Posture and locomotion in man ,evolution and

development of the locomotor apparatus in

man.Gait and gait cycles
• This chapter discusses the concepts of posture, locomotion, and voluntary movement.
Many of the ideas about human posture and locomotion are based upon observations on
quadrupeds, although some information comes from experiments on other primates and
more has been gained from a few experiments on human subjects. The human, except as
an infant, is a biped and differs from the quadrupeds in posture and in the manner of
locomotion. Tonic or static posture in man is the maintenance of an upright vertical
posture above the two supporting limbs. Locomotion is the movement of the body, by a
coordinated series of muscle activities, in a particular direction while the relation of the
body as a whole to the ground is maintained. The process of locomotion in man, as
exemplified in walking, consists in raising one leg by flexion and producing extensor thrust
in the other so that the body falls forward. Control of muscle movements can be exerted at
all levels from the cortex to the spinal cord at the segment of origin of the muscle. The
cortex is responsible for the initiation of voluntary movement. The conscious initiation of
movement is a process originating in the formulation of the will to move in the frontal
cortex. Except in the case of relatively simple movements such as rapid movements
involving very few muscles, impulses pass from the frontal cortex to the basal ganglia. The
principal muscle involved in a movement is the agonist, muscles assisting its action are
synergist, and those giving the opposite movement are antagonist.
• A number of terms should be explained before discussing these concepts. Muscles of the skeletal
system are usually described according to their function as flexor or extensor. A flexor muscle
contracts to produce bending at a joint – the biceps muscle, for example, bending the arm at the
elbow joint – whilst an extensor muscle has the opposite action of straightening the line of bones
at a joint -as the triceps muscle straightens the arm. Slightly different terms are used in the case of
the muscles of mastication which may be elevator or depressor to the mandible, or may move it
forward (protrusor) or back (retrusor). All these movements involve changes in the plane of one
muscle in relation to another; rotational or twisting movements also occur, as in the cervical spine
when the head is turned. Indeed, because of the position of insertion of muscles into the bones,
rotational movements may be included in the flexor or extensor movement. The principal muscle
involved in a movement is the agonist (or prime mover, or sometimes, protagonist); muscles
assisting its action are synergist, and those giving the opposite movement are antagonist.
• The motor neurone passing from the central nervous system (spinal cord or the nuclei of the
cranial nerves) to the muscle is often termed the lower motor neurone, a convenient clinical term
to distinguish these neurones from those whose cell bodies and processes are entirely in the spinal
cord or brain, damage to which causes different clinical effects. The lower motor neurone, as the
nerve supply to the motor unit, is the only pathway by which impulses from any source in the
central nervous system can reach the motor unit. It is therefore sometimes called the final common
pathway for motor activity.
• Several reflexes which operate at a spinal or brainstem level have been thought to be implicated in
posture and locomotion. These are:
• (i) The stretch reflex. Stretching of a muscle stimulates the spindle stretch receptors and causes
contraction of the muscle by a simple reflex usually involving only two neurones directly.
• (ii) The tendon jerk. This is the same as the stretch reflex, but is initiated by tapping the muscle
tendon to produce the stretching of the muscle.
• (iii) The flexor response. In response to a noxious stimulus, the limb is flexed to remove it from the
point of stimulation. Although the reflex is seen with noxious stimuli, it is argued that, in
locomotion, the touch and pressure stimuli to a supporting limb may initiate flexion.
• (iv) The crossed extensor reflex. This response may be associated with the flexor response,
particularly if the stimuli are intense. Flexion of the stimulated limb is accompanied by extension of
the limb on the other side, the contralateral limb.
• (v) The scratch reflex. Repetitive stimulation of touch receptors causes a pattern of limb movement
involving alternate extension and flexion. Since locomotion is a pattern of alternate extension and
flexion of the limbs, analogies have been sought between the scratch reflex and locomotion.
• These simple reflexes may be involved either by themselves or as part of a larger pattern. The more
complex and more specific reflexes involved in posture and locomotion will be dealt with in the
context of the whole process.
• Posture
• Most of our understanding of neurophysiology comes from animal experiments, of which the
vast majority have been carried out on quadrupeds. Many of the ideas about human posture
and locomotion, therefore, are based upon observations on quadrupeds, although some
information comes from experiments on other primates and more has been gained from a few
experiments on human subjects. The human, except as an infant, is a biped and differs from
the quadrupeds in posture and in the manner of locomotion.
• The most important single difference is that the quadruped has four supporting limbs, and the
vertebral column with its muscles and ligaments is unimportant, whilst man has only two
supporting limbs and depends upon the rigidity of the vertebral column to maintain the
upright posture. Reflexes relating activity of front and hind limbs have relevance to adult man
only when equilibrium has been lost completely.
• Posture may be considered to involve three processes. The first may be termed tonic, or static,
posture; the second is the correction of small changes in body position, and is given the name
of ‘righting’, or righting reflexes; and the third is the maintenance of a general postural position
during movement – statokinetic reflexes. Another term sometimes used is ‘attitude’: the spatial
relationship of the body structures to each other (Fig. 14.1). This is a part of all three aspects of
posture but is probably most important in the statokinetic responses.
• Static posture (stance)
• Tonic or static posture in man is the maintenance of an upright vertical posture above the two
supporting limbs. It may be achieved by the fixation of joints by simultaneous contraction of the
extensor and flexor muscles. This is true in the lower limbs, as it is in all four limbs of a quadruped: but
in addition the vertebral column must be fixed both intrinsically and extrinsically so that pelvis,
abdomen and thorax are positioned above the pelvic basin. This combination of activities is referred
to as the positive supporting action, and it seems to be initiated by proprioceptive impulses from the
small interosseous muscles of the feet. It is replaced by the negative supporting action when flexion of
a limb causes excitation of proprioceptive nerves from the plantar or volar muscles (on the sole or
upper side of the foot). Although the positive supporting action seems necessary for the adoption of
stance, it is probable that stance is maintained with very little effort in the muscles except when an
unusual stance is adopted. Even then the body adopts a position in which the centre of gravity is
above the centre of ground contact. Thus wearing high-heeled shoes which tilt the plane of the feet
upwards from toe to heel causes the shoulders to be held back to counteract the forward thrust of the
pelvis induced by the leg position (Fig. 14.2). Animal experiments demonstrate that a truly normal
stance cannot be established or maintained unless the cortex and basal ganglia are intact. Spinal
pathways are insufficient of themselves for standing. However, transection in the upper part of the
pons results in decerebrate rigidity which can permit standing. Stance with normal muscle tone
requires at least the forepart of the midbrain, and the positive and negative supporting reactions need
the basal ganglia and the cortex to be intact.
 Evolution and development of locomotor
apparatus in man
• The arrangement of striated muscle in modern humans conforms to
the basic plan seen in all pronograde quadrupedal vertebrates and
mammals (that is, all vertebrates and mammals that assume a
horizontal and four-legged posture). The primates (the order of
mammals to which humans belong) inherited the primitive
quadrupedal stance and locomotion, but since their appearance in
the Late Cretaceous Period some 65 million years ago, several
groups have modified their locomotor system to concentrate on the
use of the arms for propulsion through the trees. The most-extreme
expression of that skeletal adaptation in living primates is seen in the
modern gibbon family. Their forelimbs are relatively elongated; they
hold their trunk erect; and, for the short periods that they spend on
the ground, they walk only on their hind limbs (in a bipedal fashion).
• Modern humans are most closely related to the
living great apes: the chimpanzee, the gorilla,
and the orangutan. The human’s most-distant
relative in the group, the orangutan, has a
locomotor system that is adapted for moving
among the vertical tree trunks of the Asian
rainforests. It grips the trunks equally well with
both fore and hind limbs and was at one time
aptly called quadrumanal, or “four-handed.”
• There is little direct fossil evidence about the common ancestor of modern humans, chimpanzees, and gorillas, so inferences about its habitat and locomotion must be made. The ancestor
was most likely a relatively generalized tree-dwelling animal that could walk quadrupedally along branches as well as climb between them. From such an ancestor, two locomotor trends
were apparently derived. In one, which led to the gorillas and the chimpanzees, the forelimbs became elongated, so when those modern animals come to the ground, they support their
trunks by placing the knuckles of their outstretched forelimbs on the ground. The second trend involved shortening the trunk, relocating the shoulder blades, and, most important, steadily
increasing the emphasis on hind-limb support and truncal erectness. In other words, that trend saw the achievement of an upright bipedal, or orthograde, posture instead of a quadrupedal,
or pronograde, one. The upright posture probably was quite well established by 3 million to 3.5 million years ago, as evidenced both by the form of the limb bones and by the preserved
footprints of early hominins found from that time.
• human being (Homo sapiens)Human beings (Homo sapiens) are anatomically similar and related to the great apes but are distinguished by a more highly developed brain and a resultant
capacity for articulate speech and abstract reasoning.
• Encyclopædia Britannica, Inc.
• Changes in the muscles of the lower limb
• The major muscular changes directly associated with the shift to bipedal locomotion are seen in the lower limb. The obvious skeletal changes are in the length of the hind limb, the
development of the heel, and the change in the shape of the knee joint so that its surface is flat and not evenly rounded. The hind limbs of apes are relatively short for their body size
compared with modern human proportions.
• skeletons of humans and gorillas comparedThe skeletal structure of a human being (left) and of a gorilla (right). Several differences allow the human to walk erect on two legs with a striding
gait rather than move in a knuckle-walking fashion like the gorilla. In the pelvis these differences include shorter ischia, a broader sacrum, and broader, curved-in ilia with a lower iliac crest.
In the legs the femurs (thighbones) are relatively long and are set farther apart at the hips than they are at the knees.
• Encyclopædia Britannica, Inc.
• The changes that occurred in the bones of the pelvis are not all directly related to the shift in locomotion, but they are a consequence of it. Bipedality, by freeing the hands from primary
involvement with support and locomotion, enabled the development of manual dexterity and thus the manufacture and use of tools, which has been linked to the development in human
ancestors of language and other intellectual capacities. The result is a substantially enlarged brain. Large brains clearly affect the form of the skull and thus the musculature of the head and
neck. A larger brain also has a direct effect on the pelvis because of the need for a wide pelvic inlet and outlet for the birth of relatively large-brained young. A larger pelvic cavity means that
the hip joints have to be farther apart. Consequently, the hip joints are subjected to considerable forces when weight is taken on one leg, as it has to be in walking and running.
• To counteract that, the muscles (gluteus minimus and gluteus medius) that are used by the chimpanzee to push the leg back (hip extensors) have shifted in modern humans in relation to
the hip joint so that they now act as abductors to balance the trunk on the weight-bearing leg during walking. Part of a third climbing muscle (gluteus maximus) also assists in abduction as
well as in maintaining the knee in extension during weight bearing. The gluteal muscles are also responsible for much of the rotation of the hip that has to accompany walking. When the
right leg is swung forward and the right foot touches the ground, the hip joint of the same side externally rotates, whereas that of the opposite side undergoes a similar amount of internal
rotation. Both of those movements are made possible by rearrangements of the muscles crossing the hip.
• The bones of the trunk and the lower limb are so arranged in modern humans that standing upright requires a minimum of muscle activity. Some muscles, however, are essential to
maintaining balance, and the extensors of the knee have been rearranged and realigned, as have the muscles of the calf.
• The foot is often but erroneously considered to be a poor relation of the hand. Although the toes in modern humans are normally incapable of useful independent movement, the flexor
muscles of the big toe are developed to provide the final push off in the walking cycle. Muscles of all three compartments of the modern human lower leg contribute to making the foot a
stable platform, which nonetheless can adapt to walking over rough and sloping ground.
• Changes in the muscles of the lower limb
• The major muscular changes directly associated
with the shift to bipedal locomotion are seen in the
lower limb. The obvious skeletal changes are in the
length of the hind limb, the development of the
heel, and the change in the shape of the knee joint
so that its surface is flat and not evenly rounded.
The hind limbs of apes are relatively short for their
body size compared with modern human
proportions.
• The changes that occurred in the bones of the pelvis are not all directly
related to the shift in locomotion, but they are a consequence of it.
Bipedality, by freeing the hands from primary involvement with support and
locomotion, enabled the development of manual dexterity and thus the
manufacture and use of tools, which has been linked to the development in
human ancestors of language and other intellectual capacities. The result is
a substantially enlarged brain. Large brains clearly affect the form of the
skull and thus the musculature of the head and neck. A larger brain also has
a direct effect on the pelvis because of the need for a wide pelvic inlet and
outlet for the birth of relatively large-brained young. A larger pelvic cavity
means that the hip joints have to be farther apart. Consequently, the hip
joints are subjected to considerable forces when weight is taken on one leg,
as it has to be in walking and running.
• To counteract that, the muscles (gluteus minimus and gluteus
medius) that are used by the chimpanzee to push the leg back
(hip extensors) have shifted in modern humans in relation to the
hip joint so that they now act as abductors to balance the trunk
on the weight-bearing leg during walking. Part of a third
climbing muscle (gluteus maximus) also assists in abduction as
well as in maintaining the knee in extension during weight
bearing. The gluteal muscles are also responsible for much of the
rotation of the hip that has to accompany walking. When the
right leg is swung forward and the right foot touches the ground,
the hip joint of the same side externally rotates, whereas that of
the opposite side undergoes a similar amount of internal
rotation.
• Both of those movements are made possible by rearrangements of the
muscles crossing the hip.
• The bones of the trunk and the lower limb are so arranged in modern
humans that standing upright requires a minimum of muscle activity.
Some muscles, however, are essential to maintaining balance, and the
extensors of the knee have been rearranged and realigned, as have the
muscles of the calf.
• The foot is often but erroneously considered to be a poor relation of the
hand. Although the toes in modern humans are normally incapable of
useful independent movement, the flexor muscles of the big toe are
developed to provide the final push off in the walking cycle. Muscles of
all three compartments of the modern human lower leg contribute to
making the foot a stable platform, which nonetheless can adapt to
walking over rough and sloping ground.
• Changes in the muscles of the upper limb
• The human upper limb has retained an overall generalized
structure, with its details adapted to upright existence.
Among the primitive features that persist are the clavicle, or
collarbone, which still functions as part of the shoulder; the
ability to twist one of the forearm bones (the radius)
around the other (the ulna) so that the palm is turned
forward or backward, a process called pronation and
supination; and a full complement of five digits in the hand.
• Pronation and supination of the forearm, which allows the
palm of the hand to rotate 180°, is not peculiar to humans.
That movement depends upon the possession of both a
small disk in the wrist joint and an arrangement of the
muscles such that they can rotate the radius to and fro.
Both the disk and the muscle arrangement are present in
the great apes.
• In quadrupedal animals the thorax (chest) is suspended
between the shoulder blades by a muscular hammock
formed by the serratus anterior muscle. In upright sitting
and standing, however, the shoulder girdle is suspended
from the trunk.
• The scapula, or shoulder blade, floats over the
thoracic surface by reason of the arrangement
of the fibres of the serratus anterior muscle
and the support against gravity that is
provided by the trapezius, rhomboid, and
levator scapulae muscles. When the arms are
required to push forward against an object at
shoulder level, their action is reminiscent of
quadrupedal support
• The change in shape of the chest to emphasize breadth rather
than depth altered the relation of muscles in the shoulder
region, with an increase in size of the latissimus dorsi muscle
and the pectoralis major muscle. The human pectoralis minor
muscle has forsaken its attachment to the humerus, the long
bone of the upper arm, and presumably derives some stability
from attaching to the coracoid process, a projection from the
scapula, instead of gliding over it.
• The hand of a chimpanzee is dexterous, but the proportions of
the digits and the rearrangement and supplementation of
muscles are the major reasons for the greater manipulative
ability of the hand of a modern human.
• Most of those changes are concentrated on the
thumb. For example, modern humans are the
only living hominids to have a separate long
thumb flexor, and the short muscle that swings
the thumb over toward the palm is particularly
well developed in humans. That contributes to
the movement of opposition that is crucial for
the so-called precision grip—i.e., the bringing
together of the tips of the thumb and forefinger.
• Changes in the muscles of the head and neck
• The muscle group of the head and neck is most directly influenced by
the change to an upright posture. That group comprises the muscles of
the back (nape) and side of the neck. Posture is not the only influence on
those muscles, for the reduction in the size of the jaws in modern
humans also contributes to the observed muscular differences.
Generally, those involve the reduction in bulk of nuchal (nape) muscles.
In the upright posture the head is more evenly balanced on the top of
the vertebral column, so less muscle force is needed, whereas in a
pronograde animal with large jaws the considerable torque developed at
the base of the skull must be resisted by muscle force. The poise of the
human head does pose other problems, and the detailed attachment
and role of some neck muscles (e.g., sternocleidomastoid) are different
in humans from those in apes
Muscles of facial expression
• Changes in the muscles of the trunk
• The consequences of an upright posture for the support of
both the thoracic and the abdominal viscera are profound,
but the muscular modifications in the trunk are few.
Whereas in pronograde animals the abdominal viscera are
supported by the ventral abdominal wall, in the orthograde
posture most support comes from the pelvis. That inevitably
places greater strain on the passage through the muscles of
the anterior abdominal wall, the inguinal canal, which
marks the route taken by the descending testicle in the
male. Weakness in the canal can result in herniation.
• ifferences are also seen in the musculature, the levator ani, that supports
the floor of the pelvis and that also controls the passage of feces. The loss of
the tail in all apes has led to a major rearrangement of that muscle. There is
more overlap and fusion between the various parts of the levator ani in
modern humans than in apes, and the muscular sling that comprises the
puborectalis in humans is more-substantial than in apes.
• The muscular compression of the abdomen and the thorax that
accompanies upright posture aids the vertebral column in supporting the
body and in providing a firm base for upper-limb action. Anteroposterior
(fore-and-aft) stability of the trunk is achieved by balancing the flexing
action of gravity against back muscles that act to extend the spine. Lateral
stability is enhanced by the augmented leverage provided to the spinal
muscles by the broadening of the chest.
 Gait and gait cycles
• Gait analysis is the study of human locomotion. In order
to analyze and quantify how someone walks, it is
necessary to isolate the shortest, unique, repeatable task
during gait. This task is called the gait cycle. A single gait
cycle can be measured from any gait event to the same
subsequent event on the same foot, but the conventional
tacit model considers gait cycle is measured from one
foot strike to the subsequent foot strike of the same foot.
• Quantifying aspects of the gait cycle, such as time and
spatial measures, allow for analysis of gait symmetry,
variability and quality.
• Two Primary Phases of the Gait Cycle
• The gait cycle can be broken down into two primary phases,
the stance and swing phases, which alternate for each lower limb.
• Stance phase: Consists of the entire time that a foot is on the ground.
• Swing phase: Consists of the entire time that the foot is in the air.
• The observation of both the spatial and temporal characteristics of the two
lower limbs makes it possible to introduce complementary phases. When
both members are simultaneously in the stance phase, we speak of bipedal
support or double support (2 times 10%); when only one is in the support
phase, one speaks of unipedal support or single support (40%), the second
then being in oscillating phase.
• The most accurate approach is based on the functional interpretation of
events and identifies eight phases that capture three tasks related to
walking. We then distinguish (Perry & Burnfield, 2010):
• 1.Weight acceptance (0-12%):
• The objectives of weight acceptance are to stabilize the
limb, absorb shock and preserve the progression of the
body. This phase can be broken down further into initial
contact and loading response. Initial contact consists of the
first 3% of the gait cycle. In typical gait, the heel strikes the
ground and initiates the rotation over the heel to foot flat to
preserve progression. This motion is the first rocker of the
gait cycle. Loading response goes from 3-12% of the gait
cycle. In this portion, the knee flexes slightly in order to
absorb shock as the foot falls flat on the ground, stabilizing
in advance of single limb support.
• 2.Single limb support (12-50%):
• Single limb support involves progression of the body over the
foot and weight-bearing stability. The first sub-phase of single
limb support is midstance, which is seen during the 12-31% of
the gait cycle. During midstance, the shank rotates forward
over the supporting foot, creating the second rocker motion of
the cycle. This maintains the forward progression of gait. The
second stage of single support is terminal stance which goes
from 31-50% of the gait cycle. During terminal stance, the
center of mass advances out in front of the supporting foot.
The heel raises of the ground as you roll onto the ball of the
foot, creating the third rocker motion of the cycle.
• 3.Swing phase (50-100%):
• The objectives of the swing phase of gait
• Foot clearance over the ground
• Forward swing of the limb
• Preparation of limb for stance
• The swing phase can be broken down into 4 sub-phases.
• 1.Pre-swing takes place during 50-62% of the gait cycle. Pre-swing is the transition phase
between stance and swing, in which the foot is pushed and lifted off of the ground.
• 2.Initial swing goes from 62-75% of the gait cycle. During initial swing, the hip, knee,
and ankle are flexed to begin advancement of the limb forward and create clearance of
the foot over the ground.
• 3.Mid-swing goes from 75-87% of the gait cycle. During mid-swing, limb advancement
continues and the thigh reaches its peak advancement.
• 4.Terminal swing is the final phase of the gait cycle going from 87-100% of the cycle.
During terminal swing, the final advancement of the shank takes place and the foot is
positioned for initial foot contact to start the next gait cycle.