on the curious nature of our preferred in search of food.
form of transportation by referring to humans as the only The problem with the savanna hypothesis is that
“featherless bipeds.” recent discoveries show that the timing is all wrong. In 2001,
In 1871, Charles Darwin claimed that bipedality a team of French and Kenyan paleontologists announced
was the defining feature separating humans apart from our the discovery of multiple specimens of a 6-million-year-old
ape ancestors (2). Darwin theorized that the conversion to hominid they named Orrorin tugenensis (4). Discovered in
bipedal gait freed the hands to allow for tool use, which the Tugin hills of Kenya, the femur of this early hominid
in turn created an environment that favored rapid brain was remarkably humanlike, as it even possessed a groove
expansion. This line of reasoning is consistent with that on the back of the femoral neck for the obturator externus
of modern anthropologists such as Mary Leakey (3), muscle. This groove is only present in bipeds and confirmed
who states that bipedality “freed the hands for myriad that Orrorin most definitely walked upright (Fig. 1.1). In
possibilities: carrying, toolmaking, intricate manipulation... 2002, a team of paleontologists led by Michael Brunet (5)
this new freedom of the forelimbs posed a challenge. The unveiled a newly discovered skull from a 7-million-year-
brain expanded to meet it. And mankind was formed.” old hominid they called Sahelanthropus tchadensis (named
According to the classic theory of bipedal evolution, after the region in Africa where the fossil remnants were
approximately 2.5 million years ago a seismic shifting of discovered). Although no other remains have been found,
tectonic plates caused a rapid global cooling that quickly the skull of this hominid possessed a centrally located
Figure 1.1. In contrast to quadrupedal locomotion, bipedal gait forces the ischium to move down and forward
(arrow A), which significantly increases tensile strain placed on the obturator externus tendon where it passes
along the posterior aspect of the femoral neck (B).
1
Human Locomotion: The Conservative Management of Gait-Related Disorders
foramen magnum, strongly suggesting Sahelanthropus provided no benefit to the earliest hominids (approximately
was a dedicated biped; i.e., because bipeds walk with their 7 million years ago), beginning 5 million years ago, it most
heads balanced over their upright cervical spines, they likely provided a significant advantage over the quadruped
possess centrally located foramen magnums. This contrasts apes.
with almost all quadrupeds, who walk with their heads 6) Improved efficiency: This theory proposes the
down and typically possess posteriorly displaced foramen transition from quadruped to biped gait was associated
magnums. The discovery of Sahelanthropus tchadensis in with a decreased cost of locomotion that saved precious
Africa pushes back the origins of bipedality from 4 million calories and improved foraging ranges. Even a slight
years ago to a minimum of 7 million years ago. This means decrease in the metabolic cost of locomotion would have
the early hominids were bipeds approximately 4 million provided a considerable benefit to the early hominids since
years before the formation of the savanna grasslands, 5 the saved calories could have been used for improved
million years before our freed hands were manipulating growth and/or reproduction. Of all the proposed theories,
tools, and at least 6 million years before significant brain improved efficiency has been the most intensively studied
expansion. These recent findings raise significant questions and surprisingly, the early research was inconclusive.
concerning the reasons why we began to walk upright. In 1973, Taylor and Rowntree (9) evaluated the
Obviously the transition to bipedal gait has served metabolic efficiency of trained juvenile chimpanzees
us well, but why did those early hominids stand up and as they ambulated with biped and quadruped gaits.
take the first few steps? The most current theories are Because the metabolic costs of locomotion between the
summarized as follows: two gait patterns were the same, the authors stated that
1) Freeing of the hands: Bipedality freed the improved biomechanical efficiency could not be a factor
hands so the early hominids could more efficiently carry in the development of bipedality. In 1996, Steudel (10)
food and/or offspring. This reduced the metabolic cost of reviewed the literature concerning the metabolic efficiency
foraging, as larger amounts of food could be carried longer controversy and concluded that because of the shape of a
distances. While this became extremely important when chimpanzee’s pelvis, their center of masses are displaced
day ranges for foraging increased after the formation of the anteriorly, which forces them to walk in a stooped position
savanna grasslands, it would have provided little advantage with exaggerated hip and knee flexion (Fig. 1.2). This is
to the 7-million-year-old Sahelanthropus, who lived in extremely inefficient, since the chimp is forced to apply
dense forests where day ranges for foraging were so small. propulsive period forces in front of the hip joint instead
2) Improved vision: Standing upright improves of behind the extended hip, as in humans. Because of
visual range allowing for improved foraging and/or dramatic differences in skeletal alignment between
observation of predators. Again, this would have been chimps and modern humans, Steudel (10) argues the
important after the appearance of the savanna grasslands, early hominids would have faced similar challenges and
but not 7 million years ago as Toumai and Orrorin inhabited that energetic efficiency should not be considered when
heavily forested jungles. discussing the origins of bipedality, since “it is not possible
3) Aquatic theory: Originally described by Alister
Hardy and later popularized by Elaine Morgan (6) this
theory maintains that the early hominids became bipeds
in the relatively weightless environment of water. They
support this theory with the observations that unlike most
mammals, humans are hairless and possess a thick layer
of subcutaneous fat that is necessary for buoyancy and
insulation in an aquatic environment. They also note that
during childbirth, only human offspring present headfirst
in the vaginal canal making land-births hazardous. Despite
still being presented as valid in the popular press, virtually
all aspects of the aquatic theory have been dismissed (7).
4) Threat display theory: In 1974, R. Guthrie
suggested that the earliest male hominids used their newly
exposed penises as a “threat display organ” to intimidate
rival males (8). Although this may have occurred with some
of the earliest hominids, the concept seems far-fetched and
Figure 1.2. Unlike Homo sapiens, the chimpanzee’s
few paleontologists consider this theory valid.
longer pelvis displaces the center of mass forward
5) Temperature regulation: Walking upright (circle A), causing them to walk with both hips and
lessens solar skin exposure thereby allowing for increased knees flexed (white arrows).
foraging distances in the mid-day tropical sun. While this
2
Chapter One The Evolution of Bipedality
to compare such structurally different creatures as the early that 4.4 million years ago, upright walking might have
hominids and modern humans.” The findings by Taylor occasionally occurred in our hominid ancestors, but that it
and Rowntree (9) and Steudel (10) have been supported would not be the primary form of transportation. The fact
by more recent research by Nakatsukasa et al. (11), who that Ardi walked upright as a habitual biped forced scientists
calculated metabolic efficiency in a wide range of primates to change the way they viewed our last common ancestor in
and demonstrated that bipedalism was metabolically more that prior to Ardi, experts assumed that the farther back you
expensive than quadrupedal locomotion in every situation. went in the evolutionary tree, the more chimpanzee-like
Arguing in favor of improved efficiency with our ancestors became. The discovery of Ardi changed that,
bipedality, Sockol et al. (12) note the research by Taylor because her skeleton possessed an unanticipated amount of
and Rowntree (9) was flawed as they only studied juvenile modern human traits. For example, the bones of her hands
chimps (who are often less efficient than their adult revealed surprisingly short thumbs, short metacarpals and
counterparts) and the authors failed to include a complete exceptionally mobile wrists confirming that she was not a
biomechanical analysis of the metabolic expense of biped terrestrial knuckle-walker. The feet of Ardipithecus were
and quadruped locomotion. In an extremely detailed study, also unexpected as they were chimp-like in appearance but
Sockol et al. (12) evaluated the cost of locomotion (COL) her lesser metatarsals were humanlike in function as they
of 5 adult chimpanzees (ages 6 through 33 years) after they were short and extremely strong, allowing her to walk with
spent 4 months training the chimps to walk with both biped her hips slightly extended during the push-off phase of gait.
and quadruped gaits on a treadmill. The chimps wore masks The most surprising skeletal discovery as related
so their oxygen consumption could be measured and high- to bipedality was that Ardi’s lumbar spine was so mobile
speed video analysis was performed with kinematic data that she could arch her back into a lordosis, effectively
calculated by evaluating motion between markers placed balancing the weight of her upper body over her pelvis.
on each chimp’s shoulder, elbow, wrist, knee and ankle. The ability to create a lordotic lumbar spine was essential
By extrapolating information from this data, joint moments
were evaluated and the volume of muscle activated for
each step was then calculated and compared to total contact
times during biped and quadruped locomotion.
By using research on efficiency from other studies,
Sockol et al. (12) determined that when averaged out
between the 5 chimps, bipedal walking was slightly more
costly than quadrupedal walking. However, a more detailed
analysis comparing biped and quadruped locomotion
among individual chimpanzees revealed that for 3 of
the chimps, bipedalism was more expensive, in 2 of the
chimps, the cost of locomotion was similar, and in one
specific chimp (a 33-year-old female), the cost of bipedal
locomotion was significantly lower than quadrupedal
locomotion. In fact, this particularly efficient chimp
consumed only .16 ml of oxygen per kilogram per meter
when walking biped compared to .29 ml of oxygen when
walking as a quadruped. The authors conclude that even
if our early ancestors walked with a “bent knee, bent hip
form of bipedalism” the results of this study suggest that
even the earliest transitional forms of hominids could have
reaped some energy savings with the conversion to bipedal
gait. This in turn would have provided critical selection
pressure favoring bipedality.
The controversy regarding metabolic efficiency
continued until October of 2009, when the journal Science
published the results of a 15-year analysis of the 4.4-million-
year-old hominid Ardipithecus ramidus (nicknamed
“Ardi”). This important hominid predated Lucy by more
than 1 million years and biomechanical analysis of her
surprisingly complete skeleton confirmed that Ardi could Figure 1.3. Ardipithecus ramidus. Modified and
easily stand upright and was in fact a dedicated biped (Fig. redrawn from an illustration by J. H. Matternes in Science.
1.3). This finding was unexpected since scientists assumed 2009;326:101.
3
for improved metabolic efficiency since balancing the

upper body over the pelvis markedly reduces extensor
moments in the back, hip and thigh musculature, which in
turn significantly reduces the cost of locomotion (Fig. 1.4).
Because chimpanzees have such rigid lumbar spines (that
provide invaluable protection against spinal shear forces
associated with twisting between branches) they are unable
to balance effectively in an upright position and can only
walk as bipeds by excessively flexing their knees, hips and
spine.
The shape of Ardi’s upper pelvis was also
unexpected as her ilia angled forward allowing the hip
abductors to support weight more effectively during
single-limb support. This is in contrast to the chimpanzee
ilia that rest flat against the back and project up towards the
ribs (Fig. 1.5). Because the chimpanzee ilia are positioned
posteriorly, gluteus medius behaves more as a hip extensor
and is unable to stabilize the pelvis in the frontal plane
during single-limb stance (i.e., a chimpanzee cannot stand
upright on one leg).
All of these findings confirm that our last common
ancestor was not similar in shape to a modern chimp but
more closely resembled a blend between chimpanzees and Figure 1.5. The chimpanzee ilia (A) are positioned in
humans. Ardi confirmed what in hindsight seems obvious: the frontal plane, which makes it impossible for the
hip abductors to stabilize the pelvis effectively during
single-limb support.
while modern humans have spent the last 7 million

years evolving into our present form, chimpanzees and
apes have also evolved considerably, perhaps even more
so, developing features making them more efficient at
suspending themselves from branches and moving about
in high trees.
Because our last common ancestor possessed a
mobile lumbar spine and an upper pelvis that angled
slightly forward, the transition to bipedality would have
been much easier than previously believed using the
chimpanzee-centric model. This transition would have
provided the transitional hominids a slight increase in
metabolic efficiency for terrestrial locomotion while also
allowing them to use their hands, possibly supplying food
to pregnant females thereby ensuring that natural selection
would heavily favor bipedality (13). After meticulously
analyzing every detail of Ardi’s skeleton and comparing
it to past and present hominids, Lovejoy et al. (14) claim
the last common ancestor was not skilled at prolonged
suspension or climbing high branches and most likely spent
most of its time in the lower canopy, and may have even
been partially terrestrial. They suggest that the hindlimb
of the last common ancestor “remained dominant for body
Figure 1.4. By extending the lumbar spine into a lordotic mass support during bridging and arboreal climbing”
position (arrow), Ardipithecus could balance the upper allowing for an easier transition to walking upright on two
body over the pelvis thereby minimizing stress on the legs. Lovejoy et al. (14) make the interesting statement that
hip and back musculature.
if the early hominids could have adapted to climbing as
4
Figure 1.6. Time line of our hominid ancestors. Solid lines represent well-supported relationships while the dashed lines
represent suspected but not confirmed relationships. Based on information from Wong K. The human pedigree. Scientific
American. January 2009:60-63.
well as apes and chimps, “neither bipedality nor its social The only skeletal remains are a single crushed
correlates would likely have evolved.” skull, a few jaw fragments and a few teeth. Dating back
By piecing together the ever-increasing supply of 7 million years, Toumai’s skull possessed a cranial base
skeletal remains of our hominid ancestors, researchers similar to both Ardipithecus and Lucy, confirming that
have been able to assemble a family tree that chronicles Sahelanthropus was definitely a hominid, not an extinct
the 7 million year transition towards efficient bipedalism ape (16). The reconstructed skull had a centrally located
(Fig. 1.6). The following section reviews the anatomical foramen magnum, which is a strong indicator of bipedality
changes in each of our hominid ancestors and relates this as it both balances the upright head on top of the cervical
information to how these changes improved efficiency spine and allows for a perpendicular angle between the
during upright walking and running. orbits of the eye and the base of the skull (Fig. 1.7). This
angle is essential for vision with bipedal walking and is
Sahelanthropus tchadensis not seen in quadrupeds. Unfortunately, without additional
The genus name Sahelanthropus translates into bony fragments it is impossible to verify the extent to
“Man from Sahel” and refers to the African region near which Sahelanthropus walked upright.
the Sahara where this hominid was first found. The species
name refers to Chad where the majority of fossil specimens Orrorin tugenensis
have been recovered. The nickname for this early ancestor Orrorin translates into “Original Man” in the
is Toumai for “Hope of Life,” which is the name local Tugen language and tugenensis refers to the hills in Kenya
villagers frequently use for human children born just where this hominid was first discovered. Bony fragments
before the dry season (15). date back from 5.7 to 6.2 million years ago and include
5
Figure 1.7. In modern humans (A) and Sahelanthropus tchadensis (B), the lines bisecting the foramen magnum and
the anterior rim of the eye sockets form a nearly perpendicular angle. In chimpanzees (C), the intersecting lines form a
more acute angle making it difficult for them to maintain a horizontal gaze without prolonged hyperextension of the cervical
spine. Because they spend so much time looking down, the foramen magnum is positioned posteriorly in the chimpanzee
(D).
jaw fragments, a few molars, a finger phalanx, and most patches of hackberry, fig and palm trees. Surviving on a
importantly, the upper two thirds of a left femur. According diet made mostly of ripe fruit, there was little evidence
to Sawyer et al. (15), the femur was “remarkably that Ardipithecus consumed open-environment resources,
humanlike” in that it had a long femoral neck and a groove confirming that inhabitation of the grasslands was not a
for the obturator externus muscle. This bony groove driving force for the development of bipedality (18).
would only have formed if Orrorin walked upright on a Because their canine teeth were so small and
regular basis. Arguing against significant bipedality, CT because males and females were similar in size (averaging
evaluations of Orrorin’s femoral neck revealed a chimp- 4 feet tall, weighing 110 pounds), social aggression must
like distribution of cortical bone; i.e., because of the higher have been minimal and females may have chosen their
bending forces on the chimpanzee femoral neck, a chimp mates not based on physical prowess, but by their foraging
possesses a greater percentage of cortical bone on both the efficiency (13).
superior and inferior surfaces. Because of this, Lovejoy As previously mentioned, the spine and upper
(17) states that Orrorin was “frequently, but not habitually pelvis of Ardipithecus were remarkably humanlike in that
upright, and most likely spent a significant amount of time her lumbar spine could form a lordosis and the ilia flared
in the trees.” forward, becoming vertically shorter and horizontally
wider. The ability of the lumbar spine to extend enabled
Ardipithecus ramidus Ardi to balance the upper body over the pelvis while the
Discovered in the Awash river valley in Ethiopia, forwardly angled ilia allowed gluteus medius to more
Ardi translates into “ground floor” while ramid means effectively stabilize the pelvis when standing on one leg.
“root.” These names pay tribute to the Afar people who have The changes in the lumbar spine and ilia represent the most
“strong roots” in the Awash River area of Ethiopia (15). fundamental adaptations necessary for efficient bipedality.
Until recently, all that was known about Ardi was that the Because Ardipithecus was habitually bipedal, the
glenohumeral joints were elliptical, the wrists were wide, anterior aspect of the pelvis was forced to remodel as
and the proximal phalanx of the hallux tilted backwardly the constant tensile strain produced by the rectus femoris
suggesting that Ardipithecus was occasionally bipedal but muscle pulled on the anterior ilium, causing the formation
most likely behaved more like a ground-moving ape than of the anterior inferior iliac spine (AIIS) (Fig. 1.8). Analysis
a tree climber (15). of her skeleton revealed the AIIS was exceptionally strong
This all changed in 2009 when the journal Science and formed from a separate ossification center (19).
published 11 papers detailing findings from a 15-year While Ardi’s upper pelvis was designed for
evaluation of dozens of bony fragments, including the bipedality, her lower pelvis was not as it retained an
nearly complete skeleton of a female recovered from an ape-like shape in that the ischial tuberosities projected
erosional degradation. By carefully analyzing thousands downwardly, not posteriorly, as they do in modern
of samples of vertebrate, invertebrate, and plant fossils humans and even Lucy (Fig. 1.9). This downward slope
found at the site, researchers were able to determine that would have allowed Ardi to climb trees more effectively
Ardipithecus lived from 4 to 6 million years ago and spent but it would have significantly limited the ability of the
most of its time in woodland areas surrounded by small hamstring musculature to decelerate the forward swing
6
of the leg during upright walking. This in turn would

have markedly reduced Ardi’s ability to walk quickly as
it alters the length/tension relationship in the hamstrings:
In Ardipithecus, the hamstrings generate peak torque when
the hip is flexed towards the chest (as when climbing a tree)
but the downward slope of the ischial tuberosities places
the hamstring musculature in a shortened position when
standing upright, which significantly reduces force output.
The upper extremity of Ardipithecus provided
crucial evidence that we were not descendents of knuckle-
walking apes, as Ardi’s wrists were extremely mobile and
the metacarpals remained primitively short and lacked the
cortical thickenings and adaptations typical of knuckle-
walking (20). The midcarpal joint was capable of such
extreme ranges of motion that it is assumed that Ardi’s
wrists were used for “advanced arboreal palmigrady,” in
which she could maintain her grip on a tree branch by
extending her wrist as her body progressed beyond the
branch.
The fact that we did not descend from knuckle-
walking apes was also confirmed by a biomechanical
analysis of the wrists of great apes by Kivell and Schmitt
(21). These researchers evaluated the carpal bones of
modern apes and determined that contrary to all published
literature, the wrists of modern apes do not possess bony
locking mechanisms (in fact, their wrists were extremely
mobile) and stability is achieved through a vertical
stacking of their carpal bones in a manner similar to the
Figure 1.8. Prolonged upright postures increase tensile legs of an elephant (Fig. 1.10). This vertical stacking is
forces placed on the origin of the rectus femoris muscle accomplished in part by a centralized head of the capitate
(one of the 4 quadriceps muscles) eventually resulting in bone that possesses a slight narrowing in the midsection.
the formation of a bony exostosis (x) known as the anterior
The capitate of Ardipithecus more closely resembled that
inferior iliac spine (AIIS).
Figure 1.9. The ischial tuberosity in humans and Lucy (A and B) project posteriorly, allowing the hamstrings to
remain under tension during upright postures. Because the ischial tuberosity in Ardipithecus (C) and chimpanzees (D)
angle downwardly, the hamstrings play a more important role in stabilizing the pelvis during quadrupedal locomotion and
while climbing (arrow in D). Notice the well-developed AIIS in humans, Lucy, and Ardipithecus (small arrows). Partially
adapted from Lovejoy O, Suwa G, Spurlock L. The pelvis and femur of Ardipithecus ramidus: the emergence of upright
walking. Science. 2009;326:71.
7
Figure 1.10. As chimpanzees knuckle-walk, the radius Figure 1.11. While the proximal phalanx of the
displaces forwardly (A) causing the scaphoid and lunate chimpanzee (A) project slightly downward relative to
to compress into the central portion of the capitate (B). the bisection of the first metatarsal shaft, the proximal
In apes and elephants, the bones of the wrists stack neatly phalanx of Australopithecus ardipithecus (B) has an
on top of each other, thereby minimizing impingement of the upward cant that allows for an increased range of first
central capitate. Partially adapted from: Kivell T, Schmitt D. metatarsophalangeal joint dorsiflexion during bipedal
Independent evolution of knuckle-walking in African apes gait.
shows that humans did not evolve from a knuckle-walking
ancestor. PNAS August 25, 2009;106:14241-14246. chimps and has a design that seems to be choosing between
climbing and walking.
of a chimpanzee with palmar displacement of its head (20).
These findings confirm that our last common ancestor was Australopithecus anamensis
not a knuckle-walker, because this trait developed much Australopithecus refers to “Southern Ape” while
later in the large-bodied apes as a generalized adaptation anamensis translates from the Turkana language into
allowing them to remain highly arboreal while also “lake” and was chosen because all of the specimens have
effectively moving about on the ground (21). been found near lakes. Bony fragments from a tibia dating
The foot of Ardipithecus was perhaps the strangest back 4 million years clearly shows that Australopithecus
finding as it retained an extremely primitive appearance anamensis routinely moved about on two legs while
in which the first metatarsal and great toe were displaced changes in the distal humerus revealed that anamensis
medially (giving the appearance of a hand-like foot), while had lost the ability to knuckle-walk efficiently (Fig. 1.12).
the lesser metatarsals were more modern as they were The distal aspect of the tibia was also consistent with a
relatively short, extremely strong, and possessed an upward dedicated biped as the long axis of the tibia is perpendicular
cant at the proximal phalanges. The upwardly canted lesser to the articular surface of the ankle joint. Oddly, the lower
metatarsals would only be present if the leg was allowed extremity features of Australopithecus anamensis reveal
to extend beneath the pelvis during the push-off phase of significant changes associated with the transition into
bipedal locomotion (Fig. 1.11). Ardi’s foot represents a efficient bipeds, but the skull and facial features remain
true evolutionary mosaic, as her opposable great toe that primitive and more closely resemble orangutans.
remained fully functional for arboreal climbing while her
lesser metatarsals were structurally reinforced so they Australopithecus afarensis
could tolerate the accelerational forces associated with the Better known by her nickname Lucy (after the
propulsive period of walking. This unusual arrangement Beatle’s song that frequently played at the archeological
completely disappears in later hominids as the first site), Australopithecus afarensis actually translates into
metatarsal moves into a centralized position and becomes “Southern Ape from Afar”, as she was discovered in the
significantly wider and stronger, supporting the majority of Afar region of Northeast Ethiopia. While fragments of
weight during the propulsive period of walking. In fact, a Australopithecus afarensis were found as early as 1935,
recent cadaveric study by Jacobs et al. (22) confirmed that the discovery of Lucy by Donald Johanson in 1973 was
during the push-off phase of modern human locomotion, significant as it included pieces of her pelvis, femur, and
the first metatarsal head supports 119% of body weight (this tibia. In 2000, only 4 km from where Lucy was discovered,
includes accelerational forces) while the second metatarsal paleoanthropologists unearthed a significant portion of
head supports only 28% of body weight. The Ardipithecus the skeletal remains of a 3-year-old Australopithecus
foot represents the proverbial “missing link” in evolution, afarensis that became known as Lucy’s baby (which is
as it retains qualities common to modern humans and ironic as carbon dating revealed the child was more than
8
Figure 1.12. Posterior views of the left distal humeri and proximal tibiae in different species. Note the deeper
olecranon fossa in the chimpanzee humerus that allows the ulna to lock in place during knuckle-walking. Because the
chimpanzee’s proximal tibia is T-shaped, it is unable to effectively bear weight during bipedal locomotion. In Australopithecus
anamensis and Homo sapiens, the proximal tibial metaphysis flares out providing an increased quantity of cancellous bone
that significantly improves the ability to absorb shock. Partially adapted from photographs in Leakey M, Walker A. Early
hominid fossils from Africa. Scientific American. 2003;13:14-19.
100,000 years older than Lucy). Originally nicknamed the to decelerate the forward swing of the leg significantly
Dikka Child after the region in Ethiopia in which she was improved. This simple change would have provided
discovered, she was later renamed Selam, which means Lucy with a significant metabolic advantage over her
peace in several Ethiopian languages. predecessors, particularly for long distance foraging and
The 3-year-old Selam, who was most likely buried walking at faster speeds. To emphasize the importance of
shortly after death by a flood, was discovered with a even small increases in efficiency as the day ranges for
remarkably intact skeleton possessing the complete skull foraging increased, Leonard and Robertson (23) evaluated
and mandible, the entire torso and significant segments of every aspect of Lucy’s skeleton and created a model to
both upper and lower extremities including the patellae. determine energy savings as Lucy walked as a biped and
By piecing together bony fragments from Lucy and a quadruped. By calculating Lucy’s assumed speed as a
Selam, it becomes clear that Australopithecus afarensis biped (believed to be 2.3 miles per hour [14]) the authors
was most likely an accomplished biped. Although Lucy’s considered total daily expenditures for male and female
lumbar vertebrae suggested occasional support from her afarenses over a variety of day ranges and concluded that
forearms during bipedal gait, changes in her pelvis and the metabolic savings associated with bipedality would
lower extremity revealed that she was a very efficient have been relatively small over the shorter day ranges but
walker (15). By meticulously casting and rebuilding were substantially larger as day ranges increased. Because
Lucy’s innominate (which had been crushed and fused Lucy existed 3 million years ago when food sources were
into a single mass of matrix stone with about 40 separate becoming more spread out, and because modern hunter-
pieces of the innominate inside), Lovejoy (17) was able gatherers have day ranges for foraging of approximately 8
to recreate the pelvis of our 3.1-million-year-old ancestor. miles per day, Leonard and Robertson (23) calculated that
Compared with Ardipithecus, the most obvious change in the energy savings for the larger day ranges could result
Lucy’s pelvis was in the angle of her ischial tuberosities. in as much as an 8% reduction in energy expenditure per
Because they angled posteriorly in a manner similar to day. This significant caloric savings could have provided
modern humans, the ability of the hamstring musculature considerable benefit favoring bipedal locomotion as the
9
saved calories could have been used for improved growth Lovejoy (17) supports the theory that the newly
and reproduction. flared ilia improves stability by comparing CT scans from
Lovejoy’s (17) analysis of Lucy’s pelvis also revealed chimpanzee and modern human femoral necks with a
that her sacrum was becoming wider and her ilia became scan from Lucy. Because the chimpanzee’s femoral neck
more dish-shaped (Fig. 1.13, A and B), flaring laterally lacks the stability provided by the lower gluteus medius
even farther than Ardipithecus’s ilia (20). This significantly and piriformis muscles, the femoral neck is subjected
improves bipedal efficiency by increasing the mechanical to a “greenstick effect,” in which the superior surface is
advantage afforded the gluteus medius muscle for resisting exposed to tensile strains (Fig. 1.14, A). As a result, as is
vertical forces associated with single-limb stance (Fig. consistent with Wolff’s law, there is greater cortical bone
1.13, C). By having the ilia flare farther laterally, Lucy was formation present in the superior aspect of the chimpanzee
able to use her gluteus medius to stabilize the vertical upper femoral neck as it remodels to provide support at the site
body during single-limb support, and walking suddenly of tension. In contrast, modern humans have a significant
became an efficient form of transportation. decrease in tensile strains placed on the femoral neck
Figure 1.13. Superior views of a chimpanzee pelvis

(A) and Lucy’s pelvis (B and C). The outward flare of Figure 1.14. The weight of the torso during single-leg
Lucy’s pelvis allows gluteus medius (arrow X) to resist stance (W), creates a bending force on the femoral neck
the downward motion of the weight-bearing torso (arrow in which the upper and lower portions are exposed to
Y). This represents a class one lever comparable to a significant tensile and compressive loads, respectively
playground seesaw in which the fulcrum is the hip and (small black arrows in A). In Lucy and in modern humans,
forces on the opposing sides are body weight and the pull of the piriformis and lower gluteus medius muscles (small
gluteus medius. The length of the femoral neck determines white arrows in A) create a powerful compressive force that
the lever arm afforded gluteus medius (think of one person reduces the femoral neck bending moments. The ability
on a seesaw moving farther away from the fulcrum). of these muscles to lessen bending strains is evidenced
Interestingly, Lucy’s femoral neck was longer than a modern by the reduced quantity of cortical bone in the upper
human’s femoral neck. Modified from Lovejoy O. Evolution femoral neck (compare the dark lines in B-D). Modified
of human walking. Scientific American. November 1988: from Lovejoy O. Evolution of human walking. Scientific
118-125. American. November 1988:118-125.
10
because the lower fibers of gluteus medius and piriformis

create a stabilizing compressive force that lessens the
need for added support from dense cortical bone. Upon
analyzing Lucy’s femoral neck, Lovejoy noted a pattern of
cortical bone formation similar to modern humans, which
strongly supports the theory that Lucy was an efficient
biped (see Fig. 1.14, B-D).
Also supporting Lucy’s status as a dedicated biped
are the famous Laetolli footprints. Discovered in Tanzania
by Mary Leakey and her team of archeologists (24),
these 3.2-million-year-old footprints were formed as 3
Australopithecus afarensis walked along a steep slope of
volcanic ash. Despite a significant incline, there was no
evidence of hand contact with the ground confirming that
these hominids were indeed bipedal. The only consistent
finding with these footprints was that all 3 afarenses
walked with significant toe-out gait patterns. Also, while
the Laetolli footprints lacked the detail necessary to Figure 1.15. In modern humans and Lucy, a bony groove
evaluate shape of the medial longitudinal arch, recent for the tibialis anterior tendon is located distally, while
research suggests that Australopithecus afarensis, like in gorillas and chimpanzees, the groove is positioned
modern humans, exhibited variation in arch structure: proximally. Partially modified from photographs in Latimer
some afarenses possessed well-developed arches, while B, Lovejoy O. Am J Phys Anthropol 1990:125-133.
others were flat-footed (45).
In addition to the occasional well-formed arch,
Lucy’s skull and lower extremities possess a number of
changes consistent with frequent bipedality. Her foramen cuneiform. According to Latimer and Lovejoy (25), the
magnum is centrally located and the orbits of her eyes distal location of this groove is essential for bipedality,
parallel her line of vision. The metaphysis of her proximal because it allows tibialis anterior to physically block
tibia flares out allowing improved dissipation of ground- migration of the first metatarsal. The authors note that in
reactive forces through the knee during upright walking. In less efficient bipeds such as chimps and apes, the groove
the foot and ankle, the calcaneus was becoming wider and a for tibialis anterior is positioned proximally, making the
deep groove was forming in the posterior fibular malleolus first metatarsal unstable (Fig. 1.15). This would allow
and in the trochlea of the calcaneus for the peroneal the first metatarsal to shift when exposed to the forces of
tendons. Latimer and Lovejoy (25) claim that these bony propulsion.
changes are consistent with a greater percentage of ankle Although the distal articular surface of the medial
plantarflexion force coming from the peroneal musculature cuneiform has a convexity midway between humans and
rather than the Achilles tendon. chimpanzees, the proximal aspect of the first metatarsal
One of the most significant anatomical clues possesses a double concavity with an invagination of
suggesting that Lucy was a biped is the distinct groove for the articular surface that limits axial rotation of the first
tibialis anterior located on the distal aspect of her medial metatarsal shaft (Fig. 1.16). This is an extremely important
Figure 1.16. Posterior view of the left first metatarsal. Compared to the chimpanzee first metatarsal, both Lucy and
modern humans possess a bony ridge that serves to lessen axial rotation. Note the distal attachment of the tibialis anterior
tendon to the plantar medial aspect of the proximal first metatarsal in both Lucy and modern humans. Modified from Latimer
B, Lovejoy O. Hallucal tarsometatarsal joint in Australopithecus afarensis. Am J Anthropol. 1990:125-133.
11
change as it stabilizes the medial forefoot and creates a bony lower extremity, Lucy serves as an excellent example of
block that lessens muscular strain during the propulsive “mosaic evolution” in that she is exchanging the grasping
period. The changes present in Lucy’s feet suggest that skills of the foot (which Ardipithecus found so invaluable)
she is giving up her opposable great toe in exchange for in favor of improved stability necessary for bipedality (Fig.
improved efficiency during bipedal locomotion. Because 1.17).
Lucy’s hip could not extend as far back as modern humans,
she was unable to create a significant force during push- Australopithecus garhi
off and her gait pattern most likely resembled something Because paleontologists were searching for an
between modern humans and chimps. Her thick radial shaft Australopithecus afarensis when they discovered the bony
cross-sections confirm that despite the lack of handprints fragments of this hominid, they named the species garhi
present in the Laetolli footprints, Lucy frequently used from the Afar language meaning “surprise.” Although only
her forearms for support (15). Because of her blend of a few skeletal fragments have been discovered, the length
anatomical features allowing support from both upper and ratios of the upper and lower limbs are more humanlike
than Lucy. The midshaft diameter of the radius suggests
that garhi spent less time knuckle-walking than Lucy and
fragments from the tibia and foot are close in size and
shape to Australopithecus afarensis so it is assumed that
garhi moved mostly upright and was habitually bipedal.
Australopithecus africanus
In 1924 a South African miner working in the
Taung quarry near Johannesburg stumbled upon the skull
of a new species of hominid. Australopithecus africanus
literally translates into “The Southern Ape from Africa.”
Nicknamed the Taung child by the paleontologist Raymond
Dart, the skull belonged to a young child and consisted of
almost a complete face, a few teeth and a section of the
braincase. Subsequent discoveries include another nearly
complete skull, which was remarkably humanlike with
its reduced jaw length and fairly vertical face. The ratio
of upper extremity length to lower extremity length was
.85, which is about halfway between modern humans
(.76) and pygmy chimpanzees (.92). The tibiae are shorter
than the femurs and various foot bones suggest that
Australopithecus africanus had a very mobile foot with
an abducted first metatarsal and no significant presence
of a medial longitudinal arch (15). Analysis of the lumbar
spine revealed small articular cross-sectional areas with a
flattening of the lumbar lordosis suggesting limited upright
walking. When compared to Lucy, Australopithecus
africanus had limited hip extension and thicker cortical
bone in the forearms, strongly suggesting that this species
was not habitually bipedal and most likely stood upright
primarily when feeding.
Figure 1.17. Lucy walking. With her centralized foramen
magnum (A) Lucy maintains a horizontal gaze while walking. Paranthropus robustus
Her thick radial cortex (B) confirms that she frequently used Accidentally discovered by a South African
her forearms for support while her elongated and curved
schoolboy in the 1930s, this hominid was named
fingers confirms that she spent a significant amount of time
Paranthropus because it is from a side branch of human
grasping branches and climbing. Compared to modern
humans, her limited range of hip extension (C) would have lineage, and robustus because of its robust teeth, skull,
shortened her stride length and she most likely would have and jaw. Paranthropus stands out among early hominids
walked with exaggerated knee flexion during initial ground because bone tools were found at various archeological
contact (D). Because her talar neck was everted, the joints sites containing skeletal remains of this hominid. Because
of the mid and forefoot may have been able to tolerate the of heavy wear patterns along the tips and scratches along the
forces of propulsion without buckling (E). shafts, it is theorized that these early bone tools were used
12
to dig for tubers and/or possibly termites (15). The most

important finding relating to bipedality was the discovery
in 1981 of a complete, undistorted first metatarsal (26).
The morphology of this bone was remarkably humanlike
in that it had a groove for the tibialis anterior tendon, a
prominent tubercle for peroneus longus and a small area
along the lateral aspect of its base that served as a contact
point for the second metatarsal. The proximal portion of
the base also possessed separate articular surfaces that
limited mobility and the shaft was short and stout: strongly
indicative of bipedal gait. When viewed distally, the lower
section was appreciably flatter than a chimpanzee’s first
metatarsal head (compare X and Y in Fig. 1.18). Susman
and Brain (26) argue that the inferior aspect of the first
metatarsal head flattened in response to the increase in first
metatarsophalangeal compressive forces associated with
more frequent upright locomotion; i.e., when the heel leaves
the ground while walking, the distal first metatarsal head
is pushed into the proximal phalanx, which is maintained
in a fixed position by the plantar ligaments, including the
plantar fascia. The authors state that over time, the repeated
intrinsic joint compression flattened the metatarsal head.
Susman and Brain (26) state that flattening of the first Figure 1.18. Distal view of the first metatarsal head
metatarsal head represents the first sign of a functional in pygmy chimpanzee, Paranthropus robustus, and
metatarsophalangeal joint with the presence of a windlass modern human. Dotted lines represent transverse plane
mechanism described by Hicks (27). bisections. While the inferior aspect of the Paranthropus
The most important factor arguing against robustus’s first metatarsal head is relatively flat (compare
significant efficiency during bipedal locomotion is the X and Y), the superior aspect remains narrow compared
relative dorsal narrowing of the superior distal aspect of the to modern humans (double arrows). Redrawn from
first metatarsal head (compare B and C in Fig. 1.18). This photographs in Susman R, Brain T. Am J Phys Anthropol.
dorsal narrowing suggests the first metatarsophalangeal 1988;77:7-15.
joint would be unstable when the hallux reached its full
range of dorsiflexion, which would significantly limit femoral neck angle, thickness, and shape of the femoral
this hominid’s ability to apply force during the final diaphysis (more teardrop-shaped with a greater anterior
moments of the propulsive period. In humans, the wider to posterior thickness) and improved alignment between
distal surface allows for a locking mechanism in which the femur and tibia all provide significant advantages for
the metatarsophalangeal joint maintains stability as forces efficient bipedality. The patellae face forward when the hip
peak during propulsion. This finding alone would indicate is extended and there is a humanlike ankle that is aligned in
that Paranthropus robustus most likely did not have an the transverse plane so the foot moves forward in the same
effective terminal push-off and the limited articular joint plane as the lower extremity; i.e., unlike Lucy who walked
surface area present in the lumbar spine along with a with a significant toe-out, Homo rudolfensis walked with
relatively unstable hip joint suggests that Paranthropus knees and feet moving forward in the sagittal plane.
was not a particularly efficient biped and frequently walked An extremely important by-product of Homo
on all fours (15). rudolfensis’s habitual bipedality is that it decoupled the
process of breathing from locomotion. As noted by Carrier
Homo rudolfensis (28), running quadrupeds time their respiration with their
Named after Lake Rudolf in Western Kenya where strides: when their legs extend forward they inhale and
this hominid was first discovered, Homo rudolfensis when they hit the ground and their muscles contract to
possessed such a large braincase that it was the first of absorb shock, they exhale. This coupled pattern locks the
our ancestors to be classified as Homo (which is Greek for breathing cycle during quadrupedal locomotion to the stride
“same”). Skeletal remains of the pelvis and lower extremity rate. In a full-time biped, breathing is no longer coupled
suggest that Homo rudolfensis spent almost all of its time with locomotion, which is essential for the development
walking on two legs. The lumbar facet and hips have of speech as it allows for an adjustable rate of airflow.
increased surface areas capable of supporting body weight Whether or not our hominid ancestors began talking
for long periods without assistance from the arms. The 50,000 years ago or 5 million years ago is the subject of
13
Figure 1.19. In modern humans (A), heel lift occurs with the cuboid pivoting about the calcanean process until
its dorsal border contacts the overhanging calcaneus. This action essentially locks the midfoot during the propulsive
period and allows for an efficient transfer of force from rearfoot to forefoot during propulsion. In a chimpanzee (B), the first
stage of heel lift occurs with the cuboid maintaining ground contact (arrow): the calcaneocuboid joint fails to lock and the
midfoot buckles under the forces of propulsion. The second stage of heel lift occurs when the cuboid leaves the ground
transferring body weight to the forefoot through a collapsed and buckled arch.
debate but it is clear that the process of walking on two legs prominence that projects proximally into the calcaneus
was responsible for the eventual development of language. and serves as a locking mechanism for the lateral column
(Fig. 1.19, A). Unlike chimpanzees who have a double
Homo habilis push-off where the calcaneus initially leaves the ground
“The Handyman” was named by Raymond Dart followed shortly thereafter by the cuboid, human heel
because it was the first hominid to routinely use tools lift occurs as a single event with the calcaneocuboid joint
and its increased brain size suggested that this hominid locking so the midfoot does not contact the ground during
was “mentally skillful”: habilis is Latin for “able, handy, propulsion (Fig. 1.19, B). As described by Bojsen-Moller
mentally vigorous.” Discovered in Tanzania in 1959, Homo (29), the calcaneocuboid joint locking mechanism provides
habilis is somewhat controversial because it has many significant stability as the cuboid pivots about its medial
characteristics of Australopithecus afarensis (e.g., Lucy), plantar prominence until its dorsal surface collides with
causing some paleontologists to argue whether it should be the overhanging calcaneus, thus providing a bony locking
classified as a Homo or if it would be more appropriately that lessens muscular strain. Kidd et al. (30) demonstrate
named Australopithecus habilis (15). For example, its hand that the calcaneocuboid joint locking mechanism in Homo
is wide and humanlike but its fingers possess a chimp-like habilis was even better developed than in modern humans,
curvature with the thumb rotated in a manner similar to perhaps because the toe-in gait placed greater stress on the
a great ape. The midshaft diameter of the radius is thick lateral column.
confirming that Homo habilis was not a full-time biped. In Morphological evaluation of a Homo habilis foot
addition, the lower tibia and ankle internally rotate in the revealed an orangutan-like talus with an ape-like navicular
transverse plane, which would have forced Homo habilis to (30). The first metatarsal was remarkably humanlike as the
walk with a significant toe-in gait pattern. In fact, Sawyer hallux aligned with the lesser metatarsals and the dorsal
et al. (15) note that modern humans that are as pigeon-toed surface of the first metatarsal head significantly widened,
as one particular specimen of Homo habilis are unable to which would have allowed the first metatarsophalangeal
maintain balance while walking. Other skeletal remains joint to be stable through a full range of motion during the
of Homo habilis are consistent with a more humanlike propulsive period. Whether or not Homo habilis should
form: a centrally located foramen magnum with a slight be classified as an Australopithecus or Homo may be the
indentation in the posterior skull for the nuchal ligament, subject of debate, but the development of a stable first ray,
suggesting that this hominid frequently walked upright a calcaneocuboid joint locking mechanism that allowed
with a balanced head. for a single-phase push-off, along with a dorsally widened
An almost complete Homo habilis foot discovered first metatarsophalangeal joint that could provide stability
in 1960 has been extensively studied and it possesses during terminal propulsion are all extremely important
many humanlike qualities. The calcaneocuboid joint advances in the process of becoming efficient bipeds.
was remodeled so the cuboid possessed a medial plantar
14
Homo ergaster a thickening of the ridge (the nuchal ridge is absent both
Named from the Greek word meaning “workman” in chimpanzees and Australopithecus afarensis). To
(numerous tools have been found at various archeological support their case, the authors note that a computerized
sites) this hominid should have more appropriately been tomographic evaluation of a Homo erectus skull revealed
named “the traveler” as it was one of the first hominids a dramatic increase in the size of the semicircular canals
to leave Africa (31). Evaluation of a 1.5-million-year- (33). Because running often lacks a double-limb support
old skeleton of a 12-year-old boy revealed that Homo phase in which both feet are on the ground (i.e., the
ergaster was surprisingly humanlike in appearance. Unlike airborne phase is followed by a precarious single-limb
habilis, ergaster had limb proportions comparable to support phase), the improved balance provided by the
modern humans. In fact, the tibiae of Homo ergaster were large semicircular canals is essential for stability. Other
actually longer than ours, which could have allowed for authors have noted that chimpanzees lack a well-developed
more efficient bipedality. Because the growth plates of the semicircular canal, which partially explains why they are
young boy had not closed, it was impossible to ascertain incapable of balancing on one leg during static stance. This
adult articular ranges of motion but the transverse plane would also have been true for Australopithecus afarensis,
alignment of the lower extremity was similar to Homo as computerized tomographic evaluations of Lucy’s skull
rudolfensis in that the foot and leg moved together in the revealed poorly developed semicircular canals (32).
same transverse plane; i.e., unlike Lucy with her toe-out Besides changes in skull shape and the inner ear, the
gait and habilis with the extreme toe-in, ergaster walked upper extremities in Homo erectus also revealed significant
with a straight gait pattern and was most likely a very changes consistent with endurance running, as there was
efficient biped. a 50% reduction in the mass of the forearms relative to
body mass and the scapulae were less cranially oriented
Homo georgicus (32). These changes would have made Homo erectus
Discovered in volcanic deposits in the medieval less skilled at climbing but would have been invaluable
town of Dmanisi in southeastern Georgia, Georgicus is the for running, since the wider shoulders and shorter arms
first hominid discovered outside of Africa. Although recent would have allowed for independent counter rotation of
findings include a femur, patella, lower leg and various the upper extremity and pelvis while also reducing weight
foot bones, these bony fragments have yet to be studied in of the upper extremity: both of which are important with
detail. The prior discovery of a 1.8-million-year-old Homo endurance running.
georgicus third metatarsal was significant, as it was small The spine and pelvis of Homo erectus had also
but stocky and possessed a large tubercle at its base. This undergone significant changes compared with prior
may have been associated with the development of an early hominids, as the ratio of articular surface area to body
medial longitudinal arch (15). mass significantly increased in the lumbar spine, sacroiliac
joint, femoral head and tibiofemoral joints. These changes
Homo erectus would have been essential in order to handle the increased
Named erectus because of its upright stature and ground-reactive forces associated with running. The femur
narrow frame, this hominid had a remarkably long species of Homo erectus is particularly interesting as it is identical
lifespan with fossil remnants dating from 1.9 million years to that of modern humans: e.g., the femoral neck angle is
ago to as recently as 27,000 years ago. In a beautifully written 120° with approximately 10° of anteversion (which allows
article detailing specific evolutionary traits associated the patellae to face forward when the hip is extended),
with the development of bipedal locomotion, Bramble while the midshaft cross section is teardrop-shaped and
and Lieberman (32) argue that Homo erectus was the first the posterior aspect of the greater trochanter possesses
hominid capable of endurance running. This is important thickened ridging at the attachment points for the gluteal
as no primates other than humans are capable of endurance muscles.
running and the ability to jog long distances would have Compared to Australopithecus afarensis, there
provided significant advantages when scavenging for food. was a reduction in the length of the femoral neck (which
The authors support their hypothesis by noting that the skull decreased metabolic efficiency while walking because
of Homo erectus has a significant ridge along the base of the it lessened the lever arm afforded gluteus medius) but
occiput where the nuchal ligament attaches. They suggest the shorter femoral neck was essential for running since
that this ridge would only be present if Homo erectus was it decreased the bending moments on the neck during
capable of endurance running, because running results in single-limb support, thereby reducing the risk of femoral
a slight but rapid deceleration of the body following heel neck stress fracture. Interestingly, the tibiae of the Homo
strike that causes the skull to pitch forward abruptly with erectus were identical to modern humans except for the
each stride. The increased forward head pitch coupled with fact that they were more robust. Although there are no
the oscillations present at heel strike would significantly skeletal remains of Homo erectus feet, the recent discovery
stress the nuchal ligament insertion, eventually producing of 1.5-million-year-old Homo erectus footprints in Ileret,
15
Kenya, revealed well-defined medial longitudinal arches Homo floresiensis

and a modern humanlike angle of abduction of the Discovered in 2003 on the remote Indonesian island
hallux: the angle between the hallux and shaft of the first of Flores (hence the name Homo floresiensis), this enigmatic
metatarsal in the Homo erectus footprints averaged 14°, hominid stood only 3 feet 6 inches tall and possessed such
which is comparable to the modern human angle of 8° a small brain case that it was initially incorrectly suggested
(34). In contrast, the abduction angle of the hallux in the that the skull belonged to a microcephalic Homo sapien
Australopithecus afarensis footprints discovered in Laetoli (Fig. 1.20). Because Homo floresiensis lacked the bony
averaged 27° (34). Additionally, the path of the prints characteristics typically seen in pygmies and pituitary
paralleled the direction of travel and a detailed optical dwarfs (both of which have small bodies and large brains),
laser analysis of pressure patterns revealed a greater depth researchers determined that Homo floresiensis was not
of print beneath the medial forefoot, suggesting a medial related to Homo sapiens and must have descended from a
weight transfer before the push-off phase of walking prior hominid. It was assumed that this hominid managed
(34). These findings are consistent with a long, modern, to drift to the remote island at least 800,000 years ago
humanlike stride with push-off occurring over extended (which is significant since there is no evidence of boat
lower limbs and stable first metatarsophalangeal joints. building until 50,000 years ago) and, after being marooned
Typically standing 5 feet 2 inches tall and weighing for several hundred thousand years, evolved into the dwarf
only 110 lbs, Homo erectus developed a narrow, elongated species Homo floresiensis.
form that lessened solar exposure and increased heat The process of dwarfing with isolation seems
dissipation while running. The skull possessed accessory implausible but it has been repeatedly demonstrated that
“vaults and expansions” that would have allowed for when animals are isolated with limited predation and/or
improved venous cooling in the face and scalp. By competition for food, large animals become small and small
combining all of these features, Bramble and Lieberman animals become large. The reason for this is that small
(32) make a very strong case arguing that Homo erectus creatures such as rabbits and rats survive in conventional
was indeed an efficient endurance runner and was the first environments by becoming smaller and producing so many
“fully terrestrial hominoid.” offspring that despite predation, they survive because of
Homo erectus’s proficiency at endurance running their population density. Conversely, large animals such as
may have also been indirectly responsible for brain elephants and rhinos have a greater chance for survival if
expansion, since the improved scavenging skills provided they become so big that predators will no longer attack.
by endurance running would have allowed Homo erectus The only problem with this hypothesis is that the brain
to outperform other hominid and primate competitors: of Homo floresiensis was so exceedingly small that it
although non-human primates can sprint, they are not was disproportionate to all known scaling relationships
capable of long distance running (35). Because numerous associated with isolation dwarfing.
early Homo erectus archeological sites show an increase The mystery of Homo floresiensis’s tiny brain
in the presence of animal bones, it is suggested that for the was recently solved by researchers from the Natural
first time in hominid evolution, meat was now part of the History Museum in London. By analyzing fossils from
diet. Leonard (36) states that because the brain consumes hippopotamuses that had been isolated on the island of
16 times the calories of an equivalent mass of muscle, and Madagascar, Weston and Lister (37) demonstrated that
because resting brain metabolism in humans consumes compared with conventional isolation dwarfing, island
approximately 25% of the total caloric intake (in most isolation results in significantly greater reductions in
mammals the brain consumes 3 to 5%), the addition of
meat into the diet would be essential to provide the calories
necessary for brain expansion; e.g., meat provides up to 4
times the calories of an equal sized serving of fruit (36).
In addition to incorporating meat, Homo erectus was the
first hominid to utilize fire to heat food. Cooking played
an important role in brain expansion because it allowed for
a greater percentage of nutrients to be metabolized by the
body: 100% of cooked food is metabolized while only 30-
40% of the nutrients present in raw food can be digested.
The combination of improved foraging skills associated
with running and the increased nutritional gains associated
with cooking enabled Homo erectus to double its brain size
in just 600,000 years. Figure 1.20. The microcephalic skull of Homo
floresiensis.
16
brain versus body size, explaining how a large-brained ago, it is known that Homo floresiensis and Homo sapiens
hominid could have evolved into the microcephalic Homo coexisted for at least 28,000 years. It is unclear whether
floresiensis. Homo floresiensis was simply absorbed into the Homo
The only question that remained was figuring out sapien population or if they became extinct because of
which hominid evolved into Homo floresiensis. Early disease or an inadequate ability to compete for food. Despite
research suggested that Homo erectus was the most likely a brain volume comparable to a modern chimpanzee, Homo
predecessor, mainly because it was assumed that only a floresiensis used remarkably sophisticated stone tools,
biomechanically efficient hominid could travel all the way such as blades, awls, punches and even micro-blades for
to Asia. Recent fossil evidence questions this view since big game hunting, which made some researchers question
analysis of multiple Homo floresiensis skeletons makes it the connection between brain size and intelligence. This
clear that this mysterious hominid bore a much stronger paradoxical relationship was made clear when virtual
resemblance to Lucy and other australopithecines than reconstructions of skull fragments revealed that despite
to Homo erectus. Evaluation of skeletal remains from its small skull, Homo floresiensis possessed an enlarged
14 different individuals revealed that this tiny hominid Brodmann’s area 10, a section of the frontal lobe associated
had a pelvic structure similar to Lucy’s (confirming that with complex cognitive function (40).
Homo floresiensis did indeed walk upright), with an
appendicular skeleton that more closely resembled the Homo antecessor
early australopithecines (e.g., an ape-like wrist with short While cutting a railroad path in Northeast Spain,
femurs and tibiae). workers unearthed skeletal remains that included an
Perhaps the strangest feature of the Homo floresiensis upper jaw and a few teeth (the fragments were carbon-
skeleton was the foot. Despite a hallux that aligned with dated to approximately 800,000 years ago). In 1997 it was
the first metatarsal, the rest of the foot was extremely determined that these fragments belonged to a new species
primitive and bore a much stronger resemblance to the foot of hominids named Homo antecessor (from the Spanish
of a Bonobos chimpanzee in that it completely lacked a word for ancestor), since this Homo was initially believed
medial longitudinal arch and possessed long, curled toes to be the ancestor of all modern Europeans. Although little
that would have been useful for grasping (38). By far, the is known about the joint surface areas and upper vs. lower
most unusual characteristic of the Homo floresiensis foot extremity length ratios, skeletal remains from the hands
was its length: while modern humans typically have a and feet suggest that Homo antecessor was an efficient
foot measuring 55% the length of their femur, the Homo biped, comparable to modern humans (with the exception
floresiensis foot measured 70% the length of the femur that the patellae were significantly smaller) (15).
(38). Such an extremely long foot would have forced
Homo floresiensis to walk with a steppage gait in which the Homo heidelbergensis
hips flexed excessively in order to allow the toes to clear The jawbone Homo heidelbergensis was unearthed
the ground during midswing (comparable to walking while in 1903 in a quarry near Heidelberg, Germany. Evaluation
wearing swim fins). Although they may have been able to of scratch marks on the teeth suggests that this hominid
run, it would have been for extremely short distances and used stone tools to cut objects held in its mouth. Oddly,
only in emergencies. the teeth also revealed wear patterns consistent with
In an attempt to determine the hominid predecessor toothpick use (15). Standing nearly 5 feet 8 inches tall
of Homo floresiensis, Argue et al. (39) used a technique with a species lifespan from 700,000 until 200,000 years
called cladistics, in which a wide range of shared physical ago, Homo heidelbergensis was the common ancestor to
traits are evaluated to establish relationships between both Homo neanderthalensis and Homo sapiens. Despite
organisms. Using this technique, researchers determined its relationship to modern humans, Homo heidelbergensis
that Homo floresiensis most likely evolved sometime was almost identical in appearance to neanderthalensis and
between Homo rudolfensis and Homo habilis and found no skeletal remains revealed an extremely wide pelvis with
connection between the tiny hominid and Homo erectus. the lower extremity bones being shorter and stockier than
The results of their findings are significant because it proves modern humans.
that relatives of this large-footed, small-brained hominid The skeletal proportions of Homo heidelbergensis
somehow managed to leave Africa almost 2 million years were most likely the by-product of its environment, because
ago, eventually making its way to this remote Indonesian 500,000 years of living in a cold climate eventually allows
island. To confirm this theory, paleoanthropologists are natural selection to favor a skeletal shape in which the torso
searching for Homo floresiensis skeletons at other sites in gets larger and the extremities get shorter as this lessens the
Asia. surface area to volume ratio. This in turn favors reduced
Because Homo floresiensis survived until 17,000 heat dissipation because the smaller the surface area, the
years ago, and because skeletal remains of Homo sapiens less body heat lost to the environment through convection.
have been found on Flores dating back to 55,000 years
17
Homo neanderthalensis
The prototypical caveman, almost all skeletal
remains of Homo neanderthalensis have been recovered
from cave and rock shelters found throughout Europe
and Asia. Although a Neanderthal infant was discovered
in Belgium in 1829, the first adult skeletal remains
were found by miners in the Neander Valley of northern
Germany (tal is old German for valley while thal replaced
tal in the early 1900s as the German word for valley).
Skull fragments revealed a particularly short forehead with
independent bony thickenings over the orbits of each eye
giving Homo neanderthalensis the characteristic caveman
appearance often depicted in cartoons and the media.
Although originally considered a primitive brute (skeletal
remains revealed routine cannibalism), it is now known that
Neanderthals buried their dead with rituals that included
specific positioning, the use of markers and perhaps even
the incorporation of flowers with the deceased. Sawyer et
al. (15) note that Homo neanderthalensis was often blue-
eyed and frequently possessed blond and/or reddish hair
and were most likely capable of speech.
The species lifespan for Homo neanderthalensis
was from 175,000 to as recently as 27,000 years ago.
Expanding glaciers throughout Europe and Asia during
that time suggest that neanderthalensis was exposed to
snow almost year-round. As with Homo heidelbergensis,
consistent exposure to cold for tens of thousands of years
allowed natural selection to favor a skeletal shape that
lessened heat loss: the extremities (particularly the fingers,
forearms and legs) were significantly shorter than modern
humans while the torso was long, wide and barrel-chested.
These combined features make Homo neanderthalensis
a little shorter than modern humans but about the same Figure 1.21. Homo neanderthalensis walked with an
weight. Because of their short tibiae and stocky femurs upright posture similar to modern humans. The primary
(which had reduced femoral neck angles), it was originally difference in skeletal structure relates to the bulging brow
ridge (A), widened rib cage (B), short forearms (C), reduced
assumed that Homo neanderthalensis walked with a
femoral neck angle (D), stocky femurs (E) and shortened
stooped, bent knee gait. However, the recent discovery of
tibiae (F). The shorter legs and excessive muscle mass
osteoarthritis in the weight-bearing joints of a particularly significantly increased the metabolic cost of locomotion.
old Neanderthal confirms that they most likely walked
upright in a manner similar to modern humans (although
their shorter lower extremities resulted in reduced stride Neanderthal genetic code to modern humans throughout
lengths making for a fairly inefficient gait) (Fig. 1.21). the world, the authors concluded that interbreeding
Because Homo neanderthalensis and modern humans most likely took place between 50,000 and 80,000 years
inhabited the same geographic location for thousands of ago, somewhere in the Middle East. The offspring of
years, it was always suspected that occasional interbreeding Neanderthals and humans quickly spread: people from
between Homo sapiens and Homo neanderthalensis might Papua New Guinea (where Neanderthals never lived) have
have occurred. Although early DNA research suggested just as much Neanderthal DNA as people from France. The
otherwise, a team of researchers from the Max Planck only modern humans devoid of Neanderthal DNA are from
Institute for Evolutionary Anthropology in Leipzig Africa, since interbreeding occurred after our ancestors
Germany evaluated 4 billion nucleotides from Neanderthal crossed into Eurasia.
bone fragments found in a cave in Croatia and conclusively Despite occasional interbreeding, the Neanderthal
demonstrated that interbreeding did occur (45). In fact, population slowly dwindled, becoming extinct
between 1 and 4% of modern human DNA comes directly approximately 30,000 years ago. While it was originally
from Neanderthals, including genes coding for cognitive assumed that Homo sapiens unwittingly caused the
function and skeletal development. By comparing the extinction of Homo neanderthalensis through competition
18
for food and/or by the spread of disease, more recent the length of the forefoot lever arm depending upon the
research suggests that metabolic inefficiency may have locomotor requirements; e.g., walking up a steep hill
played an important role in their demise. In a detailed is more efficient when pushing off the lateral forefoot
study of the energetic cost of locomotion as related to (low gear), while rapid sprinting is best accomplished by
lower limb length, Steudel-Numbers and Tilkens (42) pushing off the medial forefoot (high gear) (Fig. 1.23).
prove that the Neanderthal’s shorter tibiae would have Compared with our ancestors, the tibiae became
increased the metabolic cost of locomotion by at least longer, the toes became shorter (which reduced weight as
30%. Froehle and Churchill (43) went on to calculate the we no longer needed them for grasping) and the patellae
daily energy expenditure of Neanderthals versus modern became wider, allowing for improved stability of the knee
humans and determined that because of their stockier in the sagittal plane. The femoral neck angles increased to
frames and increased muscularity, Homo neanderthalensis an average of 130°, which reduced bending strains on the
had to spend anywhere from 100 to 350 kcal/day more neck and improved efficiency of the gluteus medius muscle
than their Homo sapien relatives. While the Neanderthal’s during single-leg stance. A slightly adducted femoral
greater body mass and increased muscularity significantly shaft (coxa varum) positioned the femur directly over
improved thermoregulation and made them effective big the perpendicular tibia effectively converting the lower
game hunters, their inability to survive on non-meat food extremity into a stable vertical column thereby lessening
sources (meat was becoming increasingly sparse as rapid
changes in climate reduced the number of large mammalian
prey) coupled with their higher cost of locomotion may
have created an environment where they were spending
more calories on transportation then they were able to
consume. As noted by Froehle and Churchill (43), the
biomechanically more efficient modern humans “may have
been able to convert their energetic savings into a slight
reproductive advantage” thereby ensuring their survival.
Homo sapiens
“The Knowing Man” (sapiens translates from the
Latin “to know”) represents us: the remarkably successful
species of modern humans (Fig. 1.22). DNA evidence
coupled with carbon dating of skeletal remains suggests
that the first Homo sapiens emerged from Africa about
200,000 years ago. These early humans were remarkably
adept at locomotion as they were quick to cover the planet.
With a small band of modern humans leaving Africa about
100,000 years ago (estimated to be between 50 and 100
individuals [1]), skeletal remains have been dated in the
Middle East at 90,000 years ago, China 67,000, Australia
30,000, and the Americas starting at 11,000 years ago.
Within 1,000 years, our early ancestors spread from the
steppes of Russia to the lower tip of South America.
The skeletal factors that allowed for such efficient
long distance travel included a widening of the dorsal
first metatarsal head (that stabilizes the hallux throughout
the propulsive period), an effective calcaneocuboid joint
locking mechanism (that decreases muscular strain and
prevents buckling of the midfoot after heel lift occurs), a
centralized first metatarsal with a powerful peroneus longus
muscle wrapping around its base (providing improved
medial stability during push-off) and an increased mass
in the distal tibia and fibula that allowed for a larger and
more stable ankle joint complex. Also, in order to provide
variable gearing depending on the terrain and/or the desired
speed, the distal metatarsal heads formed a parabolic curve
that enabled the early Homo sapiens to rapidly change Figure 1.22. The modern human.
19
which the tendons responsible for the storage of energy

(e.g., the Achilles and the plantar fascia) became broader
and stronger so they could both absorb and return energy
during the gait cycle (getting farther on fewer calories is the
ultimate goal of efficient bipedality). Muscle attachment
sites became thicker and left clear marks on our skeletons
(particularly in males). According to Sawyer et al. (15),
these unique skeletal features all serve to increase the
efficiency of our two-legged movements. These authors go
on to explain why the world’s most successful long distance
runners are from sub-Saharan Africa (notably Kenya and
Ethiopia). They suggest that since the original Homo
sapiens were from eastern Africa, they have had the longest
time to allow natural selection to produce a skeletal shape
that could accommodate heat dissipation so essential in a
tropical environment. Just as the early Homo sapiens living
in cold environments for thousands of years eventually
Figure 1.23. Because the second metatarsal is longer developed wide, long torsos with short extremities that
than the remaining metatarsals, it serves as a pivot favored heat retention (e.g., Inuit, Mongolians and even
point allowing the human foot to choose between 2 the early European cave dwelling Cro Magnons), the
different push-off options. When the rearfoot supinates earliest Homo sapiens that remained in sub-Saharan Africa
(A), push-off occurs through the oblique axis which has a eventually developed reduced trunk volume and longer
shorter lever arm to the ankle joint (compare X1 and X2).
extremity lengths that increased the surface area to volume
This lessens strain on the Achilles and is often used when
ratio thereby providing for improved heat dissipation. As is
running uphill. Because of the shorter lever arm, use of the
oblique axis allows for what Bojsen-Moller (44) refers to as consistent with natural selection, the greater the time period,
a low gear push-off. When greater force is needed (e.g., the more pronounced the changes: the Inuit, with only a
sprinting) the peroneus brevis muscle (PB in B) everts the few thousand years in a cold environment do not have the
rearfoot thereby forcing the foot onto the transverse axis same torso extremity length ratios as the Mongolians, who
(B). Because of the longer lever arm, use of the transverse have been exposed to cold climates for tens of thousands of
axis is referred to as a high gear push-off and this axis is years. Sawyer et al. (15) claim that after 200,000 years of
used when faster speeds are required. Notice in figure B adapting to extreme heat, the increased lower limb lengths
that the plantar fascia (pf) and flexor hallucis longus muscle relative to torso volume make sub-Saharan African runners
(FHL) tense to stabilize the medial forefoot against the the most efficient endurance runners on earth. Given the
greater forces associated with use of the transverse axis.
small populations of Kenya and Ethiopia compared to the
The transition from low to high gear push-off results in a
world population, and the number of world records held by
brief period in which body weight is supported solely by the
second metatarsal. Despite its narrow width, the second these two countries in the 10 km and marathon distances,
metatarsal shaft possesses a greater percentage of cortical this theory seems more than plausible.
bone allowing it to tolerate the significant bending forces It took a long time but by standing upright and taking
associated with this brief but frequent transition phase. those first few steps 7 million years ago, Sahelanthropus
tchadensis set into motion a series of events that would
eventually convert a tree-dwelling ape into the most
bending strains. The articular surface areas increased in successful species the world has ever seen. Bipedality freed
size relative to bone length (which allowed for an improved the hands for a variety of actions from tool use to creating
distribution of joint pressure) and the spine developed 3 music and art. It uncoupled respiration and locomotion
curves that provided larger ranges of motion while more (which turned out to be essential for the development of
evenly distributing pressure between the discs and facets. speech) and improved our long distance foraging skills, so
There was a continued progressive increase in vertebral crucial in providing the calorie dense diet our expanding
girth moving from the cervical spine to the lumbar spine brains required. In 100,000 years, Homo sapiens have
that provided protection against the large ground-reactive grown from a small band of adventurous (and probably
forces associated with our upright posture. The sacrum’s 5 hungry) modern humans taking those first few steps out
vertebrae fused and our tail (coccyx) shriveled in size. of Africa, into a world population exceeding 7 billion
Unlike Lucy, the pelvis became wider and the birth and growing rapidly. Although unaware of the causes and
canal size increased in females, allowing for the delivery timing of specific events, Darwin was right: bipedality was
of our big-brained babies. Soft tissue changes occurred in the defining feature that made us human.
20
21. K ivell T, Schmitt D. Independent evolution of knuckle-

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22
Chapter Three
Ideal Motions during the Gait Cycle
The fundamental component of human locomotion 33% of stance phase, beginning the moment the heel leaves
is the gait cycle. One complete gait cycle consists of the the ground and ending when the tips of the phalanges no
anatomical interactions occurring from the moment the longer make ground contact. Although running is also
foot first contacts the ground, until that same foot again divided into the same 3 periods, the increased speed and
makes ground contact with the next step. The human the need for a more forceful propulsive period changes the
gait cycle consists of two phases: stance phase, in which timing of the events, as the contact and midstance periods
the lower extremity is contacting the ground; and swing are slightly shorter (occurring in the initial 0-20% and 20-
phase, in which the lower extremity is swinging through 45% of stance phase, respectively), while the propulsive
the air preparing for the next impact (Fig. 3.1). When a period is extended, occupying the final 55% of stance phase
person is walking, the gait cycle lasts approximately one (2). With more than 5,000 cycles performed daily, the gait
second (1). As a result, stance phase occurs in 0.6 seconds cycle is one of the most repetitive events in our lives.
and swing phase in 0.4 seconds. Because the distal end of The neurological mechanisms necessary to complete
the kinetic chain is fixed by ground-reactive forces during a gait cycle are unusual in that swing phase motions are
stance phase, motions during this portion of the gait cycle reflexive and present at birth (e.g., an unbalanced toddler
are referred to as closed-chain motions. In contrast, swing will immediately swing the lower extremity into a protected
phase motions are referred to as open-chain motions since position), while movements associated with stance phase
the distal end of the kinetic chain is freely mobile. Because represent a learned process (3). Scott (4) supports this
of the complexity of stance phase motions, this portion of statement with the clinical observation that children born
the gait cycle has been subdivided into contact, midstance, without sight make no spontaneous attempts to stand up
and propulsive periods (Fig. 3.2). and walk on their own, and will only do so when physically
When walking, the contact period represents the first guided. With or without sight, once upright and moving
27% of stance phase, beginning at touchdown and ending about, children immediately begin experimenting with a
when the entire forefoot makes ground contact. Midstance wide range of walking and running patterns, subconsciously
occupies 27-67% of stance, representing the period in analyzing the metabolic expense associated with each
which the body’s center of mass is “vaulting” over the variation in gait. This is a time-consuming process and
stance phase foot. The propulsive period occupies the final perfecting the musculoskeletal interactions necessary to
Figure 3.1. Gait cycle of the right leg. Stance phase begins at heel strike (HS) and ends when the great toe leaves the
ground. Swing phase continues until the heel again strikes the ground. The length of stride, which refers to the distance
between successive ipsilateral heel strikes, is approximately 0.8 times a person’s body height and the average cadence
is 115 steps/minute. Because size affects stride length and cadence, there is much individual variation in the gait cycle as
women typically have slightly shorter stride lengths and a more frequent cadence. Children have particularly high cadences
as the average 7 year old takes approximately 143 steps per minute. Because of the prolonged airborne phase, stride lengths
while running significantly increase and it is not uncommon for world-class runners to possess stride lengths exceeding 3.5
meters (11 feet 6 inches), which is more than one meter longer than a comparably sized running quadruped (13).
87
Figure 3.2. The various periods of stance phase. HS, heel strike; FFL, full forefoot load; HL, heel lift; TO, toe off.
become metabolically efficient can take up to a decade to stiff, the body’s center of mass would move through a series
master. Even when considering size differences, the average of abruptly intersecting arcs (Fig. 3.3A) that would greatly
3 year old consumes 33% more oxygen when traveling at increase the metabolic cost of locomotion because muscles
a fixed speed compared to an adult (5). By the age of 6, must tense to accommodate the exaggerated angular
children continue to have significantly higher ratios of displacements. Further strain would be placed on the
energy costs versus work performed (6). Fortunately, by supporting muscles since they would initially absorb, and
age 10, mechanical efficiency has improved and the cost/ then accelerate these forces as the curves reverse direction.
work ratios of 10 year olds and adults are about equal To lessen the metabolic cost of locomotion, each person
(6): after almost a decade of practice, children are finally incorporates a specific series of articular interactions that
efficient at bipedal locomotion. effectively decrease angular displacement of the body’s
In order to create a metabolically efficient gait, center of mass. These actions, or determinants, are listed as
Saunders et al. (7) claim that individuals must learn to follows: pelvic rotation; pelvic tilt; knee flexion/extension
“translate their center of mass through space along a during stance phase; hip-knee-ankle interactions; and
path requiring the least expenditure of energy.” This is lateral pelvic displacement. The following illustrations,
accomplished by modifying joint positions in the lower which were adapted from Saunders et al. (7), demonstrate
extremity and pelvis in such a way that the pathway of the how each determinant affects translation of the center of
center of mass through space is flattened. For example, if mass through space (Figs. 3.3-3.8).
an individual were to walk with knees locked and the pelvis Although the determinants described by Saunders et
Figure 3.3. Determinants of gait: pelvic rotation. Panel A represents a lateral view of the gait cycle with the knees and
hips locked. Notice how the pathway of the center of mass creates an exaggerated sine wave (M1), which is metabolically
expensive because the hip abductors must raise and lower the center of mass through the exaggerated ranges. By
incorporating pelvic rotation (arrows in panel B), the pathway of the center of mass is flattened slightly as rotation of the
pelvis decreases the amount of hip flexion/extension necessary to achieve the same stride length (W). This decreases
vertical drop during double-limb support by approximately 9 mm (the difference between the ground and the center of mass
in X and Y), flattening the pathway for the center of mass (compare M2 and M1).
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Chapter Three Ideal Motions During the Gait Cycle
Figure 3.4. Pelvic tilt. Eccentric contraction of the hip abductors during midstance lowers the pelvis on the side of the swing
leg (arrows in B). This decreases vertical displacement of the center of mass by approximately 3 mm.
Figure 3.5. Knee flexion/extension during stance phase. Part A represents stance phase lower extremity motion without
knee flexion while Part B represents the same leg with knee flexion/extension. Notice that when the lower extremity is
straightened throughout stance phase, the center of mass describes a path along the arc of a circle, with the length of
the lower extremity being the radius. This arc is effectively flattened by knee flexion during early stance phase and knee
extension during late stance phase.
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Figure 3.6. Hip, knee, and ankle interactions. As heel strike occurs, the anterior compartment muscles eccentrically
contract to slowly lower the stance leg to the ground (A and B). This action, coupled with simultaneous knee flexion,
maintains a smoother course for the center of mass during the contact period. Forceful ankle plantarflexion during propulsion
markedly elevates the leg (B-D) and is responsible for the maintenance of an almost straight pathway for the center of mass
during late stance phase (3-4). Flexion of the knee and hip during swing phase (5-7) allows for sufficient ground clearance
despite lowering of the pelvis that is normally occurring on the swing leg side. If the knee and hip were unable to move
through adequate ranges of motion, the individual would most likely compensate by circumducting the swing leg. This action
greatly distorts movement of the center of mass and is metabolically very expensive.
Figure 3.7. Lateral pelvic displacement. To maintain balance during the gait cycle, the weight-bearing leg adducts, and
the swing leg abducts. This allows the center of mass to be displaced laterally over the supporting leg (Panel A, X). If the
lower extremity were perfectly straight (as in panel A), the degree of lateral deviation necessary to maintain balance is
significant. This markedly increases strain on the hip abductors and peroneals when these muscles accelerate the center
of mass medially during late midstance and early propulsion. Fortunately, most people possess a slight degree of genu
valgum (C) that reduces the degree of lateral displacement by allowing for a more approximated base of gait (Y). A mild
genu valgum also allows the tibia to move through the gait cycle in a near vertical position (Panel B).
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Figure 3.8. Final translation of the center of mass during a single stride. Lateral and vertical displacements are
represented by A and B, respectively. Notice that these displacements form sine waves, with the frequency of vertical
displacement being exactly twice that of the lateral displacement. C represents the projection of these displacements
(which have been exaggerated) onto a plane perpendicular to the body’s line of progression. Points 1, 2, and 3 represent
the location of the center of mass during successive single, double and single-limb stance points, respectively. Because
peak vertical displacements are reached slightly before peak lateral displacements, this curve represents a slightly distorted
“lazy 8” (D). At higher speeds of walking, the amplitude of lateral displacement is decreased, and the lateral and vertical
displacements peak at the same time. As a result, the perpendicular displacement of the center of mass more closely
resembles a “U” (E). Note that even at maximal vertical displacement (X), the center of mass never reaches the level it
would assume during static stance (which is represented by 0). When walking, forward acceleration of the center of mass,
at both high and low speeds, is greatest at the low points of vertical displacement (i.e., during double-limb support) and
least at the high points (i.e., during midstance period). Another way of saying this is that kinetic energy is greatest at the low
points whereas potential energy is greatest at the high points. This is comparable to rolling an egg end over end: the egg is
moving rapidly at the low point in the cycle and gradually decelerates, so that it is barely moving by the time it reaches the
high point in the cycle.
al. (7) improve efficiency by flattening the progression of An important consideration is that the determinants
the center of mass through space, it is possible to flatten the described by Saunders et al. (7) must be modified depending
pathway so much that the resultant gait pattern becomes upon the speed of locomotion. For example, at low speeds
metabolically inefficient. To demonstrate this concept on we are most efficient when the lower extremities are stiff
yourself, try walking in a manner similar to the comedian and inflexible but at higher speeds we must increase the
Groucho Marx. Although hyperflexion of the knees and hips degree of knee and hip flexion in order to improve shock
associated with this style of gait will flatten the pathway of absorption. To determine exactly which gait pattern is most
the center of mass, it is metabolically expensive because efficient at a specific speed of locomotion, Srinivasan and
the caloric cost associated with exaggerated knee flexion is Ruina (10) created a computerized mathematical model
high. In fact, McMahon et al. (8) found that walking with to evaluate metabolic efficiency associated with every
a “Groucho gait” resulted in a 50% increase in oxygen possible type of gait (including odd patterns such as the
consumption. Excessive flattening of the pathway of the Groucho gait). As expected, at low speeds of locomotion,
center of mass accomplished by flexing the extremities walking was most efficient with an inverted pendulum
explains why small mammals are so inefficient compared gait while at higher speeds, a “bouncing” or “impulsive
to large mammals; e.g., on a gram per gram basis, a mouse running” style of gait was most efficient (Fig. 3.9). These
consumes 20 times more energy than a pony (9). It also findings correlate with the clinical observation that walking
explains why elephants, despite their enormous size, are feels more comfortable when moving slowly, while running
one of the most metabolically efficient animals on the is more comfortable as speeds increase.
planet: unlike rodents that are forced to hyperflex their Notice in these illustrations that the defining
extremities with each step, elephants keep their extremities difference between walking and running is the center of
locked during stance phase thereby lessening the muscular mass is at a low point during midstance when running,
cost of locomotion. and a high point during midstance when walking. This is
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important because the presence of an airborne phase has

historically been used to differentiate walking from running.
McMahon (11) notes that using an airborne phase to define
running is inadequate since slow running often occurs with
a double-limb support followed by a single-limb stance;
i.e., there is no airborne phase. The author suggests that
a more accurate indicator of the transition from walking
to running occurs when the center of mass switches from
a high to a low point during midstance. McMahon (11)
supports this statement with the observation that Groucho
running lacks an airborne phase and the center of mass is at
its lowest point during midstance.
The most important result of the computerized
model by Srinivasan and Ruina (10) was that walking and
running were only used at the extremes of speed: walking
at low-speed and running at high-speed. For all in-between
speeds, the computer model suggested that people would
choose a hybrid gait referred to as “pendular running” (Fig.
3.10). In this gait pattern, the stride length is significantly
shortened, there is a marked reduction in airtime, and
the lower extremities behave as inverted pendulums for
Figure 3.9. Computerized model simulating gait
brief periods during stance phase. Surprisingly, except
confirms that at slow speeds, the inverted pendulum for occasional references to “Groucho running” and
gait is most efficient, as the lower extremity remains “double-limb support slow running,” options other than
relatively straight, minimizing the metabolic cost of conventional walking and running are rarely discussed in
locomotion. When higher speeds are necessary, the the literature. Because it has only a brief airborne phase
body launches into an impulsive run in which the lower and a shorter stride length, the hybrid pendular running
extremities collapse in a spring-like manner in order to is metabolically more efficient than impulsive running
allow for the storage and return energy necessary for the and may even have been the preferred gait used by Homo
prolonged flight phase (although fast, propelling the body erectus, as Biewener et al. (12) claim that impulsive running
into flight is metabolically very expensive). Notice the length was too expensive to allow for survival of this hominid
of stride is significantly longer with impulsive running.
ancestor. By comparing muscular forces associated with
different speeds of locomotion, these authors determined
the transition from walking to running resulted in a 68%
decrease in the mechanical advantage at the knee, along
with a 5-fold increase in ground-reactive forces. These
combined factors resulted in a 5.2-fold increase in the
quadriceps muscle impulse, which would have made it
difficult for this hominid to gather the calories necessary to
fuel such an inefficient form of transportation. Because of
the inflated metabolic expense associated with conventional
running, Biewener et al. (12) claim that running efficiency
was “unlikely a key selective factor favoring the evolution
of erect bipedalism in humans.” Although this was an
Figure 3.10. Pendular running. This gait pattern extremely detailed study, a flaw with this research is the
represents a hybrid gait containing characteristics of both
authors assumed that these early hominids were forced to
inverted pendulum walking and impulsive running. At slower
speeds of locomotion, pendular running more closely
choose between the highly efficient inverted pendulum
resembles walking in that the lower extremity stiffens and walking and the metabolically expensive impulsive style of
the flight phase is reduced. In contrast, when walking running. It is much more likely that these early hominids,
speed increases, pendular running more closely resembles like most recreational joggers, developed a transitional
impulsive running because the legs become more compliant hybrid gait, which would have been significantly more
and the flight phase is increased. Notice the center of mass efficient and allowed for the improved foraging skills
fluctuates less with pendular running than in either inverted described by Bramble and Lieberman (13).
pendulum walking or impulsive running, creating two small The various types of gait available during
peaks during midstance and swing. locomotion are made apparent by stepping onto a
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motorized treadmill and gradually increasing your speed. transition occurred. The authors relate the sudden increase
At first, inverted pendulum walking is comfortable but as in force to the “improved contractile conditions” associated
you press the acceleration button to increase speed, you with a more midline position of the ankle plantarflexors.
are unable to match the speed of the treadmill so you Interestingly, the EMG output for the ankle plantarflexors
quickly respond by increasing cadence (each person has did not appreciably change despite the significant drop
a preferred stride length so it is usually more comfortable in force output, confirming that electrical activity of a
to increase turnover rate rather than lengthen the stride). muscle has little to do with the force being generated by
This only works for a short time because the metabolic the muscle since it does not take into account the length-
cost of rapidly accelerating and decelerating the lower tension relationship.
extremities is too high, so you respond by increasing your When the highest speeds of locomotion are necessary
stride length. While professional racewalkers are capable (e.g., sprinting), you launch into impulsive running. As the
of greatly increasing stride lengths by hyperextending speed of impulsive running increases, the pathway of the
their knees and exaggerating pelvic and ankle motions center of mass flattens slightly. Although metabolically
(often achieving walking speeds of 6 minutes/mile), the expensive, impulsive running allows you to increase your
average person quickly reaches a length of stride that speed simply by increasing stride length (15). By analyzing
becomes difficult to maintain. At this point, most people all methods of increasing the speed of impulsive running
transition into a hybrid slow run (either a double-limb (e.g., increasing cadence and/or shortening the time the
support slow run or a pendular run). The precise point at swing phase leg is airborne), Weyand et al. (16) conclude
which the transition to slow running occurs varies as each that success with sprinting is determined by the amount of
person has his or her own unique transition speed (which force produced during stance phase: greater force generated
tends to occur somewhere around 2 meters/second). The while making ground contact translates into longer stride
reason each person has a preferred transition speed (PTS) lengths, greater aerial time and faster speeds.
has been the subject of debate as some studies suggest Because the increased aerial phase associated with
that people switch from walking to running to improve impulsive running results in a 5-fold increase in ground-
efficiency, while other studies have shown the switch to reactive force, the body must immediately choose from
running always occurs before a metabolically optimal several different biomechanical options in order to dissipate
walking speed is achieved. these amplified forces. For example, the increased ground-
The controversy regarding the walk-run transition reactive forces can be dampened by making initial ground
was resolved in a clever paper by Neptune and Sasaki (14). contact with the forefoot, exaggerating frontal plane pelvic
By comparing ground-reactive forces beneath the forefoot motions, and/or by hyperflexing the knee and hip. The
with EMG activity of the ankle plantarflexors, these authors exact combination of biomechanical options chosen is
determined that the preferred transition speed occurs highly variable as each person has significant differences
when the degree of ankle dorsiflexion becomes so great in strength, bony architecture, and flexibility. Even prior
that the gastrocnemius and soleus muscles can no longer injury may influence which determinants are incorporated.
generate the force necessary for propulsion because their By experimenting with every biomechanical option, people
length-tension relationship has been compromised; i.e., the select a specific running pattern that is metabolically most
Weber paradox states that when a muscle is fully stretched efficient for them. This explains why runners, unlike
or shortened, contact between the muscle filaments is walkers, present with such a wide range of running styles.
diminished and the muscle is weakened. The presence of It also explains why any attempt to modify a runner’s self-
the Weber paradox is readily apparent to anyone attempting selected stride length will result in a metabolically less
a pull-up: at first, it feels impossible to lift yourself but efficient gait (17). According to Anderson (18), runners
once you move past the first few inches, the pull-up seems are able to critically evaluate all factors associated with
easier until you are almost at the bar, when it again seems “perceived exertion to arrive at a stride length which
difficult. The reason for this is that when a muscle is close minimizes energy cost.”
to a midline position (neither stretched nor shortened), a Despite differences in timing, stride lengths,
greater percentage of contractile filaments are in contact energetic criteria and the degree of joint excursions, the
with one another so greater forces can be generated. articular interactions associated with walking and running
By measuring force output beneath the forefoot, are remarkably similar. The following section reviews
Neptune and Sasaki (14) confirmed that as walking speed in detail the various kinetic/kinematic events occurring
increased, the range of ankle dorsiflexion increased and during the different phases of the gait cycle. When
the Weber paradox made it difficult for the gastrocnemius applicable, differences between walking and running will
and soleus to generate the force necessary to initiate be discussed.
propulsion. The transition to running improved the length-
tension relationship of the ankle plantarflexors and the
force generated by these muscles nearly doubled after the
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Gait Cycle Summary: In young adults, the timing between soleus

Static Stance inhibition and tibialis anterior activation is consistent
The simple task of initiating the gait cycle requires with little variation. This is not the same for healthy older
intricate interactions between the proprioceptive and motor adults, because age-related changes in the motor cortex
systems necessary for fine-tuning postural adjustments, make it difficult to inhibit the gastrocsoleus complex prior
and the motor commands necessary to produce volitional to stimulating tibialis anterior, explaining the frequent
movement. This is complex because the motor circuits prevalence of gait initiation falls in the elderly (130).
governing these two activities function independently, Immediately upon the initiation of the gait cycle,
which is clinically evidenced by the fact that people with the abdominal core muscles tense to lock the torso and
Parkinson’s disease have trouble initiating the gait cycle, pelvis, providing a stable foundation for the hip flexors,
but once started, can walk without difficulty. which are beginning to swing the lower extremity forward
In order to take the first step, the central nervous in preparation for ground contact.
system incorporates a stereotyped pattern of motor
recruitment known as the gait-initiation motor control
program, in which a series of imperceptible postural Stance Phase Motions:
adjustments shift the center of pressure posteriorly, first Contact Period
towards the foot that is about to become the swing leg, and The initial choice to be made during the contact
then back towards the soon-to-be stance leg (Fig. 3.11, A). period is deciding which part of the foot should make
Because the posterior migration of the center of pressure ground contact first. While walking and hybrid running
occurs before any noticeable movement of the center of almost always begin with initial ground contact occurring
mass, the forward component of the ground-reactive force at the back of the heel, runners often vary their initial
increases (arrow X in Fig. 3.11, B). At this time, the soleus/ point of impact, striking the ground at either the rearfoot,
gastrocnemius muscles relax, followed shortly thereafter midfoot, or forefoot. In an analysis of foot strike patterns
by activation of the bilateral tibialis anterior muscles. The in 415 runners participating in an elite-level half marathon,
sudden relaxation of the gastrocnemius/soleus muscles Hasegawa et al. (19) determined that 75% of the runners
initiates a smooth free-fall in the desired direction of travel, made initial ground contact at the rearfoot, 23.7% at the
while the delayed contraction of tibialis anterior pulls midfoot, and only 1.4% made initial ground contact at
the center of mass forward on the stance side and begins the forefoot. The authors went on to evaluate foot strike
dorsiflexing the ankle to improve ground clearance on the patterns of the fastest 50 runners, many of whom were
swing side. Olympians, and noted that the percentage of midfoot strike
patterns present in the fastest runners increased from 23.7
to 36%. The researchers also noted the fastest runners had a
tendency to strike the ground with the foot inverted, which
the authors suggested might somehow improve metabolic
efficiency.
Despite the fact that one third of the world’s fastest
runners make initial ground contact at the midfoot, little is
known about the biomechanics associated with this strike
pattern. In contrast, a significant body of information exists
on the relatively rare forefoot strike pattern; i.e., only 1.4%
of elite runners made initial ground contact at the forefoot
(19). In one of the first studies comparing different strike
patterns, Cavanaugh and LaFortune (20) measured vertical
forces as subjects switched from rearfoot to forefoot strike
patterns and noted the forefoot strike reduced vertical forces
by more than 50%. This large reduction in vertical force
was attributed to the posterior calf musculature absorbing
Figure 3.11. To initiate the gait cycle, the calf and hip impact forces that would otherwise have traveled through
abductor musculature shift the center of pressure the lower extremity.
(COP) laterally and posteriorly, first to the swing foot In a detailed evaluation of the differences between
and then to the stance foot (A). Because the posterior forefoot and rearfoot strike patterns, McClay and Manal
displacement of the COP occurs without perceptible motion (21,22) compared the biomechanics of 10 rearfoot and
of the center of mass (COM), the body tilts slightly forward, 10 forefoot strikers and noted that the forefoot strikers
pivoting about the center of mass (B). See text. Redrawn
made initial ground contact with greater degrees of ankle
from Polcyn et al. (130).
plantarflexion and rearfoot inversion, which resulted in
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increased eversion excursions and eversion velocities pattern was metabolically more efficient for several
during the contact period. Like Cavanaugh and LaFortune reasons: It improved pendular exchange of kinetic and
(20), McClay and Manal (21,22) noted 50% reductions potential energy; decreased mechanical work performed
in vertical loading rates, which they suggest may play a by the limbs; decreased the ground-reactive force moment
role in lessening the potential for tibial stress fracture. at the ankle; and it decreased muscular activity in the
They also suggest that, because knee flexion excursions extensors of the ankle, knee, hip and back. All of these
and velocities were significantly reduced with the forefoot factors greatly influence efficiency and explains why more
strike pattern, rates of knee injuries might also be lessened. than 99% of walkers make initial ground contact with the
However, the greater work performed by the ankle heel. Given the huge advantage associated with heel-first
plantarflexors coupled with the higher ankle dorsiflexion strike while walking, it is likely that the slow pendular
velocities would predispose the Achilles and posterior calf style of running described by Srinivasan and Ruina (10) is
musculature to injury: The forefoot strike pattern reduces also significantly more efficient with a rearfoot strike since
the risk of knee injury while increasing the risk of Achilles it closely resembles walking. This is clinically supported
and forefoot injury. by the fact that although one third of elite runners make
In one of the few studies evaluating the biomechanical ground contact at the midfoot (19), slower marathoners
effects of midfoot strike patterns, Altman and Davis (23) almost always make ground contact with the heel.
used tibial accelerometers to measure vertical loading The preference for rearfoot strike patterns dates
rates as 5 subjects ran with rearfoot, midfoot or forefoot back millions of years, as laser analysis of the 1.5 million-
strike patterns. As expected, the forefoot strike resulted in year-old Homo erectus footprints found in Ileret, Kenya,
significant decreases in vertical loading (at the expense of revealed that our most efficient hominid ancestor made
the posterior calf musculature), while the midfoot strike initial ground contact at the heel (27). The reason for this
pattern produced loading rates about halfway between is simple: 7 million years of evolution has molded the
those occurring in forefoot and rearfoot strike patterns. The calcaneus into a shape that is perfectly suited for absorbing
authors suggest that midfoot strike patterns may provide the forces associated with heel strike. One of the most
“a compromise between these two extremes” and may important factors making the calcaneus effective at stress
reduce the risk of injury. They support this statement with dissipation is its size: the average 100-pound human
the clinical observations that barefoot runners attempt to female has a larger calcaneus than a 350-pound gorilla. The
lessen impact loads by switching to midfoot strike patterns increased size provides more space for pressure distribution
(24), and that barefoot runners anecdotally report fewer through the thin but complex trabecular network.
injuries. Another factor improving its ability to absorb
Despite anecdotal reports, midfoot strike patterns shock is the somewhat incongruous finding that the
have never been proven to lessen the rate of injury (25), calcaneus, despite being exposed to large vertical forces
and may even increase the rate of forefoot injuries because during contact, possesses extremely thin cortical bone and
ground contact forces beneath the forefoot are prolonged sparse trabeculae. This combination creates an essentially
(increasing the risk for sesamoiditis and metatarsalgia). hollow structure in which the calcaneus functions like an
Furthermore, claims that midfoot strike patterns are more overblown cushion at heel strike; i.e., the well-developed
efficient than rearfoot strike patterns are unfounded, vascular supply maintains an elevated intraosseous
since eccentric contraction of the ankle plantarflexors pressure that reinforces the slender cortical bone, allowing
during early stance phase is metabolically expensive. it to bend in and out with the application of ground-reactive
The inability of mid/forefoot strike patterns to improve forces. Like the end plates of the lumbar vertebral bodies
metabolic efficiency was confirmed by Cunningham et (refer back to Fig. 2.91), the walls of the calcaneus bulge
al. (26). By calculating joint torque, mechanical work slightly during heel strike and the trabeculae, with their
performed and muscle activity associated with altering abundant blood supply, quickly repair the microfractures
initial contact points at various speeds of locomotion, associated with repeat impacts. The thin cortical walls are
the authors determined that running with a mid/forefoot helpful when differentially diagnosing a possible calcaneal
contact provided no clear metabolic advantage over a heel- stress fracture: because the cortical walls are so thin, lightly
first strike pattern, because the costs of transport when squeezing the medial and lateral walls elicits discomfort
using the different contact points was about the same. The when a stress fracture is present.
only biomechanical difference was that runners making The final factor making the calcaneus effective at
initial ground contact with the mid/forefoot tended to have stress dissipation is that it is protected by an incredibly
increased stride frequencies and decreased stride lengths. well-designed fat pad. Averaging 18 mm thick in the typical
In contrast to running, Cunningham et al. (26) adult male, the calcaneal fat pad is comprised of spiral
confirmed that walking with a heel-first strike pattern chambers of sealed fat surrounded by whorls of fibroelastic
reduced the metabolic cost of transport by a surprising tissue (28) (Fig. 3.12). These fibroelastic chambers form a
53%. The authors demonstrate that the rearfoot strike honeycomb pattern and are completely isolated as injections
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Figure 3.12. The calcaneal fat pad. Modified from Jahss

et al. (28).
of India ink into the individual chambers confirms there

is no leakage: these compartments behave as closed-cell
chambers that deform and return to their original shape
as they absorb shock (29). Ultrasonography reveals the
fat pad is divided into a deep, thick, highly deformable
macrochamber, and a thin, superficial, nondeformable
microchamber (30). Because it maintains its shape upon
compression, the microchamber functions as a protective
cup that serves to contain the macrochamber layer beneath
the calcaneus, while the macrochamber layer functions
as the major shock absorber, quickly deforming and
rebounding with the application of force (Fig. 3.13).
To prevent slippage and maintain its position Figure 3.13. The normal heel fat pad is composed of a
deep macrochamber and a superficial microchamber.
beneath the calcaneus, the fat pad is anchored by
When exposed to vertical force, the macrochamber
numerous fibrous reinforcements to the skin and heel. compresses significantly (compare A and A1) while the
Snow and Bohne (31) determined the heel pad’s strongest microchamber remains unchanged (compare B and B1).
attachment to the calcaneus occurred through a previously Once vertical forces are removed, the macrochamber
unidentified fibrous band they named the “medial calcaneal springs back to its original shape (C). Drawn from
retinaculum.” This structure is made from large retinacular ultrasonography images in Hsu et al. (30).
fibers that branch into the fibrous stroma of the heel pad,
essentially tethering the heel pad to the calcaneal tuberosity.
The authors claim that this medial retinaculum, present in mammalian heel pads fall to temperatures close to freezing
9 out of 10 heels, plays an important role in “counteracting when standing on snow, providing insulation and reducing
the valgus shear force on the fat pad at the time of heel heat loss from the body to the ground. Bennett and Ker
strike.” (34) evaluated the viscoelastic properties of the cadaveric
During static stance, the heel pad lessens peak contact human heel pads throughout a range of temperatures and
pressure points beneath the heel by distributing pressure noted that even at 0°C, the human fat pad retains almost
evenly over the entire surface of the calcaneus. Because all of its shock absorbing properties. The ability of the heel
plantar pressures are highest in the center of the heel, the pad to function in cold environments may in part be due
chambers in this region are tightly packed, narrower and to the higher percentage of polyunsaturated fatty acids:
positioned in a vertical manner. These densely packed Unlike conventional adipose tissue with a polyunsaturated
vertical chambers more effectively distribute ground- to saturated fat ratio of 2.5:1, the healthy human heel pad
reactive forces away from the center of the calcaneus (32). contains 4.5 times more polyunsaturated fatty acids (35).
Besides distributing pressure during static stance, The increased prevalence of polyunsaturated fat might
the heel pad may also play an important role in reducing improve function because it lessens heel pad viscosity and
heat loss to the environment. Irving (33) notes that is more stable at lower temperatures.
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Using radiographic fluoroscopy with an optical directions, respectively (39). This may not seem like
pressure display, Gefen et al. (36) evaluated the in vivo much but when multiplied by the 5,000 foot strikes taken
biomechanical behavior of the fat pad and determined each day, it translates into nearly 640 tons of pressure the
that when walking, the fat pad compressed very rapidly average person must absorb daily (10). To compound the
to a deformation of about 40%, dissipating 17-19% of problem, these forces travel through the body at speeds
the forces associated with heel strike. The deformation exceeding 200 mph (40).
occurred during the first 150 ms of stance phase and was In order to prepare itself for the sudden application
initially rapid, with a slower period of compression that of ground-reactive force associated with contacting the
continued into early midstance, when an unloading phase ground, the body aligns itself during swing phase so every
was initiated. In a study of the viscoelastic properties of the joint is in an ideal position to dampen these forces. At the
human fat pad, Jorgenson and Bojsen-Moller (37) note that moment heel strike occurs, the spine is in a neutral position;
a healthy pad absorbs shock 2.1 times better than sorbothane the hip is flexed 30°; the knee is almost fully extended;
(considered one of the best synthetic shock absorbers) and the ankle is slightly dorsiflexed; the subtalar joint is
possesses an open plexus of veins that allows it to enhance slightly supinated; and the midtarsal joint is fully pronated
the countergravitational return of blood. The venous plexus about its oblique axis and supinated (inverted) about its
enhances shock absorption as the contained blood acts as longitudinal axis (Fig. 3.14). The muscles stabilizing
a cushion. these joints are in their midline positions (maximizing
Although effective at dampening forces associated their length/tension relationships) and many of them
with static stance and walking, the 3- to 5-fold increase are pretensed in anticipation of impact forces. In fact,
in vertical forces associated with heel strike while running switching from soft to hard surfaces produces immediate
produces a jarring impact capable of damaging the walls of increases in anticipatory muscle activity prior to heel strike
the heel pad chambers. In an in vivo cineradiographic study as preactivation improves the ability to absorb shock (41).
of the heel pad in barefoot and shod running, De Clerq Animal studies confirm that variation in surface hardness
et al. (38) demonstrate that barefoot running produces a is accommodated with altered muscle function and joint
60% deformation of the heel pad, compared to the 35% excursions within a single stride length (42).
reduction when running shod. This contrasts with the 40% In an interesting study of ground-reactive forces in
deformation associated with walking barefoot (36). Over horses, Wilson et al. (43) determined that when a galloping
time, the repeated application of 60% deformation could horse’s leg strikes the ground, the impact causes the horse’s
cause the heel pad to bottom out, limiting its ability to leg to vibrate at a rate of 30-40 Hz, and these potentially
distribute pressure and absorb shock. To avoid damaging dangerous vibrations have to be dampened in order to
the fat pad when running, barefoot runners intuitively avoid fatigue failure in tendons and bones. The authors
switch to a midfoot strike pattern (24). This strike pattern demonstrate that these impact vibrations are dampened
protects the heel pad from the exaggerated vertical forces by the digital flexor muscles, which had been previously
as the initial point of contact is displaced forward, away believed to be vestigial in horses, since its muscle fibers are
from the heel pad, allowing the posterior calf musculature too short to produce motion. Although useless for creating
to absorb the increased ground-reactive forces. joint movement, the digital flexor muscles are ideal for
The ability to dampen impact is important since dampening bony vibrations, as their angled origins allow
typical ground-reactive forces when walking average for lateral force transmission of bony vibrations through
110%, 15%, and 10% in the vertical, forward, and medial their “complex 3-dimensional architecture.”
Figure 3.14. Ideal joint positions present at heel strike. OMJA, oblique midtarsal joint axis; LMJA, longitudinal midtarsal
joint axis.
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Because impact vibrations occur in the first 100 ms Plantarflexion of the ankle also serves to decelerate
following ground contact, reflex muscular activation is too the braking phase associated with heel strike; i.e., frictional
slow to provide protection and the vibration dampening forces immediately pin the heel to the ground and the
muscles, to be effective, must be preactivated prior to body’s forward progression comes to an abrupt halt. To
ground contact. In humans, barefoot running has been demonstrate the braking phase on yourself, walk across a
shown to produce similar vibrations as accelerometer and room while holding a full bowl of soup: Every time heel
kinematic data recorded from human tibiae during barefoot strike occurs, the soup has a tendency to spill forward as
running revealed horizontal oscillations between 40-50 progression of the center of mass is temporarily halted by
Hz, which continued for 100 ms following impact (44). frictional forces associated with heel strike. Pandy and
According to Wakeling and Nigg (45), tension created in Berme (48) evaluated the various determinants of gait and
antagonistic muscles preceding ground contact serves to concluded that transverse plane pelvic motion plays the
stiffen the leg in anticipation of these bony oscillations. greatest role in limiting the magnitude of the braking phase
This preactivation may be used to “increase the resonant in that posterior rotation of the pelvis during initial ground
frequencies of the soft tissues of the leg and optimally tune contact smoothly absorbs the temporary reduction in speed
the system to minimize vibrations” (45). In a separate in associated with braking.
vivo study of human walking, Wakeling et al. (41) measured When initial ground contact is made with heel, the
force transmission and muscle activity associated with anterior compartment muscles also play a role in reducing
landing on hard and soft materials and determined the braking because ankle plantarflexion allows the ankle
lateral head of the gastrocnemius and the bicep femoris to continue to roll forward, providing a more gradual
muscle were the major contributors to dampening lower deceleration of the lower extremity (refer back to figure
extremity vibrations associated with heel strike. 3.6). Because midfoot and forefoot strike patterns negate
Following the initial impact, a combination of the ankle’s ability to accommodate the braking phase with
ground-reactive forces (which are typically applied to a forward role of the ankle, the body is forced to rely more
the posterolateral heel) and inertial forces (the pelvis and heavily on transverse plane pelvic motions. The negative
lower extremity continue their internal rotation, which influence of forefoot strike patterns on braking is made
began during early swing phase) cause the ankle to obvious by running downhill while alternating between
plantarflex and the subtalar joint to pronate. Plantarflexion rearfoot and forefoot strike patterns: the moment ground
of the ankle is resisted by the eccentric contraction of the contact is made with the forefoot you feel the lower
anterior compartment musculature. These muscles play an extremity come to an abrupt halt, while a rearfoot strike
important role in absorbing shock as they smoothly lower pattern produces a smoother braking phase as the ankle
the forefoot to the ground, thereby reducing trauma to the rolls downward and forward.
plantar soft tissues. Radin and Paul (46) state that joint In addition to sagittal plane motions at the ankle,
motion controlled by muscles lengthening under tension frontal plane motion of the subtalar joint also plays a role
is the primary kinematic process responsible for shock in shock absorption when tibialis posterior eccentrically
absorption. Tibialis anterior is particularly well-suited contracts to resist calcaneal eversion. This muscle is well-
for decelerating ankle plantarflexion since its muscle designed for shock absorption, because it has the longest
architecture is arranged in such a way that it is almost lever arm for controlling subtalar pronation and prior to
impossible to damage with repeated eccentric contractions inserting beneath the arch, its tendon rotates approximately
(47). The ankle continues to plantarflex during the first 70% 45° in order to store and return energy (49,50) (Fig.
of the contact period, reaching a maximally plantarflexed 3.16). In many ways, tibialis posterior is the frontal plane
position of 10° (Fig. 3.15). At that time, ground-reactive equivalent of the Achilles tendon. The ability of tibialis
forces beneath the forefoot cause the ankle to dorsiflex posterior to absorb shock is enhanced when heel strike
slightly (although the ankle is still plantarflexed 5° by the occurs with the rearfoot inverted, since this allows the
end of the contact period). subtalar joint to move through a larger range of motion
Figure 3.15. Ankle plantarflexion during

early and midcontact period is resisted
by eccentric contraction of the anterior
compartment muscles (arrow in A). Roughly
40% of the way through contact period (B), the
fifth metatarsal head strikes the ground. The
forefoot is then smoothly loaded from lateral to
medial with the entire forefoot making ground
contact approximately 70% of the way through
the contact period (C).
98
with tibialis posterior eccentrically contracting to absorb

shock. Perhaps this explains why runners who make initial
ground contact with the rearfoot inverted are less prone to
injury (132).
Throughout the contact period, the talus continues
to slide medially down the calcaneus, adducting and
plantarflexing over the calcaneal facets. Because no
muscles attach to the talus, this motion is controlled by
bony and ligamentous restraining mechanisms (particularly
the anterior talofibular and the sinus tarsi ligaments). In
addition to improving surface accommodation by creating
a parallelism of the shared midtarsal joint axes (Fig. 3.17),
plantarflexion of the talus also produces a cushioning effect
during heel strike by gradually lowering the ankle mortise
towards the ground (Fig. 3.18).
While motions in the subtalar joint are frequently
likened to a mitered hinge in which frontal plane motion
of the rearfoot is converted into transverse plane motion
of the shank, recent research reveals that this model is
inaccurate because only 66% of frontal plane rearfoot
motion is converted into tibial rotation (52). The clinical
implication of this is that the subtalar joint plays a more
important role in dissipating torsion between the rearfoot
and leg than previously believed.
Even though some of its motion is dissipated,
pronation of the subtalar joint continues to convert frontal
plane motion into tibial rotation by forcing the talus to
adduct, which in turn causes the tibia to internally rotate. Figure 3.16. Relative lever arms to the subtalar joint
The resultant internal tibial rotation causes the lateral axis (STJA). Cadaveric analysis of lever arm lengths by
tibial plateau to glide anteriorly beneath the lateral femoral Hinterman et al. (49) confirms that tibialis posterior (TP) has
condyle, which unlocks the ligamentous restraining the longest lever arm for controlling subtalar motion (X). By
giving tibialis posterior’s lever arm a reference value of 1.00,
mechanisms and allows the knee to flex (i.e., the knee is
these authors calculated comparative lever arm lengths
not a ginglymus joint as the tibia must internally rotate for for the remaining muscles responsible for stabilizing the
the knee to flex properly). subtalar joint, which are listed as follows: flexor digitorum
Knee flexion resisted by eccentric contraction of longus (FDL) .75, flexor hallucis longus (FHL) .62, tibialis
the quadriceps muscle is by far the body’s most effective anterior (TA) .59, soleus .24, extension digitorum longus
shock absorbing system. Because ground-reactive forces (EDL) -.26, peroneus longus (PL) -.82, peroneus brevis
associated with walking are comparatively low, the knee (PB) -.85. AA refers to the ankle axis of motion while positive
flexes through a range of only 20° to 25°. In many situations, and negative numbers are used to describe muscles that
individuals move through the contact and midstance periods either supinate or pronate the subtalar joint, respectively.
Figure 3.17. Anterior view of the right talus and calcaneus. Note the parallelism of the talonavicular (TN) and
calcaneocuboid (CC) axes as the subtalar joint pronates.
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Figure 3.18. Anterior view of the right talus and calcaneus. Notice the talus is lowering as the subtalar joint pronates
(compare A, B, and C).
with their knees fully extended, producing a walking pattern during midstance when the knee was flexed approximately
that more closely resembles the classic inverted pendulum 40°), there was a consistent and unexpected 10 mm posterior
gait. While a stiff knee gait is metabolically efficient shifting of the tibiofemoral joint’s axis of motion, which
because it lessens strain on the quadriceps, it significantly improved the mechanical efficiency of the quadriceps
increases stress on the gluteus medius muscle, which muscle by temporarily increasing the length of its lever
eccentrically fires to lower the contralateral pelvis. The arm (Fig. 3.20). The authors theorized the abrupt shifting of
gluteus medius muscle, however, has a limited ability to the axis allowed the bony architecture of the tibiofemoral
absorb impact force and as shock absorption requirements joint to automatically compensate for the temporary loss
increase with the transition to running, the degree of knee in force that occurs when the quadriceps muscle switches
flexion increases in a linear manner; i.e., slower speeds of from eccentric to concentric contraction (i.e., the transition
running have lower knee flexion angles while faster speeds produces a temporary weakening of the muscle secondary
have proportionately higher degrees of knee flexion. As to the “intrinsic force-velocity relationship of muscle
knee flexion increases, the patellar tendon drags the patella strength”). The unexpected posterior shifting of the axis is
downward over the femoral condyles and patellofemoral extremely useful because it improves metabolic efficiency
contact points increase, providing improved distribution and provides significant stability against knee buckling by
of pressure (refer back to figure 2.48). Fortunately, the briefly increasing the lever arm afforded the quadriceps
patella is well-equipped to handle these forces because it muscle at a time when the muscle is temporarily weakened
possesses the thickest cartilage in the body and is stabilized as it transitions from eccentric to concentric contraction.
by the powerful retinacular ligaments.
To evaluate the 3-dimensional motions in the
knee associated with walking, Komistek et al. (53) used
fluoroscopy and computerized tomography to measure in
vivo tibiofemoral motion as subjects walked in a specially
designed apparatus. They confirmed the tibia internally
rotates about a vertical axis projecting through the medial
femoral condyle with the lateral tibial plateau moving 8.4
mm anteriorly while the medial joint remained stationary
(Fig. 3.19). This refutes prior research suggesting rotation
occurred about an axis though the center of the cruciate
ligaments and explains why osteochondral fragments
frequently form at the medial femoral condyle. In an in
vivo analysis of sagittal plane tibiofemoral motion, van
den Bogert et al. (54) plotted the shifting location of the
tibiofemoral axis as subjects ran through a functional MRI
machine and determined the axis was frequently found
close to the intersection of the cruciate ligaments and, more Figure 3.19. Vertical axis of the right knee joint (A).
importantly, it moved posteriorly as the knee flexed. They The medial tibiofemoral joint behaves as a ball and socket,
also made the interesting observation that when the knee while the lateral side of the joint moves with a gliding motion
transitioned from flexion into extension (which occurred as the tibia moves anteriorly relative to the femoral condyle.
100
Figure 3.20. Posterior shifting of the sagittal plane

axis as the knee transitions from flexion to extension.
While knee flexion resisted by the eccentric

contraction of the quadriceps muscle is the body’s most
important shock absorber, the hip, with its powerful
Figure 3.21. The upper fibers of gluteus maximus
muscular support and significant surface area, also
possess a significant lever arm for controlling frontal
contributes appreciably to the dissipation of ground- plane motion (A), while the lower fibers effectively
reactive forces. At heel strike when walking, the hip is control sagittal (B) and transverse plane motions (C).
flexed 30° allowing for greater distribution of pressure as
contact areas in the hip increase when the hip is flexed.
As mentioned, because the forces associated with walking
are relatively small, the gluteus medius muscle effectively
dampens these forces by eccentrically contracting to lower
the contralateral pelvis.
As speeds of locomotion increase, gluteus medius
is unable to resist the exaggerated forces associated with
impulsive running and the gluteus maximus muscle
vigorously tenses to stabilize the hip. Because it possesses
significant lever arms for controlling transverse, sagittal,
and frontal plane motion (Fig. 3.21), the gluteus maximus
muscle very effectively dampens triplanar forces at the hip.
In addition, the piriformis and obturator internus muscles
play an important role during early stance by decelerating
internal rotation of the femur and protecting the femoral
neck from the bending forces associated with single-limb
stance (Fig. 3.22). Because of its fibrous slips traversing
over the sacrum (refer back to figure 2.81), the piriformis
is also an important stabilizer of the sacroiliac joint; i.e.,
internal rotation of the femur during the contact period
creates a tensile strain in this muscle that is transferred
directly into the soft tissues covering the posterior
sacroiliac joint. Figure 3.22. During single-leg stance, body weight (A)
Perhaps the most interesting series of biomechanical creates tensile and compressive strains on the femoral
events necessary for stability occur at the pelvis. Following neck (small black arrows), which are resisted by the
heel strike, deceleration of the lower extremity associated compressive force created by the piriformis, obturator
with the braking phase of ground contact creates internus and lower gluteus medius muscles (white
arrows).
significant pressure in the femoroacetabular joint as the
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torso continues to move downward and forward, creating

a pivot at the hip (particularly when the knee is straight).
During normal walking, these forces are directed into a
relatively small area in the anterosuperior quadrant of
the acetabulum (55). Even though hip joint forces vary
considerably while walking (with ranges from 200 to 2200
N), the gluteal muscles function to distribute pressure over
a larger surface area while maintaining femoroacetabular
joint forces at a constant level.
Because the weight of the torso is applied
posteromedially to the acetabulum, the ipsilateral ilium
tilts upward and extends while the sacrum nutates forward
(Fig. 3.23). The innominate easily manages the transfer of
forces associated with ground contact, as the cortical shell
of the pelvic bone is thickest along a line connecting the
acetabulum and sacroiliac joint. Using data obtained from
a telemetrized hip prosthesis, Dalstra and Huiskes (56)
created a detailed in vivo model in which they evaluated
external loading of the hip, including the effect of 22
muscles on hip joint forces during normal walking. These
authors confirm that hip joint forces increase rapidly during
the contact period and are initially resisted by both gluteus
maximus and gluteus medius. By midstance, gluteus
medius supports the vast majority of force at the hip. The
authors created a computer image to demonstrate stress Figure 3.23. Because body weight (W) is applied
intensity distribution in the cortical shell of the pelvic bone posteromedially to the point of support in the
acetabulum (A), the sacrum is forced to nutate forward
during the different phases of gait (Fig. 3.24). Notice that
while the ilium extends (white and black arrows,
the preponderance of load transfer occurs from the superior
respectively).
acetabulum, through the posterior iliac pillars, into the
interosseous region of the sacroiliac joint.
Figure 3.24. Stress distribution

through the cortical shell of the left
innominate during stance phase
of gait. The dark shading represents
increased areas of stress distribution.
Redrawn from Dalstra and Huiskes
(56).
102
As noted by Vleeming et al. (125), a self-locking Fortunately, tension associated with eccentric
mechanism occurs to protect the sacroiliac joint when contraction of the bicep femoris muscle is more than
pretension in the bicep femoris, which is activated during enough to stabilize the sacrotuberous ligament because the
late swing phase to decelerate forward motion of the leg, distal aspect of this muscle has a lower point of attachment
increases tension in the spiral fibers of the sacrotuberous compared with the other hamstrings. This lower attachment
ligament. This increased tension (most of which is stored allows forward motion of the swing phase leg to produce
in the form of elastic recoil) serves to decelerate forward greater tensile strain in the long head of the bicep femoris
nutation of the sacrum, when forces are transferred (see Fig. 3.49 at the end of this chapter). Because the
through the sacrotuberous ligament into the deep fibers late swing/early contact period tension in bicep femoris
of the multifidi muscles, crossing the sacrum into the increases as ground-reactive forces increase, this muscle is
contralateral erector spinae and latissimus dorsi muscles able to provide variable stability depending upon the degree
(Fig. 3.25). While Vleeming et al. (125) claim the self- of impact force. As previously mentioned, Wakeling et al.
locking mechanism is enhanced by downward motion of (41) demonstrate that when forces present at heel strike
the fibula occurring during heel strike (Fig. 3.26), recent in
vivo research reveals there is negligible motion of the fibula
during early stance and the concept of a muscle-tendon-
fascial sling being pulled by a dropping fibula needs to be
reevaluated (57,58).
Figure 3.26. According to Vleeming et al. (125),

tension created in the tibialis anterior muscle during
the contact period is transferred into the fascia of the
peroneus longus muscle (dotted lines near A), where it
is then transferred through connecting fascia into the
bicep femoris tendon (B). Vleeming et al. (125) claim the
load transfer into the bicep femoris muscle is enhanced
by downward motion of the fibula, which early research
Figure 3.25. Tension created in the long head of the by Weinert et al. (126) suggested occurred during early
bicep femoris muscle passes through the spiral fibers stance phase. Unfortunately, the belief that the fibula drops
of the sacrotuberous ligament (A) into the deep fibers downward during the contact period has been refuted by
of the ipsilateral multifidus (M). These forces are then several high-quality 3-dimensional in vivo studies (57,58).
transferred into the contralateral erector spinae (ES) and As a result, the concept that fibular motion somehow
into the latissimus dorsi muscle (L). stabilizes the sacroiliac joint is most likely invalid.
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increase (e.g., landing on concrete), the bicep femoris and present in 60% of 50 year olds and 100% of 60 year olds,
lateral gastrocnemius muscles pretense during late swing would completely block nutation of the sacrum, and once
phase in anticipation of these forces. This anticipatory formed, would force vertical forces to transfer directly
pretension in the bicep femoris enhances sacroiliac into the lumbar spine. Of note, even without sacroiliac
stability as the muscle tenses more vigorously when involvement, the sacrum, by itself, functions as a shock
ground-reactive forces increase, providing the sacroiliac absorber because its thin cortical shell allows it to bulge in
with variable degrees of reinforcement depending upon the and out with the application of vertical force in a manner
amplitude of the impact force. similar to the calcaneus following heel strike.
Keep in mind that despite the protection provided by Forces not dampened at the pelvis travel into the
the bicep femoris, the sacroiliac joint remains a relatively spine (Fig. 3.27). In the transverse plane, the lumbar spine
unimportant shock absorber because it possesses such a rotates approximately 3° in each direction and these forces
limited range of motion that its contribution to dampening are dampened by the spinal rotators. In the frontal plane,
impact force is at best limited, especially in older adults. the contralateral pelvis drops approximately 4° while the
Furthermore, the finger-sized bony bridge that traverses spine laterally flexes less than 1° in each direction. Lateral
the interosseous section of the sacroiliac joint, which is flexion of the spine is resisted by the contralateral internal/
external obliques and by the quadratus lumborum muscle.
Because of its 110 mm lever arm to the spine, the external
oblique muscle is particularly well-suited for minimizing
frontal plane displacement of the pelvis. While early
research suggested that contralateral lateral flexion of
the spine following heel strike produced ipsilateral spinal
rotation (i.e., Fryette’e law), recent research confirms
that the coupling of spinal lateral flexion and rotation is
extremely variable and there are no preset rules defining
the coupling of lateral flexion and rotation.
Sagittal plane spinal motions also contribute to
shock absorption. Following heel strike, the braking
action associated with initial ground contact causes the
torso to displace forward rapidly. The powerful iliolumbar
ligament plays an invaluable role at this time by reducing
forward glide of the fourth and fifth lumbar vertebrae on
the sacrum, significantly reducing the risk of injury to
these very mobile vertebrae (Fig. 3.28). Flexion of the
lumbar spine following heel strike is resisted by eccentric
contraction of the erector spinae muscles. Because they
Figure 3.27. Spinal motion. As the left foot strikes

the ground, the lumbar spine is rotated 3° and laterally
flexed 1° to the right (A and B, respectively). Immediately
following heel strike, the contralateral pelvis drops 4°
(C) and the lumbar vertebrae flex forward to assist with
shock absorption (60). Because the arms are swinging in
the opposite direction of the pelvis (D and E), the upper
thoracic spine rotates in the opposite direction of the lumbar
spine. The transition from right rotation of the lumbar spine Figure 3.28. The posterior (P) and superior (S) bands
to left rotation of the upper thoracic spine typically occurs of the iliolumbar ligament protect the fourth and
at the eighth thoracic vertebra. Shortly after heel strike, the fifth lumbar vertebrae from the excessive anterior
lumbar spine reverses directions in all planes in order to translation associated with the braking phase following
dampen movement of the pelvis. heel strike (arrow).
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possess 4 inch lever arms for controlling spinal flexion efficient runners made ground contact with the spine
and attach to the lower 6 ribs, the longissimus thoracis almost vertical. Perhaps the slightly flexed spine lessens
pars lumborum and iliocostalis lumborum pars lumborum angular kinematics following ground contact thereby
very effectively decelerate forward motion of not just the reducing the metabolic cost of locomotion. The end result
lumbar spine, but the entire torso. of the various spinal movements present during the contact
Callaghan et al. (59) evaluated spinal motions period is that the head remains relatively stationary despite
at different speeds of walking and noted that when significant spinal oscillations. In an interesting evaluation
the speed of walking increased, the degree of forward of head motions associated with walking on varied terrain,
flexion present during the contact period also increased, Menz et al. (64) demonstrate that although accelerations
while the ranges of rotation and lateral flexion remained of the pelvis increase when walking on irregular surfaces,
relatively unchanged (emphasizing the role of spinal head accelerations remain unchanged. The authors state
flexion in shock absorption). McGill (60) notes that arm that “one of the primary objectives of the postural control
motions associated with fast walking may serve to lessen system when walking on irregular surfaces is head control,
strain associated with the amplified spinal motions, since and that subjects adapt their stepping pattern on irregular
swinging of the arms was shown to reduce spinal torque, surfaces to ensure that the head remains stable.” They
muscle activity, and spinal loading by up to 10%. McGill suggest elderly people might be more prone to falling
(60) confirms that slow walking is associated with reduced because of an inability to maintain smooth head motions
spinal movement, resulting in a prolonged static load of the while walking.
lumbar ligaments and muscles. Apparently, swinging the
arms while fast walking “facilitates efficient storage and Midstance Period
recovery of elastic energy,” lessening the need for muscle Midstance period begins at full forefoot load and
stabilization with each step. This explains why Nutter ends at heel lift. When walking, it is the longest period,
(61) recommends incorporating fast walking into exercise occupying 40% of stance phase and lasting approximately
protocols prescribed to manage various low back disorders. 0.24 seconds (3). While running, midstance is significantly
It also explains why slow walking (such as walking in a shorter, occupying only 25% of the gait cycle. Because the
mall) is uncomfortable to individuals suffering from low energetic criteria during walking and running are reversed,
back pain. Despite significant individual variation in upper the lower extremity joint interactions are significantly
extremity motions during the gait cycle (some individuals different. During running, the knee and hip continue to
move their wrists asymmetrically across the body while flex throughout midstance, bringing the center of mass
others symmetrically move their hands up and down in a to a low point by the middle of stance phase. Conversely,
rhythmic manner), arm motions play an important role in while walking, the hip and knee remain relatively straight,
reducing spinal torque immediately following heel strike. returning the center of mass to its highest position by the
In addition to absorbing shock through muscular middle of the midstance period.
activity, the vertebral bodies, like the calcanei and sacrum, To get the center of mass to its high point while
possess extremely thin cortical bone (i.e. less than 0.6 mm walking, the lower extremity remains in a nearly
thick) that bulge in and out with the application of vertical straightened position while the entire lower extremity
forces (refer back to figure 2.91). Reinforced by well- pivots up and over the talus. The talus is well-designed to
nourished trabeculae, the bulging end plates are capable function as a pivot point during midstance, since more than
of rapid self-repair and play an important role in shock 70% of its body is covered with cartilage. This interesting
absorption. According to McGill (60), the intervertebral bone essentially functions as an almost frictionless ball
discs play only a limited role in dampening vertical forces as bearing while the stiffened lower extremity pivots over it
they function to distribute stress evenly over the end plates (Fig. 3.29). The stiffening of the lower extremity necessary
(improving the ability of the end plates to absorb shock) to propel the center of mass upwardly during midstance
and separate the vertebrae, allowing for increased ranges is accomplished primarily by maintaining the knee in a
of vertebral motion. Adams, in a personal communication fully extended position: During the contact period while
with McGill (60), suggests the elevated hydrostatic disc walking, the knee flexes through the relatively small range
pressures associated with upright postures serves to prevent of 10° to 20°. This essentially allows the lower extremity
the ingrowth of nerves, lessening the potential for chronic to behave as a pole vault, gradually elevating the center
spinal pain. of mass until it reaches its high point during the middle of
Although some research suggests that a vertical midstance.
spine at heel strike is most efficient because it requires During the contact period when walking and the
less effort to maintain postural equilibrium (62), Wlliam midstance period when running, tension in the iliotibial band
and Cavanagh (63) evaluated efficiency in runners and decreases the metabolic costs of locomotion by assisting
determined the most economical runners made ground gluteus medius in preventing the contralateral pelvis from
contact with the spine flexed forward 5.9°, while the least dropping. Throughout midstance, tension in the iliotibial
105
Contrary to popular belief, the iliotibial band does

not “roll over” the lateral femoral epicondyle, compressing
a bursa as it shifts back and forth over this bony
prominence. Detailed cadaveric analysis coupled with
MRI evaluations of individuals with and without iliotibial
band friction syndrome confirms that the band never snaps
over the epicondyle, because the distal aspect of the band
is anchored too firmly to the femur by strong fibrous bands.
As demonstrated by Fairclough et al. (65), the progressive
increase in the degree of knee flexion during early stance
phase shifts tension from the anterior to the posterior fibers
of the iliotibial band depending on the degree of knee
flexion, creating the illusion of movement (Fig. 3.30). In
no cadaver, volunteer, or patient was a bursa ever found, as
3.29. Because 70% of the talus is covered with cartilage the subsurface of the iliotibial band was protected by a deep
(shaded areas), it functions like a frictionless ball layer of fat. Because tension was greatest in the anterior
bearing, allowing the lower leg to glide over its superior fibers of the iliotibial band at low angles of knee flexion
surface (arrow). and in the posterior band at higher angles, it is possible the
anterior portion provides stability when walking (through
band is enhanced by expansion of the contracting vastus tension in the tensor fasciae latae) while the posterior fibers
lateralis muscle, which displaces the center of the iliotibial provide greater support when running as the ranges of knee
band laterally creating a bowstring effect that increases flexion increase and the gluteus maximus muscle tenses
tension in this important structure. Because the iliotibial more vigorously.
band is connected through the lateral intermuscular septum Throughout the midstance period while walking, the
to the entire posterior femur, it acts as a powerful brace that knee gradually straightens so it is fully extended by the
lessens bending strain on the femur during early and mid time heel lift occurs. In contrast, knee flexion associated
stance phase (65). with running continues until the middle of the midstance
Figure 3.30. The iliotibial band (ITB). When the knee is flexed slightly (A), the tensor fasciae latae muscle (TFL) pulls with
more force than the gluteus maximus muscle (G Max) causing the anterior aspect of the ITB to become more prominent
(compare B and C). As the degree of knee flexion increases (D), greater tension is created in the gluteus maximus muscle and
the posterior aspect of the iliotibial band becomes more prominent (E). The shifting of tension from the anterior to the posterior
fibers of the ITB (F) creates the illusion that the band is displacing forward and backward. This can be demonstrated on yourself
by placing your index and middle fingers on the anterior and posterior aspects of the iliotibial band as you flex your knee
through a 40° range of motion. Notice that when the leg is straight the anterior aspect of the band is more prominent and tension
gradually transitions to the posterior band when the knee flexes past 30°. Drawn from photographs in Fairclough et al. (65).
106
period, reaching a range of more than 40°. The increased

range of knee flexion allows the center of mass to drop
significantly while running, reaching a low point during
the middle of midstance. At this time, the center of mass
reverses direction and the body springs up and forward.
Even though knee and hip kinematics differ
significantly between walking and running, foot motions
are very similar in that the midstance period represents
a time for continued surface accommodation and, more
importantly, energy storage. During the majority of the
midstance period, the subtalar joint continues to pronate
while the talus drops inferomedially. This motion
continues until shortly before heel lift, when subtalar
joint pronation is stopped by, in order of importance,
the congenital placement of the axes of the subtalar and
midtarsal joints, the geometry of their articulating surfaces,
and by the connecting ligaments (66). Throughout its range
of motion, the talus is sliding medially down the calcaneal
facets, pivoting about the posterior band of the interosseus
talocalcaneal ligament; coming to a halt when the talar head
drops into the superomedial calcaneonavicular ligament
and the inferior calcaneonavicular ligament (Fig. 3.31).
To be efficient during this process, the body must Figure 3.31. Superior view of the right foot with the
absorb as much force as it can during early stance so it talus removed. During early stance phase, the talus slides
can return it during the latter half of stance phase. Cavagna medially down the calcaneal facets (shaded areas), pivoting
about the posterior band of the interosseus talocalcaneal
et al. (67) demonstrate that the metabolic cost of running
ligament (A). Motion of the talus continues until the
would be 30-40% higher were it not for the storage and talar head rests in the superomedial calcaneonavicular
eventual return of energy. Energy return is accomplished ligament (B) and the inferior calcaneonavicular ligament
primarily by structural differences in muscle architecture (C). As suggested by Davis et al. (127), because the
between proximal and distal muscles of the lower calcaneonavicular ligament contains very few elastic fibers,
extremity. As demonstrated by Biewener and Daley (68), it should be renamed the “sling ligament,” rather than the
the proximal muscles of the hip and knee tend to have “spring ligament,” since it functions like a sling to stabilize
long parallel fascicles with little tendon elasticity, while the talar head. Notice the tendons of tibialis posterior (TP)
the distal muscles of the foot and leg tend to have short, and flexor digitorum longus (FDL) pass just medial to the
pennate fascicles with long, flexible tendons. head and neck of the talus.
The short fibers present in the distal muscles
allows them to generate force extremely economically by excursions when they run, there is little change in muscle
contracting either isometrically or with low shortening fiber length, since the tendons store and return energy with
velocities: energy is stored in their stretched tendons with no help from the muscles (which saves precious calories).
little work being done by their short pennate fibers. This In fact, certain tendons are so efficient they are able to
contrasts with the proximal muscles, which are recruited return 93% of the work performed when stretching them
in complex patterns, producing force by altering the length (71). Because 7% of the work done while stretching is
of their muscle fibers. This finding is consistent with prior dissipated as heat, repeat use over time causes the tendon’s
research by Ker et al. (69), who demonstrate that energy- temperature to rise gradually, potentially producing thermal
saving tendons have small cross-sectional areas that allow injury. If the tendon was less resilient (which is consistent
them to stretch through large ranges, storing and returning with prior models of muscle function in which the muscles
energy like rubber bands. Although counterintuitive, muscle performed the work and the tendons passively transmitted
fiber lengths present during midstance remain relatively force), a greater percentage of the stretch force would
unchanged while their corresponding tendons stretch and be converted into heat, making prolonged locomotion
rebound back through significant ranges, comparable to impossible since the tendon would fail when exposed to
the spring on a pogo-stick. This relationship allows tendon elevated temperatures. A study of juvenile pigs confirms
elasticity to perform most of the work while muscle action that elasticity is reduced in immature tendons (possibly
occurs isometrically. In detailed biomechanical studies of explaining the metabolic inefficiency present in children),
turkeys and kangaroos, Alexander (70) confirms that even and the capacity of tendons to store and return energy
though the joints of these animals move through large lessens with age (i.e., running economy is significantly
107
reduced in older adults because the muscular system is too small a range of motion to allow it to play a significant
unable to compensate for the diminished tendon elasticity) role in dampening ground-reactive forces. By evaluating
(72). the effect of physiological loading on cadaveric joints of
In order for tendons to store energy effectively, the foot and ankle, the authors determined the greatest
they must be stretched through very specific ranges, since degree of motion occurs in the talonavicular joint (9.4°),
excessive stretching could produce injury while too small while the least amount of motion occurs in the subtalar
a range would limit energy storage; e.g., a rubber band joints (4.4°). In a recent series of important in vivo motion
stretched through a small a range of motion would not studies, Lundgren et al. (58) and Arndt et al. (57) inserted
return significant energy and an overstretched rubber band self-drilling intracortical pins into 9 bones of the foot and
would break. To provide the range of motion necessary for ankle to evaluate motion of the rear, mid, and forefoot as
the proper degree of stretching, joints of the foot and ankle 6 male subjects walked and ran slowly over level terrain.
have been designed to move through precise ranges of Subsequent 3-dimensional analysis confirmed motion
motion. In an in vivo study of the degree of arch lowering in the talonavicular joint was significantly greater than
associated with running, Ker et al. (73) determined the subtalar motion and in some situations, transverse and
average vertical deflection of the medial longitudinal arch frontal plane talonavicular motions were twice as much
is 7 to 10 mm. as subtalar motions occurring in the same planes. The
In a separate study of the effect of surface stiffness authors state that the curvature of the talar head and its
on energy efficiency, McMahon and Greene (74) corresponding navicular facet enables this joint to function
demonstrate that this exact distance provides the perfect with little osseous constraint, allowing for a considerable
degree of tendon stretch to maximize the return of energy. range of triplanar motion that makes it highly adaptable to
By having subjects run over experimental tracks made of the functional demands of the gait cycle.
varying stiffness, the authors confirmed that overly flexible Motion between the medial cuneiform and the
surfaces absorb too much energy while rigid surfaces navicular joint was much larger than expected, moving up
absorb too little, causing running speeds to be diminished to 10° in the frontal and sagittal planes (Fig. 3.32). The
in both situations. McMahon and Greene (74) prove that the authors claim that this degree of motion is likely to have
perfect track deflects vertically 7-8 mm and it returns more a significant effect on foot function, since the joints of the
than 90% of the elastic energy stored following heel strike, midfoot dorsiflexed through a larger range than the ankle in
resulting in significantly faster running speeds. Because 4 of 5 subjects tested. There was also an unexpected amount
this distance is identical to the degree of in vivo deflection of motion between the calcaneus and cuboid and between
noted by Ker et al. (71), McMahon (75) suggests that this the fifth metatarsal and cuboid, confirming that the lateral
may be the perfect degree necessary for ideal storage of arch of the foot has some capacity for accommodating
energy. The author specifically states that tendons in the surface irregularities and dampening motion. These
arch of the foot can return 17% of the energy absorbed two studies represent for the first time, a comprehensive
during a single step. description of in vivo kinematics during walking and slow
Although difficulties in quantifying energy storage
make it impossible to determine exactly which tissues
are responsible for specific amounts of energy storage, it
is possible the flexor digitorum brevis plays a significant
role, since this muscle is important in maintaining arch
height (increasing tone though barefoot exercise results
in an elevated arch as measured on weight bearing
X-rays [76]). Also, the of role flexor digitorum brevis in
force transmission is clinically evidenced by the fact that
calcaneal heel spurs form at the origin of this muscle,
not the origin of the plantar fascia (77). Because both
actomyosin cross-bridges and tendons are capable of
storing and returning energy (78), this muscle, in addition
to the long digital flexors, is in an ideal position to store
and return energy. By increasing tone in response to stress,
flexor digitorum brevis may behave as a variable length Figure 3.32. During stance phase, the medial cuneiform
spring that functions as a secondary restraint to vertical (mc) was shown to dorsiflex and invert up to 10° relative
lowering of the arch. to the navicular (n) (white arrow). Movement between the
While early research emphasized the importance medial cuneiform and the first metatarsal was inconsistent
of the subtalar joint in dampening ground-reactive forces, and approximately half that of the medial cuneiform and the
Kitaoka et al. (79) suggest that the subtalar joint possesses navicular (black arrow).
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running, emphasizing that in vivo motions in the foot and

ankle are more complicated than previously believed, and
that subtalar joint mobility is limited compared with the
talonavicular, first metatarsal-cuneiform and even cuboid-
fifth metatarsal joint motion. Interestingly, in all 3 planes,
the ranges of motion in the joints of the feet were greater
while walking than during slow running. The clinical
implication of this is that slow running is easier on the
joints of the foot and patients with various mechanical foot
disorders should not be discouraged from running slowly.
During the latter half of midstance, ground-reactive
forces centered beneath the heel begin to lessen, and the
subtalar joint begins to supinate, causing the foot to stiffen
in anticipation of the accelerational forces associated with
the propulsive period. Subtalar supination is accomplished
in part by taking advantage of the forward motion in the
contralateral lower extremity: the forward momentum of
the swing phase leg externally rotates the pelvis (white
arrow in Fig. 3.33), which then externally rotates the
weight-bearing leg (black arrow in Fig. 3.33). Since the leg
and talus behave as a closed kinetic chain during midstance,
external rotation of the weight-bearing leg causes the talus
to abduct, which in turn supinates the subtalar joint. This Figure 3.33. Forward motion of the swing phase
motion help stabilize the tarsal by decreasing parallelism leg externally rotates the stance leg, which in turn
of the midtarsal joint axes. supinates the subtalar joint. Mann (66) emphasizes
Supination of the subtalar and midtarsal joints during the role the adductors play in producing external rotation
of the stance leg by noting their firm attachment to the
late midstance is assisted by the return of elastic energy
anterior pelvis and posterior femur allows these muscles
stored in the muscles of the arch, by concentric contraction to act as effective lever arms capable of translating forward
of tibialis posterior, and most importantly, by the return of momentum of the swing leg into external rotation of the
energy stored in the Achilles tendon. Because the fibers of stance leg femur.
this tendon rotate approximately 90° before attaching to
the posterior calcaneus (Fig. 3.34), the Achilles tendon is
the most important energy storing tendon in the body. As
demonstrated by Ker (73), this tendon is capable of returning
35% of the energy used to stretch it. Despite its short lever
arm to the subtalar joint, the Achilles tendon generates the
vast majority of the inversion torque necessary to supinate
the subtalar joint. Subtalar supination during late midstance
creates a malalignment of the midtarsal joint axes (thereby
locking the midfoot), and displaces the progression of
the center of mass laterally, allowing the cuboid to begin
dorsiflexing into the overhanging calcaneus. Dorsiflexion
of the cuboid relative to the calcaneus was demonstrated in
the in vivo studies performed by Lundgren et al. (58) and
Arndt et al. (57), when the cuboid was shown to dorsiflex
on the calcaneus during late midstance while walking
and propulsion while slow running. Malalignment of the
midtarsal joint axes and locking of the calcaneocuboid
joint are necessary prerequisites for efficient propulsion.
Throughout late midstance, the hip and knee are
extending and externally rotating, allowing the knee to
lock (through twisting of the cruciate ligaments), while Figure 3.34. Rotation of the Achilles tendon. When
also allowing the hip to assist in the storage of energy: moving proximal to distal, the medial fibers move posteriorly
rotation of the hip’s capsular ligaments provides a soft (black dots) while the lateral fibers move anteriorly (white
tissue locking mechanism that decelerates hip extension dots) This results in a 90° twisting of the tendon (131).
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while also storing and eventually returning energy in the By the end of the midstance period, the ankle is
twisted fibers (refer back to figure 2.65). Perhaps more dorsiflexed 10° (forward momentum of the body coupled
importantly, the rectus femoris muscle, like the Achilles with simultaneous knee extension throughout midstance
tendon, also possesses a natural twist that allows this muscle allows ground-reactive forces applied beneath the forefoot
to store energy in a spring-like manner. As demonstrated to dorsiflex the ankle), the subtalar joint is moving towards
by Hasselman et al. (80), the indirect head of the rectus its neutral position, and the midtarsal joint is fully pronated
femoris muscle rotates near its proximal muscle tendon about both axes; i.e., the midtarsal joint has remained fully
junction, twisting 90° before attaching to the superior pronated about its oblique axis throughout midstance
acetabular rim (Fig. 3.35). Although the exact percentage although its available range of motion has significantly
of energy stored and returned is unknown, twisting of these decreased due to subtalar supination. Because ankle
fibers suggests the rectus femoris muscle plays a role in dorsiflexion displaces the naturally wider anterior talus
lessening the metabolic cost associated with flexing the hip upwardly (creating a bony block that stabilizes the ankle),
by storing and returning energy. the distal tibiofibular articulation gaps anteriorly while
the fibula moves posteriorly during late midstance. Stance
phase functions of the fibula are reviewed in figure 3.36.
Propulsive Period
The propulsive period begins the moment heel lift
occurs and ends with toe off. This period occupies the final
33% of stance phase while walking and lasts approximately
0.2 seconds. When running, propulsion is the longest
period, occupying 55% of stance phase, with forces on the
Achilles tendon, tibia, and the plantar forefoot averaging
7, 9, and 2.7 times body weight, respectively. Although
it appears to be a simple process, there are numerous
biomechanical factors responsible for producing heel lift.
First, forward momentum of the torso displaces the center
of mass directly over the forefoot, reducing the vertical
forces responsible for maintaining ground contact at the
heel (Fig. 3.37A). At that time, continued contraction of
the soleus and deep posterior compartment muscles acts
to limit the range of ankle dorsiflexion by decelerating the
forward motion of the proximal tibia. By limiting ankle
dorsiflexion, these muscles create a new pivot point at the
forefoot that allows forward momentum of the center of
mass to be applied directly towards lifting the heel (Fig.
3.37B).
The biarticular gastrocnemius muscle also plays
an important role in initiating heel lift by flexing the
knee while simultaneously plantarflexing the ankle.
These combined actions serve to lift the knee upward and
forward, providing a significant force to assist with flexing
the hip (Fig. 3.37C). Because it flexes the knee and hip,
gastrocnemius indirectly allows for improved ground
clearance during swing phase. In fact, Neptune et al. (81)
claim that by itself, the gastrocnemius muscle is the most
important muscle responsible for initiating swing phase.
These separate actions confirm that despite their shared
insertions into the Achilles tendon, the uniarticular soleus
and biarticular gastrocnemius behave very differently
Figure 3.35. Twisting of the indirect head of the rectus during gait.
femoris muscle. Notice in the transverse sections (B), To improve muscular efficiency, the gastrocnemius
that the indirect head of the rectus femoris (black) rotates fires almost isometrically during the latter half of stance
nearly 90° before attaching to the superior acetabular rim. phase, allowing the Achilles tendon to store and return
Redrawn from Hasselman et al. (80). energy. To prove this, Roberts et al. (82) surgically
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Figure 3.36. Summary of stance phase fibular motion. During the contact period (A), the fibula remains relatively
immobile as ankle plantarflexion and rearfoot eversion limit downward motion of the fibula. As demonstrated in a cadaveric
study by Wang et al. (128), eversion of the rearfoot tilts the lateral aspect of the talar dome superiorly, creating a physical
block that limits downward migration of the fibula. By midstance, the fibula supports 17% of the total load borne by
the lower leg and the interosseous membrane (im) plays a significant role in distributing force evenly throughout the
entire fibula (128). As vertical forces peak shortly before the push-off (C), the upward migration of the anterior talar dome
associated with ankle dorsiflexion forces the fibula to glide posteriorly approximately 1 mm (129), which serves to increase
ankle stability by setting the talus firmly in the ankle mortise. This close-packed position increases the percentage of weight
borne by the fibula from 17-30% (128). Although in vivo studies show inconsistent fibular motions during the latter half of
stance phase (56,57), cadaveric research suggests the fibula drops inferiorly during propulsion (black arrow in C), thereby
improving stability by deepening the ankle mortise (128). Theoretically, downward migration of the fibula at this time is
facilitated by contraction of soleus, flexor hallucis longus, and peroneus longus/brevis, all of which originate from the fibula
and are actively tensing during late stance phase. As with coupled motions in the spine, there is much individual variation
in fibular movement patterns and there are no preset rules defining motion.
Figure 3.37. Heel lift results from the combined actions of the forward momentum of body mass (1), muscular
deceleration of ankle dorsiflexion (2), and active flexion of the knee produced by gastrocnemius contraction (3).
Note that in panel A, the pivot point for motion is the ankle joint while soleus contraction in panel B locks the ankle (square)
and creates a new pivot point at the forefoot (black circle in B and C).
111
implanted special crystals and strain gauges into

gastrocnemius muscles and Achilles tendons of turkeys and
forced them to run on treadmills. By having the turkeys run
at progressively faster speeds, these researchers confirmed
that there was little change in gastrocnemius length
(approximately 7% change with each stride) and the muscle
generated just enough force to keep the Achilles stretched,
thereby allowing the tendon to store gravitational energy.
By calculating muscle activity and force production,
Roberts et al. (82) demonstrated that storing and recovering
energy in the stretched Achilles tendon was responsible
for more than 60% of the work performed by the muscle-
tendon complex.
This study confirms the spring-like action of the
Achilles tendon appreciably lessens the metabolic cost
of locomotion by reducing the workload on the muscular
support system. According to Taylor (83), it is metabolically
expensive for muscles to generate force when tensing either
concentrically or eccentrically and isometric contractions
are significantly more efficient because they produce large
forces with little metabolic expense. By converting the
gradual eccentric contraction present during midstance
into an “isometric impulse” that allows the Achilles Figure 3.38. Cadaveric model evaluating plantar
tendon to store and return energy at just the right moment, pressures and metatarsal bending strains present
isometric contraction of the gastrocnemius allows the body during heel lift. Redrawn from Ferris et al. (84).
to take advantage of the “free elastic energy” associated
with stretching this large, well-designed tendon. According
to Anderson (18), optimizing the storage and return of pressure beneath the hallux and second toe decreased
energy requires considerable practice as it is a learned 27% and 80%, respectively. More importantly, the authors
process, involving precise timing of numerous kinetic confirmed the reduced pressure beneath the toes produced
and kinematic variables. Eccentric contraction followed a corresponding increase in pressure beneath the metatarsal
by near isometric contraction (allowing for the storage of heads, resulting in large bending strains being placed on the
energy) is not unique to the gastrocnemius/soleus, since it metatarsal shafts. Apparently, a primary role of the digital
occurs in almost every other shock absorbing muscle in the flexors is to create a compressive force along the plantar
body (e.g., tibialis anterior and vastus lateralis). metatarsal shafts that counteracts the bending moments
Once the heel has left the ground, a considerable associated with propulsion. The authors emphasize that
amount of stress is transferred directly into the forefoot. although the toe flexors are relatively weak compared to
To protect the forefoot from the extreme ground-reactive the gastrocnemius and soleus muscles, weakness of the
forces associated with propulsion, stimulation of cutaneous long digital flexors can have a profound effect on regional
receptors in the skin (e.g., Meissner’s corpuscles) cause forefoot pressures. They go on to suggest that fatigue of
the digital flexors to tense reflexively, producing a strong these muscles would result in elevated metatarsal shaft
plantarflexion force at the toes that significantly reduces bending moments, predisposing to stress reactions. Flexor
pressure beneath the metatarsal heads. hallucis longus is particularly important, since the loss of
The exact degree of protection provided by the function at the hallux results in a transfer of load to the
digital flexors was determined in an interesting study by lateral forefoot that may increase bending strains placed on
Ferris et al. (84). By mounting cadaveric feet in a heel the lesser metatarsal shafts.
rise position on a specially designed apparatus, these In a separate study of forces acting on the foot
authors measured plantar contact pressures in the forefeet during propulsion, Jacob (85) determined that flexor
before and after sequentially applying tension to a series hallucis longus and brevis exert forces of 52% and 36%
of tendons clamped to pneumatic actuators. The authors body weight, respectively, while flexor digitorum longus
also embedded strain gauges into the second metatarsal and brevis exert forces of 9% and 13% body weight,
shafts in order to measure bending forces present in the respectively. Besides distributing pressure and reducing
bone with and without simulated muscle contraction metatarsal bending strains, these muscles play an
(Fig. 3.38). Using this elaborate technique, Ferris et al. important role in postural stability. In a study of balance
(84) concluded that in the absence of the digital flexors, in the elderly, Menz et al. (86) determined that decreased
112
plantar tactile sensation and weakness of the digital medially as it abducts and dorsiflexes about the OMJA
flexors were independent predictors of impaired balance (Fig. 3.39). Notice in this illustration how external rotation
and may be associated with an increased risk of falls. By of the leg creates a screw-like motion at the midfoot that
muscularly controlling anterior displacement of the center greatly increases arch height, essentially converting the
of mass during propulsion, the digital flexors increase the foot into a rigid lever.
size of the fall envelope. After evaluating age and gender Supination about the oblique axis of the midtarsal
strength differences in the digital flexors, Menz et al. (87) joint is aided by contraction of the intrinsic muscles
determined that older individuals, on average, possess originating from the medial calcaneus (particularly
32% less plantarflexion strength in the hallux and 27% abductor hallucis) and by what is known as the windlass
less plantarflexion strength of the lesser toes; and women effect of the plantar fascia: dorsiflexion of the toes after
possess 42% less hallux plantarflexion strength than men. heel lift draws the plantar fascia around the metatarsal
According to the authors, improving strength in the digital heads, pulling the anterior and posterior pillars of the
flexors will theoretically improve balance and may reduce longitudinal arch together (Fig. 3.40). This approximation
the risk of falling. of the rearfoot and forefoot allows for continued supination
In addition to the support provided by the digital about the oblique midtarsal joint axis, with its associated
flexors, the joints of the midfoot must be architecturally increase in arch height.
stable in order to tolerate the accelerational forces associated Using fiberoptic cables embedded in cadaveric
with early propulsion. The primary factor responsible plantar fascias during simulated walking, Erdemer et al.
for stabilizing the midfoot is supination of the midtarsal (88) demonstrate that force in the plantar fascia gradually
joint. This important action locks the talonavicular and increases during contact and midstance, peaking at 96%
calcaneocuboid joints, preventing the chimp-like buckling body weight during propulsion. These researchers conclude
that would otherwise occur (refer back to figure 1.19). In that because there is a strong correlation between Achilles
order to efficiently supinate the midfoot, the body takes tendon and plantar fascial stress, the plantar fascia serves as
advantage of the external shank rotation supplied by the an intermediary, transmitting force from the Achilles tendon
forward momentum of the opposite swing phase leg. into the forefoot during propulsion. In an in vivo study of
Because the closed kinetic chain ends at the metatarsal plantar fascial mechanics, Wearing et al. (89) determined
heads after heel lift occurs, continued external leg rotation the long and short digital flexors protect the plantar fascia
supinates the subtalar joint beyond its neutral position by decelerating the range of metatarsophalangeal joint
(ground-reactive forces no longer maintain the calcaneus dorsiflexion during propulsion, thereby lessening tensile
in a fixed position so it is free to move with the rotating strain in the plantar fascia as these muscles absorb force
talus), while markedly supinating the forefoot about the that would otherwise go into the stretching plantar fascia.
oblique midtarsal joint axis: The entire rearfoot pivots While considerable stability is afforded by elevation
of the arch associated with the windlass effect of the
plantar fascia, the foot could not be considered a rigid lever
were it not for the continued forefoot pronation about the
longitudinal midtarsal joint axis. During early propulsion,
the calcaneocuboid locking mechanism is maintained
by forceful contraction of the soleus muscle, which
simultaneously plantarflexes the ankle while inverting
the subtalar joint. Despite its short lever arm, this muscle
possesses 5 times the mass of any other deep posterior
compartment muscle and is therefore an effective supinator
of the subtalar joint (90). While ankle plantarflexion allows
for a forward acceleration of body mass, subtalar joint
inversion allows ground-reactive forces to dorsiflex the
fourth and fifth metatarsals, locking the lateral column.
The effectiveness of the soleus muscle in
maintaining the midtarsal locking mechanism is only
temporary, since early in propulsion the range of ankle
Figure 3.39. External leg rotation (A) acts to supinate plantarflexion places soleus in such a shortened position
the subtalar joint (B) while simultaneously supinating that it is unable to generate sufficient force to invert the
forefoot about the oblique midtarsal joint axis (C). calcaneus. At this time, the continued forceful contraction
These motions increase arch height (black arrow), thereby of peroneus longus (which passes beneath the cuboid in
stabilizing the various articulations of the midfoot during the peroneal groove) acts to dorsiflex and evert the cuboid,
propulsion. thereby maintaining the close-packed position of the
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Figure 3.40. The windlass effect of the plantar fascia. During the propulsive period, ground-reactive forces dorsiflex
the toes, which pulls the plantar fascia around the metatarsal heads (A). This action results in the approximation of the
rearfoot and forefoot (B) and allows for the increased arch height necessary for stability (C). The amount of pull generated
by the plantar fascia is directly related to the distance between the transverse axis of the metatarsophalangeal joint and
the passage of the plantar fascia: the greater the distance, the greater the pull placed upon the plantar fascia while the
digit dorsiflexes. For example, the average lesser metatarsal has an average of 8 mm between its transverse axis and the
passage of the plantar fascia (D) while the first metatarsal, with its larger head and the presence of sesamoid bones (which
the plantar fascia invest) has a distance of nearly 15 mm between the transverse axis and the plantar fascia (E) (91). As a
result, dorsiflexion of the first digit produces a much greater tractioning effect on the plantar fascia than any of the lesser
digits (compare F and G). In order to resist the greater tensile load, the plantar fascia has its strongest attachment distal to
the first metatarsal head. The plantar fascia also has strong attachments to the skin beneath the metatarsal heads (star),
which prevents sliding on the skin as posterior shear forces are applied during the propulsive period (91). The short digital
flexors play an important role in reducing strain placed on the plantar fascia because they reinforce the arch and stabilize
the metatarsophalangeal joints, essentially behaving as a variable length buttress to lessen load borne by the plantar fascia.
calcaneocuboid joint (Fig. 3.41). Furthermore, because as strong as the remaining lesser metatarsals (21). In a
the fourth and fifth metatarsals are shorter than the remaining cadaveric study of forefoot pressures associated with the
metatarsals, the lateral column is unable to maintain ground propulsive period of walking, Jacob (85) demonstrates that
contact during mid and late propulsion and is therefore the first metatarsal head supports a force of 119% body
unable to assist with the forward acceleration of body weight while the second metatarsal head supports only
mass. Locking of the calcaneocuboid joint at this time 28%. Because of the force difference, Jacob (85) states
continues to serve a purpose, because it affords peroneus that if the first metatarsal were unable to bear weight, the
longus and brevis an effective lever arm as they now second metatarsal would fail, since it could not tolerate the
function to direct body weight medially towards the transfer of forces.
opposite foot by everting the entire lateral column (Fig. Because of its passage under the cuboid and
3.42). eventual insertion into the base of the first metatarsal
By lifting the lateral column and transferring and medial cuneiform, peroneus longus has the
weight medially, peroneus brevis plays an important role interesting ability to transfer body weight medially
in improving speed by transferring force to the medial while simultaneously stabilizing the medial forefoot
forefoot. This medial shift of body weight is necessary so it may better tolerate these forces. This stabilizing
to maintain a straight gait pattern and to allow the action is related to the improved angle of approach
final transfer of vertical forces to occur off the medial afforded the peroneal longus tendon as the subtalar joint
forefoot, which is better equipped to handle these forces is supinating (Fig. 3.43).
since the first metatarsal is twice as wide and 4 times
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Figure 3.41. Concentric contraction of peroneus longus

during early and mid propulsion serves to lift (dorsiflex
and evert) the cuboid, thereby locking the midtarsal
joint.
Figure 3.42. Because they are so short, the fifth metatarsal leaves
the ground approximately 33% of the way through the propulsive
period, with the fourth metatarsal leaving shortly thereafter. At that
time, continued contraction of the lateral compartment musculature
serves to shift body weight medially towards the opposite foot (arrow),
which is just beginning its contact period.
Figure 3.43. The effect of subtalar positioning on peroneus longus function. When the subtalar joint is pronated
(A), the nearly horizontal angle of approach afforded peroneus longus allows for the production of a strong posterolateral
compressive force (1) and a mild dorsiflectory force about the first ray axis (2). As the subtalar joint moves into a progressively
more supinated position (B and C), the posterolateral compressive force is lessened (3), and the more vertical approach of
the peroneus longus tendon allows for the development of a strong plantarflectory force about the first ray axis (4).
115
The improved ability of peroneus longus The combined actions of peroneus longus as an
to function as a first ray plantarflexor is extremely evertor of the lateral column and a plantarflexor of the
important during the propulsive period, because the first ray allow for what Bojsen-Moller (91) refers to
increased height of the medial longitudinal arch coupled as a high gear push-off. By everting the lateral column,
with the normal parabolic curve of the metatarsal heads the peroneals allow the final transfer of body weight to
(i.e., the first metatarsal is normally shorter than the occur through the transverse axis of the metatarsal heads
second metatarsal) necessitates the first ray actively (Fig. 3.45). Use of the transverse axis supplies the ankle
plantarflex in order to maintain ground contact. Besides plantarflexors with a longer, more effective lever arm for
the obvious importance of maintaining ground contact to accelerating body mass forward. Failure of the peroneals
resist ground-reactive forces, active plantarflexion of the to evert the lateral column would allow for continued
first metatarsal allows for the dorsal-posterior shifting supination of the subtalar joint with the final push-off
of the first metatarsophalangeal joint’s transverse axis occurring as a rolling action through the oblique axis
needed for the hallux to reach its required range of of the metatarsal heads. Because the oblique axis has a
dorsiflexion (Fig. 3.44). shorter lever arm to the ankle axis (the oblique axis is
15-20% closer to the ankle axis than the transverse axis), it
allows for a less efficient propulsion referred to as a low
gear push-off. It is clinically interesting that transitions
to faster speeds of running are associated with significant
increases in peroneus brevis activity, with little change
in activity of the gastrocnemius and soleus muscles (92).
According to Bojsen-Moller (91) use of the transverse axis
by means of peroneus longus/brevis contraction represents
the final evolutionary change in the process of producing a
fast, efficient propulsion.
During the final portion of the propulsive period,
the foot will ideally be supinated about the oblique
midtarsal and subtalar joint axes. In addition, the
forefoot will remain fully pronated about the longitudinal
midtarsal joint axis. This axis is maintained in a pronated
position during late propul sion as extensor digitorum
Figure 3.44. Because the first metatarsal is normally

shorter than the second metatarsal, it actively
plantarflexes to maintain ground contact during
the propulsive period (A). As the first metatarsal
plantarflexes, the metatarsal head glides posteriorly along
the sesamoids (B), which allows for a dorsal-posterior shift
of the transverse axis of the first metatarsophalangeal
joint (C). This new axis allows for an unrestrained range
of hallux dorsiflexion (D). Failure of the first metatarsal to
plantarflex during propulsion (E) inhibits the posterior glide
of the metatarsal head on its sesamoid (F), which in turn
prevents the dorsal-posterior shift of the transverse axis.
The hallux is now forced to dorsiflex about the original axis
Figure 3.45. The transverse and oblique axes of the
(G). This results in a “jamming” of the dorsal cartilage (H)
metatarsal heads.
with its characteristic resorption of subchondral bone and
dorsal lipping of the first metatarsal head.
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longus and peroneus tertius are vigorously contracting in

preparation for the swing phase of gait. The final transfer
of forces should occur through the hallux, which has been
stabilized throughout the propulsive period by vigorous
contraction of flexor hallucis longus. The propulsive
period ends when the hallux leaves the ground.
Swing Phase Motions

The swing phase begins at toe off and ends at
heel strike, occupying 38% of the gait cycle and lasting
approximately 0.4 seconds (1). The primary kinematic
actions during this phase are to provide ground clearance of
the forefoot by midswing and to position the articulations
in such a way that the supporting muscles are prepared
to dampen impact forces as the next heel strike occurs.
Because the typical lower extremity weighs 15% of body
mass, and each leg moves through a full cycle 2,500 times
each day, the average 160-pound person must accelerate
and decelerate 56,000 pounds of force per leg, per day.
The late stance phase activity of the gastrocnemius muscle
has already played an important role in lessening the Figure 3.46. During the late propulsive period, tension
metabolic cost associated with initiating swing phase by created in the psoas (A), iliacus (B) and rectus femoris
(C) extend the lumbar spine and tilt the pelvis forward
storing energy in the Achilles during late midstance, and
(black arrow above acetabulum). To protect against these
returning it during propulsion to drive the knee and hip up potentially injurious motions, the external obliques (D) and
and forward. This markedly reduces strain on the iliopsoas, rectus abdominus (E) pull the anterior pelvis upwardly,
rectus femoris and hamstring musculature, which, were it counteracting forces created by the hip flexors, allowing the
not for the energy returned from the Achilles, would have spine and pelvis to function as a stable anchor.
fired with significantly more force to swing the lower
extremity forward.
Even though the total workload performed by function is similar when walking with and without legs
the iliopsoas muscle is reduced by the highly efficient and the abdominal muscles, without contribution from the
gastrocnemius/Achilles tendon complex, contraction of iliopsoas and lower extremities, are capable of producing a
these muscles during the initiation of swing phase imposes very smooth gait. This model of muscle function suggests
substantial compression on the lumbar vertebrae as the that the iliopsoas functions in a manner similar to the plantar
psoas extends the lumbar spine while the iliacus tilts the fascia during heel lift, in that it transfers force generated in
pelvis anteriorly. To lessen strain and protect the spine a proximal area into a distal area. Of course, the iliopsoas
and pelvis from the elevated shear force associated with can add to these forces by concentrically contracting while
combined iliopsoas activity, the multifidus, erector spinae, the plantar fascia provides a more passive transfer of force.
rectus abdominus, abdominal obliques and latissimus dorsi Because the ankle reaches its maximally
muscles actively contract during propulsion, with peak plantarflexed position shortly after toe off, the anterior
forces occurring at the initiation of swing phase (60).These compartment mus cles have less than 0.2 seconds to
muscular actions lessen strain by preventing spinal and overcome inertial forces and dorsiflex the forefoot into
pelvic hyperextension, locking the spine and pelvis while a safe position by midswing. Since extensor digitorum
the lower extremity is being pulled forward (Fig. 3.46). longus and peroneus tertius are the first anterior
Gracovetsky (93) proves that activity in the lumbar compartment muscles to contract (94), the foot, in addition
spinal musculature also plays an important role in initiating to dorsiflexing at the ankle, will immediately pronate
swing phase by demonstrating that individuals born without about the oblique midtarsal and subtalar joint axes. (These
legs are capable of walking with extremely efficient gait muscles possess significant lever arms for pronating these
patterns by pivoting on their ischial tuberosities. Kinematic axes.) The dorsiflectory components of these pronatory
analysis of these individuals reveals that they maintain motions allow for improved ground clearance.
their lumbar spines in lordotic positions, converting lateral Almost immediately after extensor digitorum longus
bending moments of the spine into an axial rotation that and peroneus tertius contract, tibialis anterior and extensor
drives the pelvis forward. Referred to as the spinal engine, hallucis longus begin contracting, markedly in creasing
electromyography of their torsos confirms that although the dorsiflectory movement created at the ankle. Root et
the amplitude of spinal motions vary, abdominal muscle al. (94) claim that extensor hallucis longus is the strongest
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ankle dorsiflexor during early swing phase. Tibialis object, clearing the ground by such a small margin lessens
anterior, by virtue of its insertion on the first metatarsal and the metabolic cost of locomotion by decreasing strain on
medial cuneiform, also acts to improve ground clearance the hip, knee and ankle musculature.
by dorsiflexing the first ray during early swing phase Shortly after the forefoot has cleared the ground,
(Fig. 3.47). Notice in this illustration how the forefoot is muscles of the swing leg have a relatively quiet period
maintained in an everted position about the longitudinal when motion is maintained by inertial forces generated
midtarsal joint axis during early and midswing phase by during propulsion and early swing (96). At this time,
the continued contraction of extensor digitorum longus the vacuum phenomena present in the hip decreases the
and peroneus tertius. The dorsiflectory motion of the first metabolic cost of locomotion by temporarily supporting
ray, besides improving ground clearance, also serves to en the weight of the lower extremity (97). As discussed in
hance the efficiency of extensor hallucis longus as an ankle Chapter 2, the hip labrum creates such a tight seal around
dorsiflexor, since it results in an anterior/inferior shift of the periphery of the acetabulum that even when all of the
the first metatarsophalangeal joint’s transverse axis (Fig. hip’s supporting muscles and ligaments are cut, the weight
3.48). of the entire lower extremity can be supported solely by
By far, the most important kinematic factor the negative intraarticular hip pressure generated by the
responsible for producing ground clearance is flexion of dangling lower extremity. This negative pressure provides
the knee. In fact, if for any reason the knee is unable to stability and decreases the metabolic cost of locomotion
adequately flex during midswing, the metabolic cost of by reducing the muscular effort necessary to maintain the
locomotion skyrockets, because the individual is forced femur in the acetabulum during the middle of swing phase.
to circumduct the stiff swing phase lower extremity by Following the midswing quiet period, the hamstrings
excessively abducting both the stance and swing phase hips. begin eccentrically contracting to decelerate forward
By the time midswing has occurred, the hip and knee are motion of the swinging leg. The bicep femoris is the most
flexed 30° and 50°, respectively; the ankle is dorsiflexed to efficient decelerator of the swinging leg, since its more
a near neutral position; the subtalar and midtarsal joints are distal insertion provides a longer lever arm for decelerating
pronated (the midtarsal joint is pronated about both axes); the swinging leg (Fig. 3.49). Although prior researchers
and the first ray is dorsiflexed and inverted. In an attempt suggested that the dual innervation of this muscle was
to reduce the metabolic cost of locomotion, the large hip responsible for its high rate of injury while running (98),
and thigh muscles contract with only enough force to Thelen et al. (99) confirm the lower attachment of the bicep
allow the foot to barely clear the ground; i.e., the typical femoris produces greater tensile strains in this muscle
foot clears the ground by only 1.29 cm (95). Although this during late swing phase, explaining the higher injury
occasionally causes the individual to trip on an unseen rate. The authors demonstrate that compared with upright
postures, the bicep femoris lengthens 9.5% during late
swing phase, while semitendinosus and semimembranosus
lengthen 8.1% and 7.4%, respectively. As is consistent
with muscles responsible for dampening vibration, the
bicep femoris muscle has short muscle fibers and long
tendons, allowing it to dampen vibration very effectively.
In a separate study of the long head of the bicep femoris
during sprinting, Thelen et al. (100) determined that
following the onset of muscle activity during late swing
phase, stretching of the muscular component of the bicep
femoris slowed considerably while the tendon lengthened
and stored elastic energy. The authors emphasize that a
more compliant tendon reduces peak muscle stretch and
negative muscle work, potentially lessening the risk of
injury.
Just before making ground contact, the anterior
compartment muscles simultaneously contract in
anticipation of dampening the impact forces associated
with the contact period. Because of their relationships with
Figure 3.47. During early swing phase, the lateral
the various axes, tibialis anterior and extensor digitorum
branches of extensor digitorum longus and peroneus
tertius actively pronate the forefoot (A) while tibialis longus produce mild dorsiflexion at the ankle, with tibialis
anterior, in addition to dorsiflexing the ankle, actively anterior markedly inverting the forefoot while extensor
dorsiflexes and inverts the first ray (B), thereby allowing digitorum longus and peroneus tertius assist with ankle
for improved ground clearance. dorsiflexion and pronation of the forefoot about the oblique
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Figure 3.48. Dorsiflexion of the first ray by tibialis anterior (A) shifts the first metatarsophalangeal joint’s transverse
axis back to its original position (black dot), thereby limiting the range of hallux dorsiflexion possible. The
plantarflectory motion of the hallux improves the efficiency of extensor hallucis longus as an ankle dorsiflexor by stabilizing
its insertion on the distal phalanx (star).
Figure 3.49. Because the distal bicep femoris has a lower point of attachment than the semitendinosus (st) and
semimembranosus (sm), forward motion of the leg during swing phase creates a greater tensile strain in the bicep
femoris muscle. Modified and redrawn from Thelen et al. (99).
119
midtarsal joint axis. Basmajian and Deluca (51) state frontal plane stability of the pelvis during transitions from
that tibialis anterior acts as an ankle dorsiflexor during double-limb support to single-limb support. Peak activity
early swing phase and an inverter of the forefoot during is seen during the early and midpropulsive periods (51),
late swing phase. By positioning the foot with the ankle when the erector spinae tense to maintain frontal plane
dorsiflexed, the forefoot inverted, and the subtalar joint alignment of the pelvis. Waters and Morris (105) note a
slightly supinated, the pretensed muscles of the foot and brief burst of activity during the latter half of swing phase,
leg are now prepared to dampen ground-reactive forces when this muscle possibly contracts to assist the weakening
associated with stance phase. It is interesting that with contralateral gluteus medius in raising the pelvis in prepa
sprint running and anticipated falls, other shock-absorbing ration for heel strike. As noted by McGill (60), longissimus
muscles (lateral gastrocne mius, vastus lateralis, gluteus thoracis pars lumborum and iliocostalis lumborum pars
maximus, bicep femoris, etc.) become hyperactive prior lumborum limit forward motion of the lumbar spine
to heel strike when they pretense in an effort to more following heel strike, dampening impact forces on
effectively dampen the perceived increase in ground- individual vertebra while the spine flexes forward. Because
reactive forces (101,41). longissimus thoracis and iliocostalis lumborum attach to
the lower 6 ribs, they function to decelerate forward motion
Summary of Gait Cycle Muscle Function: of the torso.
Gluteus Maximus
This muscle contracts during late swing and early Gluteus Medius
stance phase to decelerate flexion and initiate extension at Gluteus medius is the primary frontal plane stabilizer
the hip (although it may also mildly assist with abduction of the pelvis. It begins contracting during late swing and
of the hip). Basmajian and Deluca (51) demonstrate that continues throughout midstance and into propulsion. Peak
during terminal stance (toe off), the middle fibers of gluteus activity occurs during early midstance when this muscle
maximus display a brief burst of activity. Contraction at this vigorously contracts to prevent excessive lowering of the
time possibly allows these fibers to assist gluteus medius contralateral pelvis (which is entering its swing phase).
with abducting the swing phase leg. Lyons et al. (102) note Basmajian and Deluca (51) note a brief burst of activity
that the angles of approach afforded the various sections of in the anterior fibers of gluteus medius during toe off;
this muscle allow its lower portion to act as a hip extensor possibly to assist with abducting and internally rotating
while the upper portion acts as a hip abductor. Cadaveric the femur during early swing phase. Because the lower
studies confirm gluteus maximus has the greatest capacity fibers of the gluteus medius muscle parallel the femoral
for controlling internal rotation of the hip. neck, contraction of these fibers lessens bending forces at
In an important in vivo analysis of gluteus maximus this site. Gluteus medius also plays a role in distributing
function during walking, Preece et al. (103) demonstrate pressure throughout the femoroacetabular joint during
that this muscle places large external torques on the stance phase (55), functioning in a manner similar to the
femur, ultimately leading to rapid deceleration of the tibia rotator cuff musculature of the glenohumeral joint.
during early stance phase. Although the common belief
is that rotation of the lower leg is controlled by pronation Gluteus Minimus
of the subtalar joint (i.e., talar adduction associated with This muscle functions agonistically with gluteus
subtalar pronation maintains the lower extremity in an medius during early stance phase. The brief burst of
internally rotated position), these authors confirm that activity during midswing most likely allows for continued
gluteus maximus plays a significant role in controlling internal rotation of the femur. Like the gluteus medius, this
transverse plane motions of the lower leg, overcoming muscle also functions to maintain femoroacetabular joint
the influence of subtalar pronation. This is consistent with pressures at a steady level during stance phase.
research by Bellchamber and van den Bogert (104), who
demonstrate that during the late stance phase of gait, the
hip musculature plays a more important role in controlling Tensor Fasciae Latae
rotation of the lower extremity than the foot and ankle. Because of its insertion into the anteroproximal
Because the largest portion of gluteus maximus inserts iliotibial band, contraction of tensor fasciae latae during
into the posterior fibers of the iliotibial band, the gluteus the contact period acts to balance the force placed on the
maximus muscle provides significant stability to the hip iliotibial band by the simultaneous contraction of gluteus
and knee during contact and midstance, particularly while maximus. Contraction of tensor fasciae latae during early
running as knee flexion angles increase. stance increases tension in the anterior aspect of the band,
providing stability while walking. The tensor fasciae latae
Erector Spinae also contracts with significant force during late propulsion
Because iliocostalis is the most lateral of the erector and early swing, when it assists iliopsoas with flexing the
spinae musculature, it is able to assist in maintaining hip.
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prevent impingement by drawing the medial menis cus

Iliopsoas posteriorly while the knee flexes. During propulsion,
The iliopsoas muscle demonstrates peak activity semitendinosus demonstrates a mild burst of activity (51),
during terminal stance and early swing phase, when it when it may assist gastrocnemius with flexing the knee.
assists the adductors, tensor fasciae latae, rectus femoris Elliot and Blanksby (108) note that with running, all of the
and sartorius with flexing the hip. Momentum gained hamstring muscles maintain high levels of activity during
by rapid thigh flexion during early swing phase plays an the propulsive period, when they function as powerful knee
important role in forward acceleration of the center of flexors and moderate hip extensors. Throughout late swing
mass during late swing phase (66). Although occasionally phase, the more distal attachment of bicep femoris results
referred to as a spinal stabilizer, EMG analysis confirms the in an increased stretching of this muscle compared to the
psoas muscle functions primarily as a hip flexor (60,106). other hamstrings (99), allowing it to stabilize the sacroiliac
Because contraction of the iliacus muscle tilts the pelvis joint through tension transferred into the sacrotuberous
anteriorly, extending the lumbar spine, the psoas muscle ligament. Pretensing of the bicep femoris may also play
counters these motions by anchoring the lumbar spine to an important role in dampening bony and soft tissue
the hip, allowing the iliacus to act as a hip flexor. Despite vibrations associated with foot strike (41). Because of its
its ability to externally rotate the hip, an EMG evaluation multiple attachments on the posterior oblique ligament
by Juker et al. (106) revealed only slight activity of the and the medial collateral ligament, the semimembranosus
psoas during hip rotation tasks. As mentioned, because of muscle plays an important role in resisting valgus strains
their different origins and functions, McGill (60) feels the at the knee.
iliopsoas should be referred to as two separate muscles.
Quadriceps
Sartorius The quadriceps pretense during late swing phase and
Sartorius is active only during swing phase with demonstrate peak activity during the early contact period,
peak activity shortly after toe off. Because of its origin on when they eccentrically contract to decelerate knee flexion.
the anterior superior iliac spine (ASIS) and insertion on the These muscles continue to contract until the center of mass
proximal anteromedial tibia, this muscle is able to assist passes in front of the knee (1). A brief and less forceful
with flexion of the knee and hip while simultaneously burst of activity is seen during late stance and early swing
internally rotating the tibia during the first half of swing phase as rectus femoris assists with hip flexion (particularly
phase. at faster speeds) and the quadriceps function as a group to
decelerate the range of knee flexion associated with early
Adductors swing. The quadriceps muscles are by far the body’s most
The adductors, as a group, demonstrate peak important shock absorbers, and activity in these muscles
activity during toe off, when they flex the hip and possibly increases as the degree of knee flexion increases. While
assist with internally rotating the swing phase femur. These the vastus medialis obliquus stabilizes the medial patella,
muscles again contract during late swing as they pretense the lower fibers of vastus lateralis and fibrous slips from
in anticipation of ground-reactive forces. Although there is the iliotibial band stabilize the lateral patella, preventing
much individual variation in the behavior of the adductors displacement while the knee flexes and extends.
(1), Basmajian and Deluca (51) note that with the exception
of a brief quiet period at midswing, adductor magnus fires Popliteus
constantly throughout the gait cycle. Also, it is possible The popliteus muscle is a stance phase muscle that
that the more horizontal sections of the adductors assist the has a slight peak in activity during heel strike with another
contralateral swing phase pelvis with externally rotating more sustained peak through midstance and propulsion.
the femur during the ipsilateral midstance period (66). During the contact period, popliteus concentrically
contracts to assist the posterior cruciate ligament in
Hamstrings preventing excessive forward glide of the femur on the
The hamstrings demonstrate peak activity tibia, while simultaneously assisting the subtalar joint with
during the terminal portion of swing phase, when they internally rotating the tibia. Because this latter action is
eccentrically contract to decelerate forward motion of the necessary to unlock the cruciate and collateral ligaments,
rapidly extending leg. These muscles continue to contract popliteus is often referred to as “the key to the knee.”
through the majority of the contact period, at which time Throughout midstance, popliteus eccentrically contracts
they assist the gluteus maximus with decelerating flexion to aid gastrocnemius with decelerating extension at the
and initiating extension of the hip joint. Interestingly, knee. During propulsion, popliteus again concentrically
Hollinshead and Jenkins (107) note that because the contracts, producing external rotation of the femur. This
distal semimembranosus sends fibrous attachments to action is necessary for knee flexion to occur because it
the posterior horn of the medial meniscus, it is able to restores the normal coupled motions present in the knee;
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i.e., throughout the propulsive period, the knee is flexing

while the tibia is ex ternally rotating (elevation of the
arch abducts the talus, which externally rotates the tibia). Tibialis Posterior, Flexor Digitorum Longus, and
Because these motions conflict with the normal coupled Flexor Hallucis Longus
motions associated with knee flexion (the tibia should Tibialis posterior functions during early stance
be internally rotating as the knee flexes), the femur must phase as the body’s most important decelerator of subtalar
externally rotate farther than the tibia for knee flexion to pronation. It is also important during the latter half of
occur. The greater range of external femoral rotation allows stance phase, when it assists the gastrocsoleus complex and
the normal coupled motions to occur, as even though the the Achilles tendon with plantarflexing and inverting the
tibia continues to externally rotate, it remains internally foot to allow for an efficient propulsive period.
rotated relative to the more externally rotated femur. The long digital flexors play important roles
during terminal midstance, as they assist with heel lift by
Tibialis Anterior, Extensor Hallicus Longus, decelerating the forward momentum of the proximal tibia
Extensor Digitorum Longus, and (see Fig. 3.37). The digital flexors continue contracting
Peroneus Tertius throughout most of the propulsive period, forcefully
The anterior compartment musculature demonstrates maintaining the digits against the ground and assisting
peak activity immediately after heel strike. During the con abductor hallucis with supinating the foot about the
tact period, these muscles decelerate ankle plantarflexion, oblique midtarsal joint axis. Because of its attachment
with tibialis anterior maintaining the forefoot in an inverted to the flexor digitorum longus tendons, the quadratus
position about the longitudinal midtarsal joint axis during plantae muscle significantly increases forces transferred
the early and midcontact periods. (Ground-reactive forces through this tendon. In fact, EMG analysis confirms the
maintain this inverted position during the late contact quadratus plantae muscle may act as a primary flexor of the
period.) When walking, these muscles are normally lesser toes, being preferentially recruited over the flexor
inactive during mid stance and again contract during digitorum longus (109).
terminal stance. When running, the anterior compartment
muscles remain active during midstance, during which Gastrocnemius and Soleus
time they function to accelerate the body by pulling the Both soleus and gastrocnemius demonstrate peak
proximal tibia over the fixed foot (66) activity during terminal midstance, when they function
Because extensor digitorum longus and peroneus to produce heel lift. Soleus prevents forward motion of
tertius are the first anterior compartment muscles to the proximal tibia (decelerating ankle dorsiflexion) while
contract during the propulsive period (94), they are able to gastrocnemius flexes the knee and plantarflexes the ankle
dorsiflex the ankle while simultaneously maintaining the (initiating heel lift). The femoral origin of gastrocnemius
forefoot in a pronated position about the oblique midtarsal allows this muscle to maintain a constant flexion tension
joint axis. (Extensor digitorum longus also acts to maintain on the knee throughout midstance, lessening the risk of
a compressive force on the lesser metatarsophalangeal hyperextension injury.
and interphalangeal joints, which prevents clawing of the During the contact period, soleus decelerates internal
digits.) During terminal stance, the early swing phase rotation of the tibia while gastrocnemius decelerates
contraction of tibialis anterior also assists with ankle internal rotation of the femur. These dual actions reduce
dorsiflexion, but its insertion on the medial cuneiform the buildup of torsional strains at the knee during the early
and first metatarsal allows it to produce simultaneous stance. Soleus continues to contract through midstance
dorsiflexion and inversion of the first ray. The extensor and into early propulsion, when it supinates the subtalar
hallucis longus muscle acts to maintain tension on the joint, externally rotates the tibia, and stabilizes the lateral
hallux during late stance and early swing, when it behaves forefoot against the ground (thereby locking the lateral
as the strongest ankle dorsiflexor. column). Gastrocnemius continues to contract throughout
The anterior compartment muscles usually midstance and into propulsion, when it assists with
demonstrate a brief period of inactivity shortly after subtalar joint supination and external femoral rotation. Of
midswing, which is followed by simultaneous contraction course, in addition to assisting with heel lift, one of the
of all of these muscles during terminal swing (51). This most important actions of the gastrocnemius muscle during
simultaneous late swing phase activity allows for mild late propulsion is to drive the knee up and forward, thereby
dorsiflexion of the ankle and metatarsophalangeal joints lessening strain on the hip flexors and indirectly assisting
with extensor digitorum longus and peroneus tertius with producing ground clearance by midswing.
reestablishing the forefoot in a pronated position about the
oblique midtarsal joint axis. The late swing phase activity
of tibialis anterior produces marked inversion of the
forefoot about the longitudinal midtarsal joint axis.
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abductor and adductor hallucis (oblique head) to compress/

Peroneus Longus and Brevis stabilize the first metatarsophalangeal joint during the
Just prior to heel strike, the peroneal musculature propulsive period makes it impossible for the transverse
plays an important role in limiting excessive rearfoot pedis muscle to prevent splaying of the metatarsals, since
inversion, providing invaluable protection against its unstable origin is set into motion.
inversion ankle sprain (133,134). The preparatory Because abductor hallucis has a significant lever
activation of the peroneal musculature is enhanced in arm and angle of approach to both the first ray and
individuals with a prior history of inversion ankle sprain oblique midtarsal joint axes, it functions as an important
(134). During the midstance period, peroneus longus and plantarflexor of the first ray (it assists peroneus longus in
brevis create a pronatory force at the subtalar joint (brevis this action) and supinator about the oblique midtarsal joint
more so than longus) that partially resists the supinatory axis (it is assisted in this action by flexor hallucis longus,
forces generated by the superficial and deep posterior flexor digitorum longus, flexor digitorum brevis, and
compartment musculature. This antagonistic action quadratus plantae).
decelerates the speed of subtalar joint supination and
allows the subtalar joint to return smoothly to its neutral Flexor Hallucis Brevis and
position by early propulsion. Contraction of peroneus Flexor Digitorum Brevis
longus also acts to stabilize articulations of the midfoot as By virtue of its tendinous investment of the
this muscle works synergistically with tibialis posterior to sesamoids, flexor hallucis brevis is a powerful stabilizer
create a compressive force on the tarsals: peroneus longus of the proximal phalanx. This muscle functions with flexor
applies an abductory and posterior force at its insertion hallucis longus to create a compressive force at the first
while tibialis posterior applies an adductory and posterior metatarsophalangeal joint and to maintain the hallux against
force at its insertion. These forces are resolved into a the ground during propulsion. Flexor digitorum brevis has a
straight compressive force that prevents splaying of the similar role in that it functions with flexor digitorum longus
tarsals during late midstance and early propulsion. to compress the metatarsophalangeal joints of the second
The peroneus brevis muscle is also able to create an through fifth rays and allows the lesser digits to maintain
important compressive force as it pulls the fifth metatarsal effective ground contact during the propulsive period. Un
into the cuboid and the cuboid into the calcaneus, thereby like flexor hallucis brevis, flexor digitorum brevis assists in
stabilizing the lateral column. The peroneals continue to producing a strong supinatory force about the oblique mid
contract throughout the majority of the propulsive period, tarsal joint axis during propulsion. Because it lessens the
when peroneus longus plantarflexes the first ray (improving velocity in which the toes dorsiflex during the propulsive
ground contact and allowing for the dorsal-posterior shift period, flexor digitorum brevis protects the plantar fascia
of the first metatarsophalangeal joint’s transverse axis) from high tensile strains by acting as a muscular synergist
while peroneus longus and brevis act together to evert the to the plantar fascia.
locked lateral column (thereby transferring body weight
medially and allowing for a high gear push-off). Because Interossei and the Lumbricals
the peroneals have such short lever arms to the ankle axis, The interossei function during late midstance and
they only slightly assist with ankle plantarflexion during propulsion to maintain transverse plane stability at the
propulsion. second through fifth metatarsophalangeal joints and to
compress the proximal phalanx against the metatarsal heads.
Abductor and Adductor Hallucis The lumbricals have the interesting ability to compress the
The abductor and adductor hallucis muscles function intermediate and distal interphalangeal joints while also
during the propulsive period to stabilize the proximal maintaining the lesser digits against the ground by creating
phalanx of the hallux against the ground. (They maintain a plantarflectory force about the metatarsophalangeal joints
a plantarflectory tension on the first metatarsophalangeal (94). Because the lumbrical tendons pass medially to the
joint.) These muscles are also responsible for transverse metatarsophalangeal joints, they are also able to generate
plane stabilization of the hallux, since they act to create a mild adductory force to resist the abductory shear
equal and opposite rotational components of force on the force associated with ground contact. Since the tendons
proximal phalanx (which resolve into pure compressive of the interossei pass below the transverse axis of the
force). Because of its origin on the proximal phalanx metatarsophalangeal joints, they act as plantarflexors of the
of the hallux and insertion into the distal metatarsal proximal phalanx and, in conjunction with the lumbricals,
heads (refer back to figure 2.116), the transverse head of play an important role in maintaining extensor rigidity of
adductor hallucis (transverse pedis) prevents splaying of the digits during midstance and propulsion.
the metatarsals as it pulls medially on the metatarsal heads
from its stable anchor on the proximal phalanx. Failure of
123
Graphic Summary of the Gait Cycle while Walking
Figure 3.50. Sagittal plane motions.
124
Figure 3.50. Sagittal plane motions, cont.
125
Figure 3.51. Frontal plane motions.
126
Figure 3.51. Frontal plane motions, cont.
127
Figure 3.52. Transverse plane motions. *Although the pelvis in this graph is internally rotated only 2° at heel strike, at
higher speeds of locomotion, the pelvis is often maximally internally rotated at heel strike, which allows for a significant
increase in length of stride.
128
Figure 3.52. Transverse plane motions, cont.
129
Summary of Muscle Function during the Gait Cycle:
Figure 3.53. Muscle function. TFL= tensor fasciae latae.
130
Figure 3.53. Muscle function, cont.
131
Figure 3.53. Muscle function, cont.
132
Summary of Biomechanical Factors Associated with than doubled the metabolic costs of locomotion. Additional
Improved Running Economy: studies have confirmed that increasing shoe weight by
only 2 ounces increases the metabolic cost of running
In closing this chapter on ideal motions during the gait approximately 1% (114,115). These findings explain why
cycle, it seems fitting to discuss the various biomechanical endurance runners with small feet are more efficient than
factors responsible for maximizing speed and metabolic their large-footed rivals (63).
efficiency (which are often mutually exclusive since 3) Running efficiency is associated with low
sprinters make terrible long distance runners, and long vertical oscillation of the body’s center of mass. In an
distance runners are often terrible sprinters). Even though interesting study of efficiency in middle and long distance
natural selection has relentlessly modified each person’s runners competing in a 5 km race, Miayashita et al. (116)
musculoskeletal system for over 7 million years, there determined that the center of mass in the best runners
is significant individual variation in running skill: some moved with a vertical displacement of only 6 cm, while the
people are fast and tire easily while others are slow but can less efficient runners averaged vertical displacements of 10
run extremely long distances without fatiguing. Because cm. The length of stride between fast and slow runners was
there are dozens of anthropomorphic and movement also different in that the average stride length for a good
variables capable of affecting performance, the following runner was 1.77 m compared to 1.60 m for the less skilled
section reviews kinetic/kinematic traits responsible for runners. The authors determined that the good runners ran
success in endurance running, followed by a list of factors 5,000 meters in 2,825 steps while the poor runners required
associated with successful sprinting. 3,125 steps. The added work associated with lifting
the center of mass the additional 4 cm with each stride
Endurance Running produced an increased workload roughly the equivalent
1) According to Anderson (18), the best male long to the cost of running up a 50-story building. While this
distance runners tend to be slightly shorter than average seems impressive, the notion that increasing stride length
while females tend to be slightly taller than average. will automatically improve efficiency may be flawed.
Females tend to be ectomorphic while males tend to be Because the less skilled runners generated less force with
ectomesomorphic. Both males and females present with their shorter strides, the metabolic expense associated with
lower percentages of body fat. Sawyer et al. (110) note long versus short strides is difficult to compare. Remember
that sub-Saharan Africans (e.g., Kenyans and Ethiopians) that every runner selects a stride length that maximizes
possess increased limb lengths relative to torso volume, efficiency and any attempt to modify an individual’s freely
which markedly improves efficiency because a smaller chosen stride length invariably increases the metabolic cost
torso is easier to move long distances. Although longer of locomotion (17).
limbs relative to torso volume improve efficiency while 4) Efficient runners plantarflex their ankles through
running, the benefits associated with longer tibiae are less a smaller range during propulsion (117) and this reduced
clear. Despite the fact that walking efficiency improves movement occurs at a faster velocity (18). While studying
with longer legs (111), anthropomorphic evaluation of kinematic differences during propulsion, Cavanagh et al.
tibial lengths in runners provides conflicting results: a study (117) determined that efficient runners plantarflex their
of Olympic level male runners revealed that long distance ankles 10° less than inefficient runners. In a separate study,
runners were short-legged, middle distance runners were Williams and Cavanagh (63) evaluated 3 groups of runners
long-legged and sprinters were short-legged (112). In a based on economy and determined that the best runners
detailed study comparing metabolic efficiency in runners again plantarflex their ankles through a smaller range of
of different abilities, Williams and Cavanagh (63) found motion during propulsion. The decreased range of motion
no connection between leg length and efficiency when present in the ankle coupled with the increased angular
running. excursion velocity noted by Anderson (18) may represent
2) The best long distance runners possess leg a kinematic marker associated with improved storage and
morphology that distributes mass closer to the hip joint. return of energy, as isometric contractions produce smaller
Runners possessing muscular hips with relatively thin changes in movement with more rapid joint excursions.
lower legs are more efficient because accelerating and 5) Running efficiency is associated with reduced
decelerating the distal segments contributes greatly to the angular excursions of the arms and wrists. Williams and
metabolic cost of locomotion. Since the distal segments Cavanagh (63) correlated running efficiency with decreased
have long levers to the proximal muscles, even a slight wrist excursions while Anderson and Tseh (119) confirm
increase in weight applied to the foot will greatly reduce that the most economical runners present with the smallest
efficiency. To prove this, Martin et al. (113) measured arm movements. Although arm motions lessen strain on
oxygen consumption before and after adding weights the back musculature (60), exaggerated movements require
to either the foot or thigh of recreational runners. These muscular effort to initiate and dampen, and are therefore
authors determined that adding weight to the feet more associated with reduced metabolic efficiency.
133
Sprinting
1) Several studies (120,121) reveal that sprinters Conversely, sprinters possess toes that are almost 1 cm
have significantly longer fascicles in their gastrocnemius longer than non-sprinter controls. While counterintuitive,
muscles compared with non-sprinters. The longer fibers the 25% shorter lever arm allows the Achilles to effectively
allow the calf muscle to operate more efficiently through plantarflex the ankle with little change in length occurring
a larger a range of motion, improving the force-velocity in the gastrocnemius and soleus (Fig. 3.54). The reduced
ratio. Another possibility is that longer fascicles are more lever arm may decrease mechanical efficiency of the
efficient at storing and returning energy in the actomyosin Achilles tendon, but it allows the gastrocnemius and soleus
cross-bridges because they can stretch farther than shorter to move the ankle with a nearly isometric contraction.
muscle fibers. The longer fascicles may be inherited but On the opposite side of the fulcrum, the longer toes
more likely result from training as muscles rapidly adapt to allow for greater force production in the forefoot because
high intensity training by increasing fascicle length (122). the increased toe lengths provide the digital flexors with
2) Although peroneus brevis is important with significantly longer lever arms that allow for a more
sprinting because it everts the rearfoot allowing for a high powerful push-off. Even though the added metabolic cost
gear push-off, EMG analysis of a wide range of muscles of accelerating and decelerating the longer, heavier toes
utilized while running confirms that the majority of force would lessen efficiency while walking and running long
associated with forward propulsion is produced by hip distances (which is why evolution has favored shorter toe
flexion and knee extension (123). This is consistent with lengths), the longer toes provide increased force production
research demonstrating the distal muscles act as springs to during propulsion, thereby allowing the elite sprinter to run
store and return energy while the proximal muscles act as at the fastest speed possible. The combination of a short
force generators, muscularly accelerating and decelerating Achilles lever arm coupled with long toes is also found in
the lower extremity with each step (68). nature; e.g., cheetahs, which are capable of sprint speeds
3) In an interesting study of foot morphology in exceeding 70 mph, have shorter posterior calcanei and
sprinters, Piazza (124) determined the distance from the longer toes than lions. Although it takes millions of years,
posterior calcaneus to the center the ankle is 25% shorter natural selection eventually matches form to function with
in elite sprinters compared with the non-sprinter controls. the simplest possible design.
3.54. Because the distance from the Achilles tendon is 25% longer in non-sprinters (compare A and B), the
gastrocnemius and soleus must move through larger ranges of motion to plantarflex the ankle (compare C and D).
Notice the toes of sprinters are 1 cm longer than non-sprinters.
134
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TABLE of CONTENTS

Chapter One The Evolution of Bipedality 1

Chapter Two Structural and Functional Anatomy 23

Chapter Three Ideal Motions During the Gait Cycle 87

STANCE PHASE MOTIONS 94

SWING PHASE MOTIONS 117

SUMMARY OF MUSCLE FUNCTION DURING THE GAIT CYCLE 120

GRAPHIC SUMMARY OF THE GAIT CYCLE 124

Chapter Four Abnormal Motion During the Gait Cycle 139

DEVELOPMENTAL TRENDS IN LOWER EXTREMITY ALIGNMENT 140

Transverse Plane Alignment 140

Frontal Plane Alignment 147

DEVELOPMENT OF THE MEDIAL LONGITUDINAL ARCH 151

ALIGNMENT OF THE METATARSAL HEADS 172

VARIATION IN METATARSAL LENGTH 180

LIMB LENGTH DISCREPANCY 184

EXCESSIVE MOBILITY 189

Tibiofemoral Joint 189

Ankle Joint 190

Subtalar Joint 191

Midtarsal Joint 191

Hip Joint 195

Tibiofemoral Joint 197

Ankle Joint 197

Subtalar Joint 199

First Metatarsophalangeal Joint 200

TREATMENT of LIMITED MOTION 201

Restricted Motion Resulting From Muscular Contraction 202

Muscle length assessment techniques 205

Home stretches 212

Restricted Motion Resulting From Osseous Block 219

Restricted Motion Resulting From Joint Stiffness 221

Joint manipulations/mobilizations 223

NEUROMOTOR COORDINATION and STRENGTH 237

Muscle Testing 244

Rehabilitative Exercises 251

Chapter Five Biomechanical Examination 271

SUPINE EXAMINATION 271

PRONE EXAMINATION 277

STANDING EXAMINATION 280

DYNAMIC EVALUATION 293

Chapter Six Foot Orthotics 301

MECHANISM of ACTION 301

ORTHOTIC CASTING TECHNIQUES 304

LABORATORY PREPARATION and ORTHOTIC FABRICATION 309

Modification of the Positive Model 309

ORTHOTIC POSTING 311

ORTHOTIC SHELLS 316

ORTHOTIC ADDITIONS 318

IN-OFFICE FABRICATION TECHNIQUES 323

ORTHOTIC DISPENSING 324

ORTHOTIC PROBLEM-SOLVING 324

Chapter Seven Shoe Gear 333

HISTORY OF FOOTWEAR 333

MODERN SHOES 335

MINIMALIST SNEAKERS 341

Chapter Eight Treatment Protocols 345

ACHILLES TENDINITIS 346

METATARSALGIA and METATARSAL STRESS FRACTURES 349

PLANTAR PLATE RUPTURE 350

21. K ivell T, Schmitt D. Independent evolution of knuckle-

prevent impingement by drawing the medial menis cus