Lecture Notes on Biomechanics and Syllabus for Gross Anatomy I

Prepared by Dr Subhash C Dubal, Professor of Anatomy

DEPT. OF ANATOMY COLLEGE OF VETERINARY SCI. & A.H., ANAND AGRICULTURAL UNIVERCITY, ANAND (GUJARAT) INDIA

"It is almost a miracle that modern teaching methods have not yet entirely strangled the holy curiosity of inquiry; for what this delicate little plant needs more than anything, besides stimulation, is freedom ..Einstein (14 March 1879 18 April 1955)

SYLLABUS

For a superficial observer, scientific truth is beyond the possibility of doubt ; the logic of science is infallible, and if the scientists are sometimes mistaken, this is only from their mistaking its rules. (H. Poincare, 1921)

Preface
The days when medicine was left to the physicians, and engineering was left to the engineers, seem to have passed us by (Wait and Fine, 2007). Now biomechanics is also under the preview of biologists. It is a curious feature of science that it is widely assumed

that the newer the subject the more difficult it must be, and conversely the older the subject the easier it is assumed to be (Bondi. 1968) This little lecture note is meant for under graduate students of First Semester (Gross Anatomy) of Veterinary Science & A.H. (VCI). Biomechanics is an extremely vital subject in understanding the biological and physical performance of cells (hence the animal body). One of the most important discoveries in biological science in general and molecular and cell biology in particular is that of mechanotransduction. The gene expression (i.e., performance of various functions) is under the action of mechanical stimuli. The cells express different genes in response to extremely small changes in their mechanical environments. Thus the importance of biomechanics in veterinary sciences is far reaching. Mathematics is a science as well as a tool to explore the mechanism by which the nature expresses its beauty. Mathematics, is the essential key to the science without which nature cannot be properly understood. The book which is the universe is written in mathematical language (Galileo, 1564 1642). However, hardly 10% of biologists (particularly veterinary anatomists) desire to enjoy the nature through mathematics. Anatomists try to avoid the mathematics. They have tones of morphological data, but could build comprehensive predictions. I therefore have given the mathematical expressions in terms of description of the conclusions (which is hardly 50% of the total that can be seen by mathematical expressions). The lecture note mainly concerns with the elementary biomechanics of locomotor system and deformation and fracture of bone (rupture of tendon and ligament). There is also a brief description of biomechanics of lubrication of synovial joints. An isolated fact can be observed by all eyes; by those of the ordinary person as well as of the wise. But it is the true physicist (read Anatomist) alone who may see the bond which unites several facts among which the relationship is important though obscure. Facts are sterile until there are minds capable of choosing between them and discerning those which conceal something and recognizing that which is concealed; minds which under the bare fact see the soul of the fact ( Poincare, 1908). It appears that these statements are written for anatomists learning the biomechanics. Biomechanics is intellectually stimulating and challenging, and vitally important discipline. The final achievement of biomechanics is to contribute to the improvement of health care delivery to the animals. This goal deserves our best effort. I hope the student will find this little lecture note on biomechanics helpful for their academic curriculum and it will increase some interest for deeper persuasion of the vast subject of biomechanics. Criticisms and suggestions are most welcome. They will improve the quality and quantity of the notes. I am thankful to all for their materials used in this lecture notes. Dr. Subhash C Dubal Veterinary College, Anand (Gujarat) India 02.03.2012

 Biomechanics aims to explain the mechanics of life and living. From molecules to organisms, everything must obey the laws of mechanics Y.C.Fung (1990) the Father of modern Biomechanics .

I. INTRODUCTION

1.1. Cells are the structural and functional unit of living organisms. The cells are physically neither the solid nor the fluid (liquid and gas). The body cells are constantly under the action of forces (for example the universal gravitational force) and are, in turn, under the influence of mechanical stimuli. The mechanical stimuli affect the structures and functions of the cells (even the gene expressions). Mechanobiology is the branch of biology which deals with the structure and functions of cells (animal body) controlled by mechanical stimuli. The mechanical stimuli act upon the cells in accordance with laws of mechanics. Biomechanics is a branch of mechanobiology that deals with the application of laws of mechanics to the living organisms. Biorheology is also a branch of mechanobiology which deals with the deformation and flow of materials. BIOMECHANICS CELLS BIORHEOLOGY MECHANOBIOLOGY Besides the traditional (classical) branches of biomechanics (Flow chart 1), the biomechanics is more than the simply application of laws of mechanics. The biomechanics is vibrant field- one with great promise. The main branches (research areas) of biomechanics (slightly modified from Humphrey, 2003 and Humphrey and Delang, 2004) are as follows: 1. 2. 3. 4. 5. 6. 7. 8. 9. Molecular and Cell Biomechanics Developmental Biomechanics Biomechanics of Injury and Rehabilitation Biomechanics of Functional Tissue Engineering Biomechanics of Tissues, Organs and Systems Biomechanics of Solids-Fluid Interactions Thermobiomechanics Electrobiomechanics Chemobiomechanics

Molecular and Cell Biomechanics concerns with study of basic processes namely cell adhesions, contraction, division, migration, spreading, phygocytosis, cellular apoptosis (programmed cell death), synthesis and degradation of matrix (ground substance), production of growth regulatory ,olecules, cytokines, cell surface receptors and gene expression under the influence of mechanical stimuli. Developmental Biomechanics deals with study of effects of mechanical forces on the development of tissues, organs and the organism as a whole. Injury refers to failure and damage of biosystems as in broken bones, torn muscles, ligaments, and tendons, and organ impairment. Injury studies thus include evaluation of tissue properties. They also include studies of accidents and the design of protective devices. 4

Rehabilitation refers to the recovery from injury and disease. Rehabilitation thus includes all applications of mechanics in the health care industries encompassing such areas as design of corrective and assisting devices, development of implants, design of diagnostic devices, and tissue healing mechanics The tissue engineering corresponds to the concept of substitute tissue developed in vitro, from bioresorbable or non bioresorbable scaffolds and from cells harvested in a physiologic mechanical environment such as from cartilage, bone and vessels. At the same time, the problems of cell grafting in tissue repair and especially the use of stem cells have led to new therapeutic fields. Definition or research areas of other branches of biomechanics are self explanatory. 1.2. The mechanobiology involves four principal processes: 1. Mechanical coupling which generally implies the transformation of the applied force into a force which is detectable by the cells or the induction of a physical phenomenon 2. Mechanotransduction which concerns with the transmission of a mechanical stimulus to a physiological phenomenon. 3. Signal transduction, i.e. the conversion of the mechanical signal into intracellular physiological signals. 4. Cellular response: it deals with the regulation of a gene, release of autocrine or paracrine factors, expression of specific receptors In brief, the generated by: Cell membrane integrin-ECM (extra-cellual matrix) adhesions, Stretching of cell membrane, or /and Shear stress is transmitted to components of CSK (cytoskeleton) with the help of link molecules, which in turn transmit the force to the molecules of nuclear membrane. Thorough theses molecules, the force will be transmitted to the molecules of chromosomes and there will be uncoiling the particular DNA of the desired gene(s). This whole process will lead to gene expression.

Research in mechanobiology leads to medical and therapeutic applications.

1.3. Biomechanics: It is a branch of mechanobiology which deals with the application of laws of mechanics to the living organisms. The biomechanics is divided into the following branches (Figure 1):

Figure 1.1: Flow chart showing the branches of biomechanics

1.4. Study of Locomotion requires knowledge of study of biostatics and biodynamics. 1. Biostatics: It is that branch of mechanics, which deals with study of forces and their effects, while acting upon the animal body (or part of body) at rest. It is assumed that the body is acted upon by ` FORCES` which balance one another. The primary conceptions of biostatics are forces and the body upon which they act. 2. Biodynamics: It is that branch of biomechanics, which deals with study of forces and their effects, while acting upon the animal body (or part of body) in motion. It is divided into two major branches. i. Biokinemetics: It is the branch of biodynamics, which deals with the study motion of animal body regardless of the causes, the causes of motion (forces) are not studied. The measurements of linear motion are displacement (distance), velocity (speed) and acceleration, while the corresponding measurements of curvilinear motion are angular displacement, angular velocity and angular acceleration, ii. Biokenatics: It is that branch of biodynamics, which deals with the study the causes (forces) of motion of animal body. The motion of animal body and the causes of motion (forces) are studied. It is now widely recognized that biomechanics plays an important role in the understanding of the fundamental principles of animal movement; the prevention and treatment of musculoskeletal diseases; and the production of implements and tools 6

that are related to animal movement mechanics, and skeletal joint kinematics. In addition, performance connotes global activities such as operating vehicles or tools, and sport mechanics. 1.5. Principle of Body Design: The components of the body are designed to provide adequate strength with minimum materials to save on weight, bulk and metabolic requirements. This principle is important in analysis of skeleton. For example the bone material develops along the line of action of applied force . The following chapters mainly concern with the Biomechanics and its application with reference to quadruped locomotion, kinetics of locomotion, stress and strain falling on locomotor apparatus, landmarks, angulation and weight bearing bones of ox, buffalo and comparison with other animals particularly horse and dog.

One of the principal objects ... in any department of knowledge is to find the pointof view from which the subject appears in its greatest simplicity, Josiah Willard Gibbs (1839-1903) (the father of thermodynamics)

II (A). LOCOMOTION
The act of changing place or position by the entire body or by one or more of its parts is called movement. Movement is one of the characteristic features of living organisms. Study of movements is called kinesiology. In animals, movement is of two main types: 1. Motion: Movement of part (or parts) of body is called as motion. 2. Locomotion: Movement of whole body from place another place is known as locomotion. Muscular movement is the method used in most of the vertebrates, including man. Muscular movement is based on the use of muscle fibres. Muscle fibres have the unique property of ability to contract and relax, which exerts a force. This force is responsible for movement of body parts and locomotion. There are a number of descriptive anatomical features that can be linked to the locomotor type. Thus animals that leap tend to have long hind limbs; animals that move quadrupedally tend to have hind limbs that are the same length as their forelimbs and brachiators tend to have long forelimbs. Locomotor behaviour has also been linked to body weight. Quadruped movement and locomotion also result from co-operation between muscles and bones and joints (plus nerve and blood supplies). The movements of animals are controlled by a complex system of muscles and their tendons, bones and joints The study of locomotion requires solutions of a multiple approach of mechanics of multiple joint systems, hydrodynamics, aerodynamics and automated synchronizing controlling system. The muscles act as motor for power generation that is transmitted with help of their tendons to the bone and joint system. The bones act as links and the joints as kinks. They form kinematic chains. The totality of links so connected that if one of them is secured and another is set into motion, the remaining ones will of necessity move in a preset manner. Hence study of LOCOMOTOR APPARATUS (Bones, Joints and Muscles) and their Mechanobiological Properties in the animal body becomes essential. The animals that move on the ground are called as cursorial quadrupeds are cursors and evolved from walkers and are either predators (e.g. lion, dogs) or herbivores (ox, buffalo, horse). Animals that jump or hop are known as saltatorial. Saltators are frequently bipedal and if both the hind limbs are used in a single jump or a succession of jumps (kangaroo), the gait is called as ricochet. 2.1. FORCES: Force: A force is that changes or tends to change the state of rest or uniform motion of body. KIND OF FORCES: There are mainly three kinds of forces, viz., 1. Attraction or Repulsion 2. Tension or Thrust 3. Action and Reaction

1. Attraction and Repulsion: when one body exerts a force on another body without any visible and physical means, the force is called the force of attraction (the bodied tend to approach each other) or repulsion (the bodies tend to separate from each other). In biostatics, this type of force is weight. The weight of a body is the force with which the earth attracts it towards its center. It is equal to mg (m = mass and g = gravitational acceleration). 2. Tension: If a force tries to stretch (increase in size)of bone, tendon or ligament is called as Tension or Pull (force). 3. Thrust. The force which pushes a body is called as Thrust or Push (force). Important Properties: 1. The tension of a light inextensible tendon (ligament) is the same throughout its length along the tendon (ligament). 2. If a weight is suspended from a light inextensible tendon (ligament), its other end being tied to a fixed point, the tension remains the same throughout its length, and equals to the weight suspended 3. When the tendon (ligament) is heavy, its tension varies from point to point on its length and depends upon the weight per unit length. 4. When two or more tendons (ligaments) are knotted together, the tensions are different in different portions on the two sides of the knot, though for each separate portion it continues to have a constant value. 5. If the tendon (ligament) is extensible, the tension will vary with the extension. 6. The Tension in a tendon (ligament) with a weight attached to one of its extremities, passing over a smooth surface like peg (sesamoid bones), pulley (trochlea), grooves etc., remains the same throughout its length and is equal to the weight suspended. 4. Action and reaction: When one body is in actual contact with another body, the force exerted at the point of contact by the first body on the second is called the Action b and the second on the first is called the Reaction. The two forces of action and reaction are equal in magnitude, opposite in direction and acts along the common normal at the point of contact of the bodies. 2.2. MOMENT OF FORCE: Force acting on a rigid body can produce different kinds of effect depending upon its location of point of action even though its magnitude and direction is kept the same. For examples, the force applied along any axis, say X-axis, produces pure translational motion and if it acts at any point, it produces translational as well as rotational motion The measurement of tendency of a force to tilt, turns, or rotates a body about some point, line, or an axis is called as the moment of the force (torque). Thus, mathematically: (Magnitude of) moment of force F = (Magnitude of) Force Perpendicular Distance between F and the point about which the moment is measured = Force Length of perpendicular distance. One of the examples of moment of force is the lever a simple machine. The concept of moment of force is important in understanding the lever system of the body. The animal can move without wheels. It is possible because of angulated 9

joints and the lever system which comprises of acting force (efferent force produced by the muscles), bones (lever arm) and joints (pivot). Lever is a machine that is used to lift a heavy load by providing an effort force of low magnitude through the pivot. 2.3. Lever : A lever is a rigid structure that transmits forces by turning (or tending to turn) at a point knows as pivot. The perpendicular distance between the in-put-force, Fi (applied force) and the pivot is called as in-put lever arm (Li) and the perpendicular distance between the load, Fo, (out-put force) and the pivot is called as output lever arm (Lo). The product of force and its lever arm is called as torque (). The in-put-torque (i) is equal to Fi Li and the output torque (o) is equal to Fo Lo. When Fi Li > Fo Lo , the lever rotates in the direction of Fe ; when FI Li > Fo Lo, the lever rotates in the direction of Fo ; when Fi Li = Fo Lo; the system is in equilibrium and there is no rotation. Thus when tetrapodes stand, the forces of all postural muscles are adjusted so that effort torque equals the load torque and the animal stands without motion. Types of lever system: There are three types of lever system (Figure2.1). 1.Type-I lever system: The effort-force and the load-force are acting at the ends of the lever arms and the pivot(P) is present between them, e.i., Fi P Fo , (Figure 2.1a). The efforttorque and the load-torque are directed in opposite directions from each other. Further, the effort-lever arm is, in general, shoter than the load-lever arm. In other words by applying a small force Fi = (Fo Lo / Li), a large amount of load can be lifted. Mostly, the extensor muscles form such type of lever system. For example, the extensor muscles of elbow joint (mainly triceps brachii, anconeus and tensor fascia antebrachii muscles) insert at summit of olecranon process of ulna and act as effortforce to lift the forelimb below the elbow joint. The elbow joint acts as a pivot. The load-force is the weight of the forelimb. The type-I lever system is the most efficient lever system.

Figure 2.1a: Type-I lever system

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The size of head and length of neck of different quadrupeds can best be understood on the basis of moment of force. However, it requires the understanding of concept of centre of gravity (CG; Figure 2.1b). The centre of gravity is defined as the imaginary point in the body where the total weight of the body resides.

Figure 2.1b: Centre of gravity In most of the quadrupeds CG is located nearer to the forelimbs. In other words the forelimbs bear more weight than the hind limbs. The ratio of weight born by the forelimbs and the hind limbs is about 60:40 in the horse; 58:42 in the buffalo, 56:44 in the ox and the dog. Thus the CG acts as a pivot for type-I lever system. Let Li is approximately equals to the length of neck and head and the weight of these structures together is about Wi. The corresponding values of part body caudal to CG are Lo and Wo, respectively. Therefore, when the body is in equilibrium condition ( no motion), Wi Li = WoLo; hence Li = Wo Lo / Wo. Thus, the animals having larger head have shorter neck (e.g., elephant) than the animals having lighter head (e.g., giraffe). 2. Type-II lever system: The effort-force is present between the pivot (P) and the load-force, e.i., P Fo Fi , The direction of the effort and load torques is in the same direction that is both torques are parallel to each other in the same direction. The load force is closer to the pivot than the effort force. This type of lever system is less found in the body. The lateral movement of neck and head is one of such lever systems in the animal body. It is less efficient than the Type-I lever system. 3. Type-III lever system: The effort-force is present between the pivot (P) and the load-force, e.i., P Fi Fo , (Figure 2.1c). The direction of the effort and load torques, are in the same direction that is both torques are parallel to each other in the same direction. The load force is farther from the pivot than the effort force. This type of lever system is frequently found in the body. Most of the flexor muscles act as Type-III lever system. 11

Figure 2.1c(i): Type-III lever system

Figure 2.1c(ii): Type-III lever system For examples: (1) teres major muscle inserts at the teres major tuberosity, which is present between shoulder joint (pivot) and the remaining part of forelimb and acts as a flexor muscle of shoulder joint. The limb below the shoulder joint acts as a load force (2) biceps brachii muscle has its insertion at radial tuberosity, which is nearer to the elbow joint (the pivot). The limb below the elbow joint acts as a load force.

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It is less efficient than the Type-I and Type-II lever systems, however, the gain in the form of distance moved (longer arch as limbs act as composite pendulum) by applying Type-III lever system is larger than the other two types of lever systems. The important thing is not to memorize the diagrams but to learn to identify the pivot, effort- and load-forces, and effort- and load- lever arms in any specific bone-joint-muscle system. Since the velocity is also a vector quantity (force is a vector quantity), there is also velocity lever system but in reverse order of force lever system that is Vi Lo = Vo Li, so that Vo = Vi Lo / Li. For example, for high velocity at the foot, the olecranon process of ulna should be short and the forearm long Figure 2.2. The same lever cannot enhance both the force and the velocity of the same muscle. It is often desirable to know the speed of an animal as a whole. It may also be useful to know the maximum velocity gained by the animal during the motion. Maximum stride speed of an animal (vmax) = (5

h P / m)1/ 3

(Dubal, 1997)

Where h = hip length; P = maximum power effort and m = mass of hind limb. The increase in hip length increases the speed, while increased mass of the limbs decreases the speed. We will learn more in the following pages when we disuse muscle mechanics.

Figure 2.2: Point of application of in-force and out-force in digitigrades and ungulates. 2.4. STAY APPARATUS (Eschbach, 2012):

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Horses can stand on their feet for long periods, much longer than other domestic animals. This enables them to rest and doze and still be prepared take flight in a second's notice if need be. If they were lying down when a predator arrived, the time it would take to get to their feet and get moving could cost them their lives. Three features of equine leg anatomy allow horses this advantage. They are the stay apparatus, the reciprocal mechanism, and the locking mechanism of the stifle joint. The stay apparatus consists of ligaments and tendons that stabilize all the joints of the forelimb and the hind limb. Minimal muscular activity is needed to hold tension on these ligaments and tendons, which in turn prevent flexion of the joints and collapsing of the leg. The function of this is to bear weight in a very efficient manner, most economical method. Animal can lock one limb and keep the other one relaxed. The Stay Apparatus is a unique mechanism that allows the equine to "lock" both the front and the rear limb. Although most commonly discussed as a mechanism that allows the horse to "sleep" while standing, the stay apparatus is also an integral part of movement. During the gait phases, as the leg impacts the ground, the stay mechanism engages to transfer extreme amounts of force from the foot to the muscles of the body, thus dissipating energy. At rest, while the horse is standing still, the stay apparatus engages and allows the horse to stand, exerting very little energy. It has been estimated that the stay apparatus can save up to 90% of energy that would have to be used without it. I. Forelimb stay apparatus It includes mechanisms to prevent flexion of the shoulder and elbow joints, and to prevent overextension or flexion of the carpal joint. The horse uses its forelimb stay apparatus to support the weight of the front end of its body while using a minimal amount of muscular activity. The stay apparatus helps prevent fatigue of the limb muscles by reducing the amount of energy required to keep the limb stable. The stay apparatus transfers weight from the forelimb muscles to connective tissue structures that do not tire, namely tendons, ligaments or bone. With the weight shifted from muscle to connective tissue elements or bone, the horse requires less muscular activity to keep the limb stable and hold up the front end of the body. The stay apparatus helps the limb resist gravitational forces that would otherwise cause the thoracic limb joints to flex and allow the body to collapse to the ground. The serratus ventralis muscle is the major component linking the forelimb with the body. Its muscular belly is interspersed with inelastic fibrous tissue which functions to support the body when the muscle is relaxed. This means that body weight hanging from the upper end of the scapula tends to flex the shoulder joint (glenohumeral joint). II. Hind limb stay apparatus It includes mechanisms to lock the stifle and hock joints including the presence of a reciprocal apparatus, which ensures that the two joints perform in unison (Figure 2.3). The horse uses its hind limb stay apparatus to support the weight of the caudal end of its body while using a minimal amount of muscular activity. The pelvic stay apparatus is more significant than that in the fore limb. When employed by one hindlimb, the stay apparatus allows the other hind limb to be placed in a "resting" position with just the tip of the hoof touching the ground. Although the stay apparatus 14

reduces the amount of energy required to remain standing, the amount of muscular effort is not reduced. This explains why you see horses switching their weight from one hind limb to another. The hind limb stay apparatus has three essential elements. The first element, the stifle joint locking mechanism, allows the weight of the caudal body to rest, essentially, on the locked joint. The second element, the reciprocal mechanism, ensures that the stifle and hock joints will move in unison, and the leg will move in a smooth, coordinated manner. The first and second elements work together. The third element involves other ligaments/tendons in the distal limb. In the hind limb the tendinous peroneus tertius muscle can be broadly compared to the biceps in the forelimb and extends from the femur to the cannon bone. Weight pushing down tends to flex the stifle and is counteracted by tension building up in the inelastic fibers of the peroneus tertius muscle. This action on the stifle is reflected as a mechanical flexing action on the hock, but the hock is prevented from flexing by the gastronemius muscle extending from the rear of the femur to the point of the hock (calcaneus). III. Reciprocal apparatus It consists of two tendinous cords in the hind limb of the horse, the peroneus tertius and the superficial flexor, which connect the distal end of the femur to the hock, one on the cranial face of the tibia, the other on the caudal face; they ensure that the two joints always move in unison. This mechanism is present in both the fore- and hind limbs and consists of a series of muscles and ligaments which can "lock" the main joints in position. At least part of the mechanism is practically identical in both limbs, based on the suspensory ligament at the back of the cannon bone (MC3). Unlike normal ligaments, running from bone to bone, check ligaments run from bone to tendon so that at the limits of muscle extension they come into action to support the tendons making the tendon function as a ligament by cutting off the muscular attachment above. The stay apparatus of the limbs stabilize the joints below the knee (carpus) and hock (tarsus); the remaining joints are maintained in an extended position by a system of muscles. The shoulder is maintained in extension due to the action of the supraspinatus muscle and to a cooperative antagonism between the biceps and triceps muscles. The triceps contract to extend the elbow, and, since the biceps is largely composed of tendinous inelastic tissue, this will maintain the shoulder in extension. The carpal is predisposed to effortless weight bearing since the radius and the cannon bone are in the same vertical line. Vertical weight bearing at the carpus maintains extension due to the "stack" of the carpal bones which in neutral are in an extended position. Further, it is prevented from buckling forward by the inelastic lacertus fibrosus arising from the biceps tendon and running through the radial carpal extensor muscle and tendon onto the front of the cannon. Tension built up in the superficial and deep flexor tendons acts on the accessory carpal bone, pushing in forward, balancing the forces that are forcing the rest of the carpals back. The gastronemius is accompanied by the almost entirely tendinous superficial digital flexor muscle which attaches to the point of the hock before passing down into the shank. During standing it relieves the constant strain otherwise imposed on the gastronemius and maintains the hock in extension. The overall results is a purely mechanical opposition between the peroneus tertius muscles the superficial digital flexor muscle, so that the reference to the hock and stifle joints, one cannot flex or extend without the other following suit. For this reciprocal apparatus to function adequately the stifle must be fixed. This can be 15

accomplished simply by contraction in the quadriceps femoris muscle; however, this muscle may be relieved by the straight patellar ligaments, is raised sufficiently to hock on top of the enlarged upper end of the medial trochlear ridge of the femur. Once the lock has occurred little or no muscular effort is required to maintain posture, more weight falling on the limb merely tightens the hock. To unlock the system the quadriceps femoris muscle contract to pull the patella laterally off the ridge. The hip joint (coxofemoral joint) is maintained in extension by action within the hamstring (semimembranosus and semitendinosus muscles) and gluteus medius muscle.

Figure 2.3. Hind limb stay apparatus

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2.6. Kinematics of Locomotion (Alexander and Goldspink, 1977; Hildebrand, 1995; Baxter 2011): 2.6.1. Forelimb vs. Hind limb Differences (Fletcher T.F., 2010): Forelimbs and hind limbs have different roles in cursorial quadrupeds. Consequently they are anatomically different: Forelimbs (thoracic limbs) carry about 56 to 60% of static body weight. They are designed to catch body weight as it is thrown forward by pelvic limbs. Forelimbs improve gait efficiency by minimizing wasteful up/down energy expenditure (they absorb kinetic energy of downward movement, store it as potential energy in stretched ligaments, and in turn convert that to upward kinetic energy). Anatomical features of the forelimb include: (i) shorter and straighter than the pelvic limb (ii) connected to the trunk only by muscle and ligament (not bone to bone articulation) broader, more rounded hoof (horse) Hind limbs (pelvic limbs) are the "motors" of locomotion. They drive the trunk forward and propel the body up/over obstacles during jumps. Anatomical features of the hind limb include: (i) longer and more angular than the thoracic limb (ii) osseous articulation to the trunk (axial skeleton) through a sacroiliac joint (iii)musculature capable of simultaneously extending hip, stifle, and hock (iv) narrower, more pointed hoof. 2.6.2. Requirements of Cursors (Hildebrand, 1995): For being an efficient runner, an animal must have the following requirements: 1. It must overcome the inertia of its body to gain speed. 2. It must overcome the movement and inertia of limbs. 3. It must abolish transverse oscillations of any parts. 4. It supports the body during motion. 5. Its body must be adopted to reduce the forces of deceleration (wind resistance and ground reaction of feet) 6. It must control its course and movement and 7. It must maintain these functions during motion. The carnivore lifestyle requires limb flexibility and control for grasping and manipulating prey and defending self, offspring, and territory. Thus limb elongation, bone reduction, digit elimination, and conversion of muscle to ligament is moderated. The unguis (nail) in not used for locomotion and remains available as a "tool". To catch long-leg prey, the carnivore must obtain long stride length physiologically, by trunk flexion and extension (which requires much energy expenditure). The herbivore exhibits extreme anatomical specialization for running. It walks on the hoof of a single digit of an elongate manus / pes in which muscle is replaced by ligament. The digestive apparatus predisposes it to continual grazing (in the wild). Accordingly, it has evolved ligament structures that facilitate such a lifestyle, including prolonged standing with minimal expenditure of muscular energy (the stay 17

apparatus, particularly in the equine), mechanical joint linkage (reciprocal apparatus), and mechanical energy conservation through potential/kinetic energy exchange (e.g., fetlock translation). Some Definitions (http://vanat.cvm.umn.edu/gaits/glossary.html): 1 .Step: One limb undertaking one cycle in a stride (cycle = lift, swing, support, thrust). 2. Stride: A unit of locomotion during which all limbs each complete one step (at the completion of a stride, limbs have the same relative positions as when they started); also, 3. Stride length: The linear distance between two successive ground impacts of the same foot. 4. Cycle (stride cycle) = sequence of movements that limbs undergo during one stride; movements a limb undergoes in returning to its original position. [To the left, phases of the step cycle that a paw undergoes are illustrated.] The time taken in one complete cycle is the gait period The inverse of the period is the gait frequency (1/period). Normally, in one gait cycle, each leg goes through exactly one complete step cycle 5. Sequence: The series of (four) steps comprising a single stride (of any gait). 6. Beats = the number of ground impacts during a stride (number of "hoof-beats" one might hear during a stride), e.g., if two legs simultaneously impact the ground & then the other two impact simultaeously, the gait is two-beat (trot or pace). 7. Angulation = refers to the angles formed by bones of the limbs with respect to themselves and with respect to the ground. 8. Gait = a particular sequence of limb movements repeated to produce locomotion, e.g., walk, trot, pace, canter, gallop, etc. 9. Walk = a four-beat gait in which all legs step sequentially, used for leisurely travel. 10. Amble = a four-beat gait which is essentially an accelerated walk (preferred by elephants and some horses; only a transition gait in the dog). 11. Trot = a two-beat gait that uses only diagonals for support, used for traveling long distances at a fair rate of speed. 12. Pace = a two-beat gait that features sagittal support (combined use of ipsilateral fore and hind limbs); employed instead of the trot by some animals. 13. Power walk = the type of walk used by animals pulling a load; steps are shorter and slower than in a normal walk, and sagittal support is minimized. 14. Canter = a three-beat gait, often used for non-strenuous, playful locomotion; involves rear limb, diagonal, and front limb supports. 15. Gallop = a four-beat gait that features suspension phase(s); also, the fastest gait. Two types of gallop exist (see: Transverse gallop and Rotatory gallop). 16. Rotatory gallop = a four-beat gait of bovines, carnivores, swine, rodents, and small ungulates that features trunk flexion/extension and two suspension phases; a suspension phase occurs after the second-landing hind limb is lifted, as well as after the second-landing (leading) fore limb is lifted. 17. Suspension = phase of locomotion during which no limb is touching the ground (supporting the trunk).

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Since a full cycle of movement of an animal is called as a stride, the Stride speed is equal to the product of stride length and stride frequency. Animals with long feet emphasize stride length, while short feet ones emphasize stride frequency. A fast speed requires both. Stride speed: The horses run about 70 km / hr, several antelopes run at 85 95 km / hr. The cheetah seldom runs more than km, but for short distance is the fastest of animals (110 km / hr but not more than 10 15 minutes). Maneuverability is the capacity for rapid and controlled change in speed and direction. It is favored by small body size, because inertia increases with mass. Horse cannot be as agile as mice, monkeys, dog, or cheetah. Nevertheless, horses do well for such large animals (Hildebrand M., 1987) (Figure 2.4). Endurance: Man runs 30 km at 19.5 km / hr Horse can run 80 km at 18.2 km / hr. A camel travels 186 km in 12 hr. Stride Length: A galloping horse moves about 7 m per stride. The fastest, but smallest cheetah moves approximately the same distance per stride about 10 times its shoulder length. The stride length depends on the length of the limbs. Longer the limb, the longer is the stride length. However the limbs must be relatively long in relation to the other parts of the body.

Figure 2.4: Maneuverability of body posture for longer stride length.

Running is mainly about speed -- obtained by combining stride rate and stride length. Stride rate is favored by reducing limb mass (especially distally). Stride length is gained by increasing limb length (anatomically or physiologically). Rate and length are antagonistic, i.e., enhancing one compromise the other. 19

Domestic mammals exhibit cursorial (running) adaptations to varying degrees: Carnivores require multipurpose limbs (both running & manipulating). Carnivores are fast because they have flexible trunks, which is possible because they have a meat diet (which is why they need multipurpose limbs in the first place). Herbivores, with roughage diets and bulky abdominal viscera, have relatively limited trunk flexion, but their limbs are freed to be devoted entirely to locomotion. Equine limbs have become so specialized; they resemble "machines" (reliance more on bone & ligament and less nerve and muscle). Since the mass of the limbs and the speed are negatively related, the mass of the limbs is reduced by deleting bones (e.g., fibula, digital and metacarpal bones of camel, ox and horse) or reducing the effective length of bones (e.g., ulna of camel and horse) of the distal part of the limbs. Further the muscles of the distal part of the limbs are not bulky and thus it leads to reduction in mass of the limbs. 2.6.3. Methods used to increase the effective stride length of limb: 1. Modification of Length of limbs: The distal segments of the limb are usually longer than the proximal segments. The cursorial ungulates usually have a radius that is as long as or longer than the humerus, and a tibia that is as long as or markedly longer than the femur. Furthermore the foot skeleton is about equal to or even longer than the middle limb segment. It is the metacarpal or metatarsal that lengthens most. The carpals and tarsals never lengthen 2. Adaptation of Foot posture: The relative length of distal segment of limb can also be made longer Bears, opossums, raccoons, and other vertebrates that walk well but seldom run, have feet that do not contribute in lengthening the limb. In standing animal, the heel is on the ground and strikes first in each stride. Such feet are called as plantigrade (= sole + walking). Running birds and carnivores increase effective length by standing on digits. These animal are called as digitigrades (= finger + walking). Perissodectyls and artiodactyls further increase effective length by standing on the tips of the digits. This foot posture is known as unguligrade ( = hoof + walking) they are called as ungulates (Figure 2.5). The ratios of bone length is important to understand speed propulsive force of the hind leg is effective if the length from the hock to the ground is 1.7 times the length of the femur (hip-to-stifle). Equal length from elbow to knee and knee to ground in the front leg is desirable and a short humerus (shoulder to elbow) with a long forearm (elbow to knee) provides speed advantage. Fast horses can bring their legs forward more quickly in preparation for the next stride. Looking at bone length and skeletal proportions, naval engineer Larry Wellman shed light on how nature has selected for leverage and speed efficiency in hooved runners. Certain bone ratios indicate the fast running ability of the animals. For the hind limb, the most speedefficient formula was a hock-to-hoof length of 1.7 times the length from the stifle to the hip. In the forelimb, the length from the point of the elbow to the carpal and from the carpal to the ground should be equal approximately. A long humerus is defined as over 50% of the length of the scapula or shoulder. For better speed leverage a shorter humerus (shoulder to elbow) and longer elbow to carpal length provide better lever advantages. The length of the forearm or radius from the elbow to the knee - gives the forward pendulum action necessary for ground-covering strides. Length in the forearm is more advantageous than long metacarpal when breeding for height and is more beneficial for soundness. For maximum speed efficiency, the length from the 20

carpal to the ground should be 38% of the ground to wither height. In overall conformation, the total hind limb length should be 25% more than back length (defined as wither-to croup), bounding recoil is superior a long back just providing excess baggage and withers higher than the croup is a feature in outstanding fast running.

Figure 2.5: Adaptation of Foot posture in mammals.

3. Adaptation of Shoulder: The scapula of most mammals is free to move to some degree. In cursors this free-movement is increased by (1) reducing the clavicle to a vestige (carnivores) or absent altogether (ungulates) and (2) Reorienting the scapula so that it lies flat against the side of narrow and deep thorax, where it is free to rotate in the same plane in which the forelimb swings. The shoulder joint then moves in the sagittal plane and lengthens the limb by moving its pivot from the shoulder joint to a point on to the scapula. 4. Adaptation of Spines: The smaller and swifter quadrupeds, particularly carnivores, have their limbs positioned under the body instead of to the side, and consequently undulate the spine in the vertical plane. The back advances at the same time the limbs are swinging back and forth. The body of the animal is longer when the back is extended than when it is 21

flexed. However, the galloping animal can extends its back only when its hind feet are on the ground. Muscles of hind limbs pressing on the ground and friction at the ground prevent the hindquarters from moving backward (deceleration. All of increase in body length is added to stride length. Similarly, the forelimbs prevent or try to reduce, deceleration of the forequarters as the back is flexed. Therefore the shortening of body is also added to stride length. The cheetah is so adept at this maneuver that it could, theoretically run about 10 km / hr without any legs at all. Flexion and extension of the spines and their timing relative to the thrust of the limbs also make the speed of the animal that is its centre of gravity, is slightly greater than the speed of the associated girdle during the time that a pair of limbs is propelling the body. The flexion and extension of spines add extra rotation to hip and shoulder girdles, which increases the swing of the limbs proper so that the limbs reach out farther, craniad and caudad, and strike and leave the ground at more acute angles than they would if the spines were held rigid. This also increases the stride length. The ungulates cannot exhibit body maneuverability similar to cheetah. However, the horses run faster for longer period. This is achieved by reducing the effective length of the caudal portion of trunk from hind limbs.by complete fusion of caudal lumbar vertebrae with sacrum. The forward thrust propelled by the hind limbs causes the faster movement of centre of gravity forwards than the girdles. It increases the surge of faster proper positioning of forelimbs which adds the stride speed through increased stride length and stride frequency.

5. Distance between the Insertion of muscle and joint: Muscles move the joints through angles when they insert close to the joint than when they insert farther away. If the load is not too great this arrangement also benefits the rate of the stride (see below). Further the muscle functions with less shortening. It is characteristics of cursors to have limb muscles that insert relatively closer to joints. 2.6.4. Gear ratio: The stride speed is the product of stride length and stride rate. Fast runner must take long stride rapidly. The galloping race horse completes about 2 strides per sec and the fast-running cheetah completes about 3 strides per sec. When an animal starts getting speed, it at first increases the rate of its stride by causing its muscles to contract faster. However, continuous fast contraction consumes lot of energy that cannot be maintained for longer period. To achieve the fast stride rate, muscle uses high gear ratio mechanism. High gear is used when the vehicle is driven fast. Cursors have higher gears than most other animals. Since muscles move joint through wider angles when they insert close to joints than when they insert farther away. From the the velocity lever system, Vl = VeLl / Le; therefore the load velocity or rather say out-put velocity Vl is also increased by moving the insertions close to the joints(that is by reducing the value of Le the in-put velocity). The value of ratio Ll / Le for teres major muscle is 4.45 in cheetah and 3.36 in lion (Fig2.6).

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Figure 2.6a: The velocity ratio of teres major muscle in lion (Lo/Li = 3.36)

Figure 2.6b: the velocity ratio of teres mojor muscle in cheetah (Lo / Li = 4.45) There is reciprocal relationship between the velocity and force. The cursorial animals can add further speed only by sacrificing the torque. All muscle are not of high-gear system. There are some muscles which are of low-gear system. For example, semimembranosus muscle, with its long effective lever arm, is low-gear system relative to the middle gluteus muscle that has the same action but a much shorter effective lever arm. The ratio Ll / Le is 11and 44 in semimembranosus muscle 23

and gluteus muscles, respectively. Cursors reduce the requirement of force by reducing the loads on their muscles. The stride speed is further increased by moving the different limb muscles simultaneously different joints in the same direction, the independent velocities produced by them are added to derive the total velocity at the foot. Further, the cursors add an pivot to the limbs by adapting the digitigrade or ungulate posture. Furthermore, the scapula rotates through 20 to 25 on the thorax. In the cursorial carnivores, there is timing flexion of the spines with when the thorax and pelvis rotate in the direction of swinging limbs. It also increases the speed. 2.6.5. Design for Economy of effort: The energy cost of locomotion is inversely related to body weight. The large cursors have superior endurance. It is achieved by: 1. There is reduction or elimination of many oscillating motions. The limbs swing back and forth, but the limbs are not lifted so high. 2. The back is relatively stiff. 3. The centre of gravity has less verical displacement. 4. Suspensions are reduced or avoided. 5. The mass of the limbs are reduced in several ways. Adductors and abductors of the limbs are reduced or are converted to move the limbs in the direction of motion. Muscles that control the digital movements, rotate the forearm, or twist the feet inward or outward are also reduced or eliminated altogether. The bones and associated muscles of distal part of the limbs are either lost or reduced or fused. The fleshy parts of the limbs are kept close to the body. The result is a slender, light and strong foot. Examples are ulna, fibula, some metacarpals and metatarsals, and bones of digits and pronator and supinator muscles, soleus muscle etc. in the ox, horse, and camel. 6. The loads on the muscles cause to develop the oscillatory forces that are reduced by reducing the masses and velocities of the limbs. This the principal reason of distal segment of limb to lengthen more than the proximal segment. 7. Most of the joints of limbs act as hinges, allowing motion only in the line of movement. The articular surfaces are modified to interlocking splines and grooves or replacing spherical surfaces to flat or cylindrical surfaces. 8. Moving quadrupeds save energy of motion (about 40% ) by recycling between the gravitational potential and kinetic energy or storing and releasing elastic strain energy of impact. Though these adaptations necessary for endurance, there are also some disadvantages. The water buffalo, ox, camels, horses, giraffes and rhinoceroses cannot start, wheel, dodge, and cannot stop the motion quickly. Small antelopes and some carnivores can maneuver their bodies. These animals have fairly light and supple bodies, alertness, and rapid neuro-muscular co-ordination 2.6.6. Biomechanics of Curvilinear Motion: During turning of a moving animal, the centrifugal force (outward force) must be reduced or nullified by an equal and opposite centripetal force (inward force) to avoid skidding out of its curved course of movement. For turning sharply, the running animal must lean into the turn, adjusting the angle of its body so that the outward component of the thrust of its feet onto the ground equals the centrifugal force. This is opposed by the inward component of of the force of the ground on the feet. Therefore there must be sufficient friction between the feet and ground to skip slipping. Any force F that acts against the side of the animal (e.g., centrifugal force, wind pressure jostling of another animal) tends to imbalance the animal with a torque 24

of F h, where h is the height of the centre of gravity. Stability is maintained when F h is less than W b, where b is half of the width of the animal`s stance and W is the body weight. Thus broad and short legged animals are most stable. However, the cursors are long legged and often lift both right or both left limbs at once to move fast. Further, they must lean into turns when running. This is not possible for highly stable animal. The stability is sacrificed for maneuverability and the cursors adjust lateral forces by adjusting posture. For turning sharply, the galloping mammal leads with its inside forelimb. Thus the successive footfalls are in the direction of the turn and the animal cam balance over its support. The cheetah follows almost the tracts of the chasing antelope. The hapless antelope, therefore cannot evade the cheetah`s dash by dodging. 2.6.7.GAIT (Nunamaker Blauner, 1985; Alexander and Goldspink, 1977; Hildbrand 1995; Baxter 2011; http://en.wikipedia.org/wiki/Horse_gait; http://petcaretips.net/horse-gaits.html): Gait is defined as a foot-fall pattern of limb actions that an animal uses repetitively during locomotion (http://vanat.cvm.umn.edu/gaits/index.html). Gait is the coordinated, cyclic combination of movements that result in human locomotion. The movements are coordinated in the sense that they must occur with a specific temporal pattern for the gait to occur. The movements in a gait repeat as a walker cycles between steps with alternating feet. It is both the coordinated and cyclic nature of the motion that makes gait a unique 2.6.7.1. Types of gaits: The types of gaits in cursorial quadrupeds, in increasing order of speed, are (figure 2.7): 1. Walk (& amble), 2.Trot, 3.Pace (rack), 4.Canter, and 5.Gallop. In order to understand the gait, horse will be used as an exemplified. The canter and gallop are regarded as asymmetrical gaits because right and left limbs have different actions (the actions are not mirror images). Asymmetric gaits favor turning toward one side or the other, and an animal is said to be in a "right/left lead" according to which side it is predisposed to turn toward, at the canter or gallop. Animals control which lead they are using, and they switch leads to suit external circumstances or to minimize fatigue. . Walk The walk is a four-beat gait that averages about 6.4 km/h. When walking, a horse's legs follow this sequence: left hind leg, left front leg, right hind leg, right front leg, in a regular 1-2-3-4 beat. At the walk, the horse will always have one foot raised and the other three feet on the ground, save for a brief moment when weight is being transferred from one foot to another. A horse moves its head and neck in a slight up and down motion that helps maintain balance Trot The trot is a two-beat gait that has a wide variation in possible speeds, but averages about 13 km/h, or, very roughly, about the same speed as a healthy adult human can run. A very slow trot is sometimes referred to as a jog. An extremely fast 25

trot has no special name, but in harness racing, the trot of a Standard bred is faster than the gallop of the average non-racehorse. In this gait, the horse moves its legs in unison in diagonal pairs. From the standpoint of the balance of the horse, this is a very stable gait, and the horse need not make major balancing motions with its head and neck. The trot is the working gait for a horse Canter The canter is a controlled, three-beat gait that usually is a bit faster than the average trot, but slower than the gallop. The average speed of a canter is between 16 27 km/h, depending on the length of the stride of the horse. In the canter, one of the horse's rear legs the right rear leg, for example propels the horse forward. During this beat, the horse is supported only on that single leg while the remaining three legs are moving forward. On the next beat the horse catches itself on the left rear and right front legs while the other hind leg is still momentarily on the ground. On the third beat, the horse catches itself on the left front leg while the diagonal pair is momentarily still in contact with the ground. The more extended foreleg is matched by a slightly more extended hind leg on the same side. This is referred to as a "lead". Except in special cases, such as the counter-canter, it is desirable for a horse to lead with its inside legs when on a circle. Therefore, a horse that begins cantering with the right rear leg as described above will have the left front and hind legs each land farther forward. This would be referred to as being on the "left lead". Gallop

Figure 2.7a: The suspension phase, seen in the canter and the gallop The gallop is the fastest four-beat gait with all four feet are off the ground in the suspension phase of the gallop, the legs are bent rather than extended. It is very much like the canter, except that it is, more ground-covering, and the three-beat canter changes to. It is the fastest gait of the horse, averaging about 40 to 48 km/h, and in the wild is used when the animal needs to flee from predators or simply cover short distances quickly. Horses seldom will gallop more than a mile or two before they need to rest, though horses can sustain a moderately-paced gallop for longer distances before they become winded and have to slow down.

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Figure 2.7b: Gallop . The fastest galloping speed is achieved by the American quarter horse, which in a short sprint of a quarter mile or less has been clocked at speeds approaching 89 km/h. Like a canter, the horse will strike off with its non-leading hind foot; but the second stage of the canter becomes, in the gallop, the second and third stages because the inside hind foot hits the ground a split second before the outside front foot. Then both gaits end with the striking off of the leading leg, followed by a moment of suspension when all four feet are off the ground. A careful listener or observer can tell an extended canter from a gallop by the presence of the fourth beat. When all four feet are off the ground in the suspension phase of the gallop, the legs are bent rather than extended. The movement of each limb is partitioned into a stance phase, where the foot is in contact with the ground, and a swing phase, where the foot was lifted and moved forwards. Each limb must complete a cycle in the same length of time, otherwise one limb's relationship to the others can change with time, and a steady pattern cannot occur. Thus, any gait can completely be described in terms of the beginning and end of stance phase of three limbs relative to a cycle of a reference limb, usually the left hind limb. Pace: The Pace is a lateral two-beat gait. In the pace, the two legs on the same side of the horse move forward together, unlike the trot, where the two legs diagonally opposite from each other move forward together. In both the pace and the trot, two feet are always off the ground. The trot is much more common, but some horses, particularly in breeds bred for harness racing, naturally prefer to pace. Pacers are also faster than trotters on the average, though horses are raced at both gaits. Among standardbred horses, pacers breed truer than trotters that is, trotting sires have a higher proportion of pacers among their get than pacing sires do of trotters. A slow pace can be relatively comfortable, as the rider is lightly rocked from side to side. A slightly uneven pace that is somewhat between a pace and an amble is the sobreandando of the Peruvian Paso. On the other hand, a slow pace is considered undesirable in an Icelandic horse, where it is called a lull or a "piggy-pace". With one exception, a fast pace is uncomfortable for riding and very difficult to sit, because the rider is moved rapidly from side to side. The motion feels somewhat as if the rider is on a camel, another animal that naturally paces. However, a camel is much taller than a horse and so even at relatively fast speeds, a rider can 27

Figure 2.7c: Pace follow the rocking motion of a camel. A pacing horse, being smaller and taking quicker steps, moves from side to side at a rate that becomes difficult for a rider to follow at speed, so though the gait is faster and useful for harness racing, it becomes impractical as a gait for riding at speed over long distances. However, in the case of the Icelandic horse, where the pace is known as the skei, "flying pace" or flugskei, it is a smooth and highly valued gait, ridden in short bursts at great speed. A horse that paces and is not used in harness is often taught to perform some form of amble, obtained by lightly unbalancing the horse so the footfalls of the pace break up into a four beat lateral gait that is smoother to ride. A rider cannot properly post to a pacing horse because there is no diagonal gait pattern to follow, though some riders attempt to avoid jostling by rhythmically rising and sitting. Gaits are generally classed as "symmetrical" and "asymmetrical" based on limb movement. It is important to note that these terms have nothing to do with leftright symmetry. In a symmetrical gait, the left and right limbs of a pair alternate, while in an asymmetrical gait, the limbs move together. Asymmetrical gaits are sometimes termed "leaping gaits", due to the presence of a suspended phase. The key variables for gait are the duty factor and the forelimb-hind limb phase relationship. Duty factor is simply the percent of the total cycle which a given foot is on the ground. This value will usually be the same for forelimbs and hind limbs unless the animal is moving with a specially-trained gait or is accelerating or decelerating. Duty factors over 50% are considered a "walk", while those less than 50% are considered a run. Forelimb-hind limb phase is the temporal relationship between the limb pairs. If the same-side forelimbs and hind limbs initiate stance phase at the same time, the phase is 0 (or 100%). If the same-side forelimb contacts the ground half of the cycle later than the hind limb, the phase is 50%. 2.6.7.2. Differences between species: Any given animal uses a relatively restricted set of gaits, and different species use different gaits. Almost all animals are capable of symmetrical gaits, while asymmetrical gaits are largely confined to mammals, which are capable of enough spinal flexion to increase stride length (though small crocodilians are capable of using a bounding gait). Lateral sequence gaits during walking and running are most common in mammals, but arboreal mammals such as monkeys, some possums, and kinkajous use diagonal sequence walks for enhanced stability. Diagonal sequence walks and runs (aka trots) are most frequently used by sprawling tetrapods such as 28

salamanders and lizards, due to the lateral oscillations of their bodies during movement. Bipeds are a unique case, and most bipeds will display only three gaits walking, running, and hopping - during natural locomotion. Other gaits, such as human skipping, are not used without deliberate effort. Gait is the way in which animals move our whole body from one point to another. Most often, this is done by walking, although they may also run, skip, hop etc. Gait analysis is a method used to assess the way animals highlight biomechanical abnormalities. Being able to move efficiently is important in avoiding injuries. Having joints capable of providing sufficient movement and muscles capable of producing sufficient force is vital to generate an efficient gait cycle. If joints are stiff (usually caused by muscle tightness), limiting range of motion, or muscles are weak, the body must find ways of compensating for the problem, leading to biomechanical abnormalities. Biomechanical problems such as these are usually caused by muscular imbalances (tight muscles working against weak muscles), although they can sometimes be caused by structural problems, such as leg length discrepancies resulting in hip hiking. Many injuries are often caused, at least in part, by poor biomechanics. The following are a list of common overuse injuries associated with poor gait biomechanics: Examples of biomechanical abnormalities include: Over pronation Over supination Increased Q angle Hip hiking (or hitching) - lifting the hip on one side Hock equinus - limited hock dorsiflexion Pelvic tilt - can be either cranial, caudal or lateral Shin splints Plantar fasciitis Iliotibial band syndrome (runners stifle) Patella tendonitis (jumpers stifle) Patello-femoral stifle pain Achilles tendonitis Caudal back pain

2.6.7.3. The Gait cycle in walking and running: The gait cycle is the continuous repetitive pattern of walking or running. The gait cycle is split into two main phases, stance and swing, with one complete gait cycle including both a stance and swing phase. The stance phase is the period where the foot is in contact with the ground and equates to 60% of the cycle when walking. The swing pases makes up the remaining 40%. During walking there is a period called double stance, where both feet are in contact with the ground. The swing and stance phases can be further divided into:

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Stance Heel strike - The point when the heel hits the floor Foot flat - The point where the whole of the foot comes into contact with the floor 3. Mid stance - Where animals are transferring weight from the trunk, to the front of feet 4. Toe off - Pushing off with the toes to propel them forwards
1. 2.

Swing: i. Acceleration - The period from toe off to maximum stifle flexion in order for the foot to clear the ground ii. Mid-swing - The period between maximum stifle flexion and the forward movement of the tibia (shin bone) to a vertical position iii. Deceleration - The end of the swing phase before heel strike When running, a higher proportion of the cycle is swing phase as the foot is in contact with the ground for a shorter period. Because of this there is now no double stance phase, and instead there is a point where neither feet are in contact with the ground, this is called the flight phase. As running speed increases, stance phase becomes shorter and shorter. When a mammal is walking or trotting, increases in speed are achieved through increases in stride frequency. At a certain speed, it becomes more efficient for the mammal to gallop rather than to trot, and it changes its gait. Beyond this trot-gallop transition, increases in running speed are achieved by increases in stride length, while stride frequency remains almost constant (Heglund and Taylor, 1988). The relationship between stride frequency at the trot-gallop transition (fs, in Hz) and preferred galloping speed (fp, in Hz) with body mass (M, in kg) of terrestrial mammals can be described by the equation (Heglund and Taylor, 1988): fs = 4.19M-15. And, fp = 4A4M-016. The cycle frequency for maximum power output (fopt, in Hz) of mammalian diaphragm muscle performing oscillatory work and body mass is related as (Altringham and Young, 1991): Popt = 4.42M--16. 2.6.7.4. Biomechanical Considerations (): Cursorial quadrupeds are designed for forward locomotion and they have relative difficulty moving backwards. The normal center of gravity (CG) is located just caudal to the thoracic limbs. The CG can be shifted forward by lowering the head and neck and backward by raising the head. Moving the head to the side shifts the CG laterally. The tail (depending on its length and mass) also contributes to longitudinal and lateral shifts in the CG. Forward motion is generally initiated by one hind limb, which shifts the CG forward and toward the contralateral forelimb (which reaches forward to support the shifted CG). At slow gaits (walk) the CG rhythmically oscillates left/right and the trunk, head, and tail swing from side to side to maintain equilibrium. At rapid gaits, forward momentum and inertia are increased and there is less lateral oscillation (just like a bicycle). As speed increases, fewer limbs provide simultaneous support and the

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"feet" impact the ground nearer the median plane to maintain balance during propulsion. The pelvic limbs are designed for propulsion. They accelerate the CG forward and upward. Pelvic limbs are relatively long and angular, heavily muscled, and connected directly to the vertebral column (coxofemoral and sacroiliac joints). Epaxial muscle contraction assists in elevating the CG. Locomotion proceeds by repetitively throwing the CG forward and then catching it. Elevation of the CG is necessary to extend the duration of forward motion which is temporally limited by the pull of gravity. The thoracic limbs are designed for support (catching the CG). Relative to pelvic limbs, they are shorter and straighter and connected to the trunk by fibro-muscular attachments. They regularly provide upward propulsion and directional stability, but they contribute to forward propulsion only under certain circumstances (when retracting in contact with the ground while forward of the CG; when the animal is pulling a load that shifts the CG caudally). 2.6.7.5. Energy conservation: Both fore and hind limbs are designed to recapture some of the energy expended in elevating the trunk to counteract gravity. The weight of the falling trunk does work in stretching limb muscles and ligaments (storing potential energy) which can rebound (become kinetic energy) to elevate the trunk as the CG passes forward of the weight bearing limb. Due to elastic rebound, galloping gaits are metabolically more economical (less oxygen consumption) than the trot gait at high speeds. 2.6.7.6. Gait velocity: During locomotion each limb is swung forward and then retracted to contact the ground. As gait velocity increases, limbs are increasingly flexed during forward swing to decrease angular mass and facilitate speed of protraction. Ideally, at the moment of paw/hoof impact with the ground, the velocity of limb retraction should equal the forward speed of the animal (i.e., the speed with which the ground is receding). Thus the paw/hoof would impact the ground at zero velocity which minimizes potentially traumatic accelerations.

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The ultimate object is to measure something which we have already seen ... to obtain a numerical estimate of some magnitude, . (James Clark Maxwell)

II (B) BIOMECHANICS OF ANIMAL DRAUGHT POWER 2.7. Biomechanics of Draught ability: Various agricultural operations depend upon the draught ability of the animals. Strength (load bearing capacity) and endurance (stamina- ability of doing the work for longer period) are most important measurements to understand the draught ability. Work (W) is defined as the product of force (F) and the displacement in the direction of force S cos (where is the angle between the line of action of force and the displacement). Thus W = F S cos. The endurance is the measurement of continuous ability of doing the work. In other words the endurance is speed sustained through economy of effort. The economy of effort depends on body shape, on the ways the parts of body are moved and on the magnitude and distribution of masses. The ability of doing the work is known as energy. Thus the endurance depends upon the energy consumed during the motion. 2.7.1. Biomechanics of Muscles: The organs of locomotion and load bearing are mainly bone, joints and muscles as stated earlier. The force (F) generated by a muscle is directly proportional to the cross-sectional area (A) of muscle. That is F A; hence F = k A, where k = constant of proportion. If A is measured in cm2 and F is measured in kgf, then k = 2 10 kgf.cm2 = 20 100 N.cm2. Or approximately, F = 6 A. For example, a man having A of biceps brachii muscle = 4 cm2, then the force generated by the muscle is 20 kgf. In other words the man can lift a weight of 24 kg by a single hand. Thus the strength of the animal depends upon the cross-sectional area of the muscles. Exercise increases not only the mass of the muscle (hyperplasia) but also the cross-sectional area (hypertrophy). The strength of a muscle depends on the cross-sectional area. The arrangement of muscle fibres is important in achieving this goal. 2.7.2. Muscle Organization: Groups of fibers organized into fascicles (bundles). The fibers in fascicle run parallel to fascicle, but fascicle can be arranged in 4 different shapes with respect to tendon: 1. Parallel muscles: Fascicles run parallel to length of muscle. Most of the skeletal muscles are arranged in this way (e.g., sartorius and Sternocleidomastoideus muscles, fusiform muscles e..g., biceps brachii, brachialis and psoas major muscles). The muscle shortens as a direct relationship of the shortening of the muscle fibers. They are able to change length extensively and can move load over a great distance. 2. Convergent muscles: Fascicles spread out like fan on one end and converge to single point on other (e.g., long head of triceps brachii muscle). They produces less tension and distance than parallel muscle but independent contraction of fascicles can produce different movements from same muscle and provides versatility. 3. Pennate muscles (feather like) Muscles fibres are arranged at an angle to tendon (the angle is known as angle of pennation). There are three types of pennated muscles. They rotate about their

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tendon attachment or attachments, progressively increasing their angle relative to the tendon. There are three types of pennated muscles: A. Unipennate : Fascicles angled on one side of tendon (e.g., semimembranosus, Biceps femoris, extensor digitorum longus, Extensor Digitorum Longus muscles). B. Bipennate : Tendon in middle with angled fascicles on either side (e.g., rectus femoris muscle). C. Multipennate : Branched tendon with fascicles organized around each branch (Deltoideus muscle). Pennate muscles produce more tension per unit mass than parallel muscles but cannot move as far. 4. Circular muscles (sphincters): Fascicles are arranged in concentric arrangement. The function is to decrease diameter of openings, guard entrances and exits. The diameter of paralleled fibres muscle can be increased by adding more muscle fibres parallel to the long axis of the muscle. However, the synthesis of muscle protein is costlier affair and it is achieved to minimum extent. Further addition of muscle fibres also increases bulk of the muscle that directly increases the mass of the limbs. The increased mass decreases the speed of the limb. From Figure 4, it is easily understood that with help of a few fibres, A of the muscle can be increased without increasing the bulk of the muscle. The crosssectional area of multi-pennated muscle is the highest followed by bi- and unipennated muscles. For non-pennate muscles: A = m / L Where m is the mass of the muscle, is its density (1.056 g.cm-2) and L is the length of the muscle fibres. For pennate muscles: A = m cos / L Where is the angle of pennation. The length of the muscle fibres depends on the exact geometry of the muscle but tends to be much smaller in pinnate muscles so although cos is 1.0, the A of a pinnate muscle of given mass will be larger than the same sized non-pennate muscle. The amount a muscle can contract depends on its fibre length. Muscles fibres can contract to about 60% of their resting length. Since the muscle fibres in pennate muscles are shorter than the non-pennate equivalent the amount of contraction is similarly reduced. In addition since the line of contraction is not the same as the line of action you need to put in the cos factor too. All in all pennation is a non-ideal mechanism. The most efficient option is to have the line of action parallel to the muscle fibres any angular difference means that energy is wasted producing tension in directions where it cannot be used. However the physical layout of the skeleton is such that we often need high absolute forces rather than contraction distance and pennation is a mechanism that allows this.

Figure 2.8: Types of pennation (from http://education.yahoo.com/reference/gray/subjects/subject/102) 33

Energy production depends upon: 1) blood flow in the muscle, 2) amount of oxygen supply, 3) amount of myoglobin, 4) amount of oxidative enzymes and 5) number of mitochondria. The muscles having more value of these parameters will appear relatively more red. Such muscles are called as red muscles and the corresponding lower valued muscles are called as white muscles. The skeletal muscles are not formed by single type of muscle fibres. The types of muscles fibres can be distinguished histologically by localizing oxidative enzymes (e.g., succinate dehydrogenase - SDH), Fig. 5. There are basically three types of muscle fibres: 1) Red fibres (aerobic slow-twitch oxidative (SO), type I fibres), 2) Intermediate fibres ( fast-twitch oxidative glycolytic (FOG) type IIA fibres) and 3) White fibres (anaerobic fast-twitch glycolytic (FG) type IIB fibres) The red fibres are rich in oxidative enzyme (and hence rich in mitochondria). They are slow twitch fibres and can work for longer period. Their diameter is the shortest amongst the three types of fibres. Hence they are more suitable for low weight carrying for longer period. Muscles of migratory birds are good examples of red type fibres. The migratory birds have to fly long distances (from North Pole to equator). However, most of the muscles of the dog and wolf are entire composed of red fibres, but they are not migratory cursors. The reason is that their native place is North Pole. The main physiological constrain is the extreme cold. They have to sustain in the very cold environment and therefore require continuous production of heat. The red fibres act as one of the thermo-regulators. The white fibres have low

Figure 5: Muscle fibre types

oxidative enzymes and are of larger diameter than the intermediate and red fibres. They are fast twitch fibres for short period. They are good for heavy load carrying for shorter period. The intermediate fibres show intermediate properties. The red fibres are the only fibres which develop first in the embryo. The red fibres, under physiological weight loading, are continuously transformed into white fibres. On the basis of per cent population of fibre types, the muscles having more than 30% of red fibres are generally called as red muscles. For examples the serratus ventralis muscle is red muscles. The masater muscle is a red muscle in young and adult animal, however, because of continuous loading of mastication; they are white muscle in the old animal.

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An animal can be selected for draught work on the basis of fibre typing. For example, there are two bullocks A and B. The fibre profile of bullock A is as follows: red fibres (25% and diameter is 25 m), intermediate fibres (35% and diameter is 55 m) and white fibres (50 % and diameter is 65 m). The fibre profile of bullock B is as follows: red fibres (30% and diameter is 22 m), intermediate fibres (32% and diameter is 50 m) and white fibres (38 % and diameter is 55 m). The bullock A is more suitable for heavy loading for short duration, while the bullock B is more suitable for low loading for longer duration. It is found that the muscles of hind limbs are major muscles for forward thrust, while the muscles of forelimbs are for maintenance of right footing on the ground. For forward movement, a powerful single thrust can only be achieved by unison contraction of bulky muscles. That is why the muscles of proximal part of the body ( above elbow joint in case of forelimbs and stifle joint in case of hind limbs) are bulky. The muscles of distal part of the limbs are just for right footing on the ground. Hence, they are thinner and because of this, they are pennated muscles. It also causes the increased slenderness of the distal part of the limbs. The deer, antelopes and birds have distally very slender limbs.

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Everything should be as simple as possible, but not simpler . An equation is for eternity, .. (Einstein - the greatest scientist since Newton)

III. BIORHEOLOGY Biorheology: It is a branch of mechanobiology which deals with the study of deformation and flow of materials. 3.0. Introduction (Barnes, 2000; Schramm, 2000; Waite and Fine, 2007): Rheology describes the deformation of a body under the influence of stresses. Bodies in this context can be either solids, liquids, or gases. Ideal solids deform elastically. The energy required for the deformation is fully recovered when the stresses are removed. Ideal fluids such as liquids and gases deform irreversibly they flow. The energy required for the deformation is dissipated within the fluid in the form of heat and cannot be recovered simply by removing the stresses. The rheological behavior of materials related to their response to applied stresses must be further extended by the introduction of the time-scale of any deformation process: It is written in the bible that everything flows, if you wait long enough, even mountains For all materials a characteristic time factor l can be determined which is infinite in size for ideal elastic solids and almost zero for liquids such as water ( lw=10-12 s). On the other hand deformation processes relate to characteristic time values t. A high Deborah number (l/t) defines a solid-like behavior and a low Deborah number defines a liquid-like behavior. An example may help to improve the understanding of the above: The famous glass windows of the Cathedral of Chartres in France have flown since they were produced some 600 years ago. The glass panes had a uniform thickness from top to bottom in mediaeval times, but today the glass molecules have flown under the influence of gravity so that the thickness at the top is now paper-thin while the pane thickness has more than doubled at the bottom. The very long time t of this flow process results in a small Deborah-number. Thus one can state that solid glass, in spite of its high lambda-value at room temperature under conditions as stated above, belongs to the group of fluids if we wait long enough! An important conclusion of the concept of Deborah numbers is: substances such as water or glass cannot be classed as liquids or solids as such, but rather they exhibit a liquid or solid behavior under certain conditions of stress, shear rates or time. 3.1. Definition and Measurement: Stress: Stress is a measure of force per unit area within a body. It is a body's internal distribution of force per area that reacts to external applied loads. There are three main stresses: 1. Compressive stress, which decreases the length of the material body, 2. Tensile stress, which increases length of the material body stretching) and 3. Shear stress (it is the surface force that is parallel to the plane area). Stress is often broken down into its shear and normal components (compressive and tensile stresses) as these have unique physical significance. In short, stress is to force as strain is to elongation (11).

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Figure 3.1: Tensile stress strain curve Strain: Strain is the measurement of deformation caused by the action of stress on a physical body. Strain is a dimensionless quantity. It has no units of measure because in the formula the units of length "cancel out". Calculation of Strain : It is calculated as the change in dimension (e.g., length) divided by the original dimension (e.g., length): = L/L. The extension (L) is positive if the material has gained length (in tension) and negative if it has reduced length (in compression). Because is always positive, the sign of the strain is always the same as the sign of the extension. In physics, Hooke's law of elasticity is an approximation that states that the amount by which a material body is deformed (the strain) is linearly related to the force causing the deformation (the stress). Thus, Stress () = E , where E = Youngs modulus of elasticity ---------------------------------------------------Extra reading: Static fluids support normal stress (hydrostatic pressure) but will flow under shear stress. Moving viscous fluids can support shear stress (dynamic pressure). Solids can support both shear and normal stress, with ductile materials failing under shear and brittle materials failing under normal stress. All materials have temperature dependent variations in stress related properties, and non-Newtonian materials have rate-dependent variations.

volume change (or dilatation) p = K V, where G is the shear modulus and K the bulk modulus. All three of these moduli have the same dimensions as stress, that of force per unit area (N/m2 or Pa). It is convenient to use a larger unit, that of 109 Pa, Giga-Pascals, or GPa. Youngs modulus, the shear modulus, and the bulk modulus are related, but to relate them we need one more quantity, Poissons ratio. When stretched in one direction, a material generally contracts in the other two directions. Poissons ratio, v, is the negative of the ratio of the lateral or transverse strain, , to the axial strain, , in tensile loading: v
= trans / long

Materials for which Hooke's law is a useful approximation are known as linear-elastic or "Hookean" materialsThe linear elastic regime, strain is proportional to stress, but stress can be applied in more than one way (Figure 12).The tensile stress produces a proportional tensile strain and the same is true in compression. The constant of proportionality, E, is called Youngs modulus. Similarly, a shear stress s causes a proportional shear strain s = G and a pressure p results in a proportional fractional
:

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Definition of Poisson's ratio: Poisson's ratio v is the ratio of transverse contraction strain to longitudinal extension strain in the direction of stretching force. Tensile deformation is considered positive and compressive deformation is considered negative. The definition of Poisson's ratio contains a minus sign so that normal materials have a positive ratio. Poisson's ratio is usually represented as a lower case Greek v (figure 13). v = - trans / longitudinal

Figure 3.2: A cylinder under Tensile force.A. Longitudinal strain L = (L - Lo) / Lo; B. Transverse strain T = (D - Do) / Do Here, L > Lo and D < Do Normal length = Lo Normal transverse Diameter = Do Deformed Length = L and Deformed Transverse Diameter = D Hence, isotropic positive Poissons ratio = ( -) L / T

Ultimate Load: The load (force) at which the material body ruptures or breaks or fractured is called as ultimate load (UL). Strength (s): It is ratio of ultimate load and the original cross-section area (A) of the material body: s = UL / A. The strngth, as applied to supportive materials, is the capacity to resist force without breakage or permanent deformation. The compressive, tensile and shear strength of a fresh compact bone is about 1330 to 2100 kgf/ cm2, 620 to 1050 kgf/ cm2and 500 to 1176 kgf/ cm2.. thus bones are strongest under compressive stresses and the weakest under the shear stresses. Tendons and ligaments merely crumple under the compression applied lengthwise. However, they are as strong as bone under the tensile stresses. Cartilages have lower strength than bones. Working Load (WL): The force exerting on a body for doing the work. Factor of safety: It is the ratio of ultimate load and the working load. Factor of safety = UL / WL The factor of safety is the measure of design of safety margin of working load. It should always be more than unity. Strength economy: It is the ultimate stress divided with d ensity of material. Strength economy = Ultimate stress / Density (D) Stiffness economy: It the Youngs modulus of elasticity divided with density of material:. Stiffness economy = E / D Buckling: In engineering, buckling is a failure mode characterized by a sudden failure of a structural member subjected to high compressive stresses, where the actual compressive stresses at failure are smaller than the ultimate compressive stresses that the material is capable of withstanding. This mode of failure is also described as failure due to elastic instability. Mathematical analysis of buckling makes use of an axial load eccentricity that introduces a moment, which does not form part of the primary forces to which the member is subjected.

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Bending stiffness (BS) = It is the product of the Youngs modulus of elasticity (E) and the moment of inertia(I). BS = E I. It is also called as flexural rigidity. Slenderness ratio: The ratio of the length of a column to the least radius of gyration of its cross section is called the slenderness ratio (sometimes expressed with the Greek letter lambda, ). This ratio affords a means of classifying columns. Slenderness ratio = Length (L) / Radius of gyration () Radius of gyration (): It is whole root square of ratio of moment of inertia to the area : = [ Moment of inertia (I) / Area (A)] 1/2 Torque () = It is the moment of force which causes rotation in the material (e.g., bone): T = GJ / L Where G = Modulus of Shearing = E / {2(2 + v)} = E / {2(1 1/3)}; L = Length; J = Centroidal polar moment of inertia of cross section = Angle of twist and V = Poissonratio ( approx. = 1/3) Shear Stress for twisted material: = T outer / J = G outer / J Shear Strain: It is the measurement of deformity caused by twist. The deformation is the angle of twist. Plasticity: In physics and materials science, plasticity is a property of a material to undergo a non-reversible change of shape in response to an applied force. Viscoelasticity: Viscoelasticity, also known as anelasticity, is the study of materials that exhibit both viscous and elastic characteristics when undergoing deformation. Viscous materials, like honey, resist shear flow and strain linearly with time when a stress is applied. Elastic materials strain instantaneously when stretched and just as quickly return to their original state once the stress is removed. Viscoelastic materials have elements of both of these properties and, as such, exhibit time dependent strain. Whereas elasticity is usually the result of bond stretching along crystallographic planes in an ordered solid, viscoelasticity is the result of the diffusion of atoms or molecules inside of an amorphous material Depending on the change of strain rate versus stress inside a material the viscosity can be categorized as having a linear, non-linear, or plastic response. When a material exhibits a linear response it is categorized as a Newtonian material. In this case the stress is linearly proportional to the strain rate. If the material exhibits a non-linear response to the strain rate, it is categorized as Non-Newtonian fluid. There is also an interesting case where the viscosity decreases as the shear/strain rate remains constant. A material which exhibits this type of behavior is known as thixotropic. In addition, when the stress is independent of this strain rate, the material exhibits plastic deformation. Many viscoelastic materials exhibit rubber like behavior explained by the thermodynamic theory of polymer elasticity. Creep: Creep is the term used to describe the tendency of a material to move or to deform permanently to relieve stresses. Material deformation occurs as a result of long term exposure to levels of stress (physics) that are below the yield strength or ultimate strength of the material. Creep is more severe in materials that are subjected to heat for long periods and near melting point. Creep is often observed in glasses. Creep is a monotonically increasing function of temperature. Unlike brittle fracture, creep deformation does not occur suddenly upon the application of stress. Instead, strain accumulates as a result of long-term stress. Creep deformation is "time-dependent" deformation. Fatigue: In materials science, is the progressive and localised structural damage that occurs when a material is subjected to cyclic loading. The maximum stress values are less than the ultimate tensile stress limit, and may be below the yield stress limit of the material.

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3.2. Fracture: Fracture is the (local) separation of a body into two, or more, pieces under the action of stress. Types of fracture: In brittle fracture, no apparent plastic deformation takes place before fracture. In ductile fracture, extensive plastic deformation takes place before fracture. Biomechanics of rupture of tendon: Under the action of tensile stress, a tiny internal void is created, which grows into a micro crack in tensile strain. The factors responsible for formation of tiny flaws in the tendons are cross-sectional area and collagen content, composition of extra-cellular matrix), longitudinal heterogenecity, inter-fibre differences and elevation of core temperature. Further formation of a crack relieves the elastic stress. Increase in the transverse length of the tiny micro-crack requires energy. Further application of tensile stress results into fusion of the tiny flaws to form relatively large micro-crack. Further extension of tendon results in a reduction of its energy. Thus, a rise in temperature will enhance the progress of micro-crack. The cyclic tensile loading of equine tendon has been shown to result in an elevation of core temperature. The cyclic overloading creates cumulative micro-damage. When the transverse length of this micro-damage reaches a critical value (say critical length of destructive process or simply critical length of microdamage), there is spontaneous rupture of the material. An increase in tensile strength decreases in the plastic work done in initiating fracture at the tip of flaws.

Figure 3.3: Relation between energy of propagation of micro-crack and transverse length of micro-crack

----------------------------------(Extra reading: The rate of growth of micro-crack is related with a measurement known as stress concentration factor (K). It describes the distribution of stresses at the crack tip. When the value of K achieves a critical value, it is known as fracture toughness, there is catastrophic failure of the material The extensor tendons of fore limb, the gastrocnemius tendon and Achilles tendon had high tensile strength. Hence, these tendons appeared to be highly susceptible to develop flaws).

. -----------------------

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Tensile strength (TS = s) of a tendon is obtained by dividing the UL with its original CSA. Factor of safety (FS) is obtained by dividing the UL with the WL. The critical length (CL) of destructive process (the transverse length of micro-damage) is calculated as follows: 2 CL() = 2 G E / (TS) , G = Surface free energy of the tendon per unit area The exact value of G is not known. The most probable value of G is assumed to be 0.50 10 3 J / m2. Hence, = E / s2

A low but repeated (continuous or intermittent) stress can cause the rupture of Extra reding: the tendon. This minimum stress is known as fatigue tensile stress, which is obtained by stress-strain (SN) curve. From clinical point of view, the critical length of micro-damage and the number of cycles of fatigue failure are important. The critical length of micro-damage indicates the maximum transverse damage, which a tendon can sustain under tensile strain. When the tendon attains the critical length, it ruptures under the applied stress. The higher the stress the less is the durability of the tendon The ultimate load when expressed in terms of unit area is known as tensile strength. Theoretical value of TS should about 0.1 E. However, the observed TS is always several times less. This discrepancy is due to the presence of micro-cracks which reduces the strength.. The extensor tendons of fore limb, the gastrocnemius tendon and Achilles tendon had high tensile strength. Hence, these tendons appeared to be highly susceptible to develop flaws. A low but repeated (continuous or intermittent) stress can cause the rupture of the tendon. This minimum stress is known as fatigue tensile stress, which is obtained by stress-strain (SN) curve. From clinical point of view, the critical length of micro-damage and the number of cycles of fatigue failure are important. The critical length of micro-damage indicates the maximum transverse damage, which a tendon can sustain under tensile strain. When the tendon attains the critical length, it ruptures under the applied stress. The higher the stress the less is the durability of the tendon 3. The fracture toughness (FT) can be calculated as follows: (TS) ( CL) = (2 G E) The fatigue tensile strength (FTS) of a tendon is obtained by its minimum WL (assumed to be equal to 4 CSA of its belly. The number of cycles (N) of repeated loading that could produce the failure is calculated as follows: d(CL)/dN = C ( FTS ) n ( CL ) n/2 C = Constant. Since CL<< diameter of the tendon, then for n = 2, N = (CL) / C ( FTS ) 2 FT =

----------------------------------------------3.3.Biomechanics of bones structures: Adequate strength to resist all stresses is provided to all supporting tissues. However, very large tetrapods (e.g., elephant or very heavy ox, buffalo or horse) modify structure, posture and behavior to minimize stresses on the skeleton and to resist the remaining stresses effectively. 3.3.1. Compression or Tension (Biomechanics of Vertebrae): The vertebral column is made up of vertebrae, which provide attachment to various small to large muscles. Therefore, they should be stronger and heavier. The bulky vertebral column is mechanically and physiologically not effective for locomotion. The body of vertebra should be not only lighter (neither solid nor

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hollow), but also strong enough to withstand the compressive forces. Thus, the body (centrum) of the vertebra is formed by the plates (or spicules) of bones known as trabeculae, which are arranged along the line of action of the acting compressive forces. 3.3.2. Bending in One Plane ( I-Beam Mandible, Zygomatic arch, Ilium and Neural spines): There are some bones, which have large surface area (broad) and are located in the horizontal direction (e.g., mandible and pelvic bone- the os coxae). They, therefore, act as beam and are always under bending forces (Figure 15).

Figure 3.4: Beam-like bone (e.g., mandible) under bending force. The resistance to bending (R) is directly proportional to the square of the height (the dimension in the line of action of force = h) and inversely proportional to the width (dimension perpendicular to the line of action of force = w), that is R =c w h2, where c =constant. These bones are generally flat bones. The bones are always directed in such a fashion that the maximum amount of bone material is placed in the line of action of force. 3.3.3. Bending in Multi-Planes (Column Long bones of appendages): Long bones of appendages are almost cylindrical in shape and are mainly under compressive stress. Since the stress or the strength is inversely proportion to the cross-sectional area (CSA), the structure must have such a geometrical figure which has the greatest CSA. Amongst all the geometrical figures, the circle has the largest area. That is the reason that weight bearing bones and stem of trees are cylindrical in shape. When the bone is under axial compressive force (compression in the line of long axis of bone), there is only lengthwise decrease in the length. However, when the bone is under abaxial compressive force (compression away from the line of long axis of bone), it not only under the action of compression but also it experiences a bending force (Figure 16). When the compressing force becomes larger than the limiting value Fl, the internal elastic forces of the bone cannot equalize the external load and the bone buckles.

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Figure 3.5: Bending in multi-planes (Column Long bones of appendages) For a bone of length L, modulus of elasticity E and moment of inertia I, the limiting force (Fl) is equal to Fl = 2EI / L2. Thus the resistance of a bone to bending is inversely proportional to the square of the length of bone. This the reasone that animal cannot have very long to increase its speed. Other structural material and devices are used to achieve the fast speed goal. Because of bending, the bone materials of convex surface are under tensile stress, while the bone materials of concave surface are under compressive stress. There are some bone materials which neither under the compressive stress nor tensile stress. These are present along central long axis of the bone, which may be known as neutral axis. Thus the bone materials present at the neutral axis do not resist the stresses and are without use. This bone material is appropriately shifted towards periphery creating a hollow space in the centre of the bone. The space is known as medullary cavity, which provides home for the most delicate organ og the body, that is, marrow (one of the organs of haemopoiesis). This is the reason that the shaft of the long bones of appendages is hollow. The part of bone which is continuously under various forces (including bending force) is provided more bone materials and therefore becomes thicker along the line of action of dominant force. The shaft of such bone is thicker at mid-length by increasing inner and / or outer diameter. In some bones, the stress is greater along the concave surface of a curved bone (e.g., radius, third phalanx), then the wall of the bone may be thickest there. Since most muscles insert near the extremities of long bones and since forces transmitted by one bone to another bone across an angulated joint are hardly parallel to the long axis of bone (say shaft). The stress lines arc across the extremities of these bones. Thus the extremities of long bones cannot act like pure column (or cylinder). The solid bony extremities are therefore stronger and heavies than needed. The most economical design is a network of interconnecting trabeculae and thin sheets of bones that are laid down along the lines of stress. This is reason that the extremities of long bones are mainly composed of spongy bones, which are protected by a thin layer of compact bone. Since stress lines change as loads change, the body adopts the dominant lines as compromise of the more usual loads.

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It is a capital mistake to theorise before one has data,

(Arthur Conan Doyle)

IV. MECHANOBIOLOGY OF HISTOGENESIS: Mesenchymal cells are transformed into Fibrous tissues, Cartilage and Bone. Under the influence of compression (decrease length), tension (stretching) and shear stress (lateral deformation) or combination of these stresses, the mesenchymal cells produce the fibrous tissues, while under the hydrostatic pressure, the mesenchymal cells produce cartilage. The mesenchymal cells, fibrous tissues and the cartilages, under the condition of scaffolding and immobilization, are transformed into bones. The mechanical stimuli are responsible to set up the physiological and biochemical activities that lead to formation of these connective tissues.

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Difference of opinion leads to enquiry, and enquiry to truth, . (Thomas Jefferson)

V. BIOMECHANICS OF SYNOVIAL JOINT LUBRICATION

5.1. SYNOVIAL JOINT Several disorders of mammalian musculo-skeletal systems are mechanbiological in origin. The basic function of the skeleton is to sustain the load that the body must bear. Therefore the fundamental mechanical and physical microorganization and remodeling properties of all connective tissues must be characterized. The following pages will illustrate the mechanobiology of lubrication of synovial joint. Locomotion is one of the fundamental functions of the body. It is a result of an extremely complex process of muscular interaction with the skeletal structures. Locomotion of the musculoskeletal system is made possible because of the mammalian body is endowed with a multitude of freely moving joints. The joint is a union of cartilage and cartilage, cartilage and bone and bone and bone. The joint is bound by a uniting medium. The various types of movements of joints depend upon the nature of uniting medium. If, the uniting medium is (1) a fibrous tissue, the joint is practically immovable (synarthrosis); (2) a cartilage, the joint is partially movable (amphiarthrosis) and (3) a synovial membrane with a cavity between the articular segments, the joint is a freely movable joint (diarthrosis). The synovial joints must transmit heavy loads at low operating speeds. In order to make the synovial joint practically frictionless, the synovial membrane secrets a lubricant known as synovial fluid (SF), which maintain the articular segments at a distance. The SF, in the form of a film layer, may of few micrometers to molecular thickness. Articular cartilage is the main structure that maintains the SF film between the articular segments. In healthy conditions they may function for seven or eight decades. 5.2. Articular cartilage (Mow and Ratcliffe, 1997; Klisch et al., 2005 and 2008): The articular cartilage is a hyaline cartilage and lacks perichondrium. It is a solid soft connective tissue which surrounds the articular surfaces of bones at synovial joints. It is composed of mainly collegen fibres, proteoglycans, water and electrolytes. The purpose of this cartilage is to provide a suitable covering to the articular surfaces so as to facilitate motion. These are the actual load carrying surfaces and can be regarded as the bearing surfaces. Embedded in the intercellular matrix of hyaline cartilage, there are cartilages forming cells called chondrocytes. The ratio of cell mass to organic tissue mass ranges from 1 to 10 per cent. The thickness of cartilage layer ranges between 1 to 7 mm and varies from species to species and also from joint to joint. Exercise increases the thickness of the cartilage. The central part of articular cartilage is thicker on convex articular surfaces and the peripheral part is thicker on concave articular surface. The articular cartilage growth is dependent on biomechanical factors that are directly affected by local conditions such as stress, strain, or fluid velocities. These conditions themselves are in turn affected by the material properties which are evolving with growth (Davol et al.,). The synovial membrane is also called as synovial intima. It contains four layers of synovial cells embedded in a granular, amorphous, fibre-free intercellular matrix. These cells also called synoviocytes and form an incomplete layer of 1 to 4 cells thick and no basement membrane can be detected. 45

5.3. Mechanical properties: Functional adaptation of biomechanical properties takes place in early life. Depending on the type of loading the site-independent homogeneous cartilage of foetuses adapts during development and maturity to a functionally heterogenic tissue. The joint loading at young age may be the best prevention for joint injury in later life. Daily exercise prevents early cartilage degeneration and maintains normal cartilage. Thickness: The thickness of the articular cartilage varies with species, the particular joint, and the location within the joint. The thickness of the cartilage varies from a few micrometers to several millimeters in different animal or human joints and anatomical locations. Cartilage stiffness is also known to exhibit site-dependent variation within the joints The thickness of articular cartilage of femoral condyle (of adult) is as follows (Stockwell, 1971): Man (2.260.49 mm); Cow (1-68 0-12 mm); Sheep(0.840.26 mm); Dog (0.67+0.28 mm); Rabbit (0.2140.067 mm); Cat (0.327 0.150 mm); Rat (0.0720.013 mm) and Mouse (0.0580.007 mm) The thickness of the articular cartilage is related to the congruence of a joint; thin cartilage is found in congruent joints such as the ankle, whereas thick cartilage is found in incongruent joints such as the knee. The correlations imply that the larger and heavier is an animal the thicker is the cartilage in the lower limb joints. Collagen and PGs are the most important components determining the mechanical integrity of cartilage. Their interactions with the tissue water determine the resilience and compressive stiffness of cartilage as well as establish the anisotropic and poro-viscoelastic behavior of the tissue. The equilibrium modulus of cartilage increases along the cartilage depth. The mechanical properties, the composition and morphology of articular cartilage vary with age, species and joint and within a joint, from one anatomical location to another. When cartilage is loaded at high rates, it behaves as an incompressible material with no significant fluid flow. The dynamic mechanical properties of cartilage are strongly determined by the integrity of collagen network. Collagen lacks compressive properties, but demonstrates tensile viscoelastic and nonlinear behavior. Instantaneous stiffness of the tissue is significantly reduced due to the fibrillation of superficial collagen. 5.3. Mechanical properties are the result of: (1) Collagen content, (2) Collagen orientation, (3) Proteoglycan content and (4) Water content. 5.4. Essential interaction aspects of the collagen-proteoglycan are: 1. The proteoglycans regulate fluid transport in the cartilage, and hence control the time-dependent viscous properties of the material; 2. An unsupported collagen network cannot sustain compressive stress states, so that complete extraction of the proteoglycan complex leads to compressive buckling of the network and substantial degradation of mechanical performance of the material; 46

3. Unloaded matrix configuration, the collagen network is highly stretched by the proteoglycan swelling stress, so that even after substantial volume reductions (25%) the collagen network is still in tension and influences the macroscopic compressive response of the tissue and 4. The long-term response of the material under tensile stress states is entirely controlled by the collagen network, and is not affected by proteoglycan extraction. The articular cartilage tissue is non-linear with strain, both in tension and in compression, non-linear with direction of stimulus, anisotropic in tension and compression, non-homogeneous with depth, resulting in depth dependent mechanical properties, and presents fluid dependent and fluid independent viscoelasticity. The articular cartilage therefore exhibits complex mechanical properties such as anisotropy, inhomogeneity and tensioncompression nonlinearity. All local material properties are the direct consequence of the local composition and structure of the tissue. The articular cartilage shows the following mechanical behaviors: 1.Viscoelastic behavior. 2. Swelling behavior, 3. Compression behavior, 4. Tensile behavior and 5. Shear behavior Application of pure shear under small strains does (ideally) not cause volumetric changes of the tissue samples. Thus, no significant pressure gradient or fluid flow through the matrix occurs. The intrinsic shear stiffness of the PG-Collagen matrix is mainly associated with the collagen fibrils. PGs by themselves may not contribute directly to the overall stiffness in shear. However, they prestress the collagen network because of their ability to generate swelling pressures 5.5. Importance of inhomogeneity and anisotropy in the matrix components: Composition and structure of cartilage varies through the depth of the tissue, and location of the tissue in the joint. Hence, the mechanical properties of AC are nonhomogeneous and anisotropic. Due to the nonhomogeneous distribution of PGs, there is a nonhomogeneous distribution of FCD through the depth of the tissue, which results in depth-dependent swelling pressures. As the orientation of the collagen fibers changes over the height of the tissue, and because they can only resist load in tension, the collagen network plays an important role in the mechanical inhomogeneity and anisotropy of AC. In the superficial zone the collagen fibrils are arranged parallel to the articular surface, thereby giving this layer a high tensile stiffness in this direction, while it has very low stiffness in the perpendicular direction. These fibrils therefore give a large contribution to the mechanical resistance against indentation. The fibrils in the deeper zones are mainly oriented perpendicular to the articular surface, and therefore do provide great resistance against swelling 5.6. Interaction in shock absorption and lubrication: Articular cartilage and synovial fluid interact in the functions of shock absorption and lubrication. Articular cartilage is the main component for shock absorption, but maintains its properties partly because of imbibition of (an extract of) synovial fluid, due to the osmotic pressure discussed above. During loading of the cartilage, fluid will be pressed out of the cartilage into the synovial cavity, which is called exudation (opposite of imbibition). Exudation of synovial fluid is important for the lubrication process. However, permeability of cartilage is low and decreases more with increasing load. Furthermore, osmotic pressure increases with increasing load. Thus, with increasing load extra exudation of fluid becomes harder and harder. In other words, the 47

exudation process has a built-in 'safety break', so that articular cartilage will always be tensioned by and filled with fluid. Still, exudation allows a small but functional amount of fluid to be squeezed into the synovial cavity where it is needed for lubrication. In summary, articular cartilage has good shock absorption properties because of Elastic properties of collagen proteoglycans strengthening the collagen matrix (enhancing elastic properties). The viscous properties of the (synovial) fluid content of cartilage are due to: 1) Electromagnetic repulsive forces of glycosaminoglycans, 2) Pretensioning the matrix osmotic pressure by attracted calcium and sodium ions and 3) Pretensioning the matrix by fluid attraction (enhancing viscous properties via electromagnetic mechanism). Thus, shock absorption is achieved via visco-elasticelectromagnetic tissue. During its long life, a synovial joint must transmit quite large loads of varying sizes often changing cyclically at a frequency of one cycle per second for many thousands of cycles. For example, the knee joint during walking flexes and extends through 60 degrees or more, and reaches an angular velocity of about 800 degrees per second during the swing phase. Natural joint lubrication is a complex process that allows the joint to operate in different conditions. Such conditions include: joint surface velocity and load applied on the joint. The function of the lubricant is to maintain the articular surfaces at a distance apart so that there will be no physical contact of the articular segments at anytime. 5.7. Types of joint lubrication: Depending upon the structures involved, there are two types of joint lubrication. 1. Boundary lubrication: Lubrication is maintained by synovial fluid that is mainly released from synovial membrane ((Glycoproteins separate joint surfaces, few molecules thick): At high loads and low velocity, boundary lubrication is used as a means to protect the surfaces and minimize their contact. Boundary layer lubrication is one of the major low-friction characteristic of normal joints. The lubricant binds to articular cartilage where it retains a protective layer of water molecules. This type of lubrication relies mainly on the chemical properties of the lubricant. For low speeds and high loads, the thickness of the lubricant film decreases. Eventually the bearing surfaces will come into contact at "high spots" in the irregular, "wavy" articular surfaces. In this situation frictional resistance to sliding motion increases and, therefore, wear of the articular surfaces will increase. For boundary lubrication, the friction (and resistance to wear) is no longer governed by the viscosity of the bulk fluid but the physical and chemical properties of the thin surface layers of the lubricant. Synovial fluid is a very good boundary lubricant after being allowed to soak into the articular cartilage. When the disordered forces acting between the two solids at the interface are relatively strong, the force of static friction should be large, but at smaller values of these forces, the system switches over to a regime of weak static friction Boundary lubrication may occur during high constant loading when the synovial fluid is squeezed out from between the articular surfaces, such as during toeoff in walking.

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2. Cartilage-on-cartilage lubrication: Lubrication is maintained by synovial fluid that is mainly released from articular cartilages. There is finite layer of lubricant between articular surfaces At higher velocities, a fluid film is generated between the surfaces because of pressure build-up, and completely separates the surfaces. Since articular cartilage is highly deformable under pressure, this deformation may enhance the thickness of the fluid film. Articular cartilage is filled with"water"that is squeezed from the surface upon loading. Some amount of synovial fluid (fluid film) is always present between the articular segments. This fluid film lubrication occurs when the bearing surfaces are separated by a layer of fluid that is many times thicker than the surface irregularities, or asperities. Fluid film lubrication is dependent more on the physical properties of the lubricant. The most important property of a lubricant in fluid film bearings is its viscosity. This is because it governs the resistance to motion and the ability of the bearing to develop load-bearing pressures in the fluid. The different types of phospholipid adsorbed onto the surface of cartilage have been isolated by extraction and identified. Phosphatidylcholine (41%), phosphatidylethanolamine (27%) and sphingomyelin (32%) comprise the major components of the lipid layer on a normal cartilage surface for boundary lubrication (or surfactant). These molecules act as a protective layer for the cartilage surfaces and basically provide the final line of defense against destructive contact and possibly wear of the articular cartilage The biomechanical properties of the articular cartilage are important in maintaining the fluid film. During heavy loading, the synovial fluid (SF) is squeezed out between the articular surfaces. Simultaneously, there is in-flow of SF into the articular cartilage and some amount of the SF is trapped onto the articular cartilage. (boosted articular cartilage). Further high loading results into oozing of absorbed SF from the articular cartilage (weeping of articular cartilage). Both boosted and wept SF maintain at least molecular fluid film lubrication. If the opposing bearing surfaces can be prevented from touching each other by the fluid, wear is almost totally prevented and the resistance to sliding is low. Because of this situation, fluid film lubrication is often described as the ideal mode of lubrication. Synovial fluid in the joint is necessary for effective levels of fluid film lubrication which is the primary lubrication mechanism at high speeds. In conditions of low speed, high load and rest, this mechanism is assisted by boundary lubrication achieved in a layer of molecules attached to the surface of the cartilage, also known as surfactant. Boosting and weeping of SF depend upon the following factors: 1. Elasticity of the cartilage 2. Porosity of the cartilage (60 ngstroms diameter pores) 3. Thixotropic property of synovial fluid The poro-elastic bearings with couple-stress fluid as lubricant provide enhancement in pressure and ensure the increased load carrying capacity compared with viscous fluids. This may be one of the reasons in the efficient lubrication and proper functioning of synovial joints. Animals suffering from the pain and loss of mobility due to diseased joints may benefit from implants designed to improve their situation.
5.8. Biomechanical Behavior of Lesioned (Diseased) Articular Cartilage: The contact area between articular cartilages is linearly correlated with the applied load. However, the lesioned part of the articular cartilage dose not participates in load bearing. It therefore decreases the effective load bearing area, the healthy part of the articular cartilage, which in turn, is under the influence of greater loading. The increased stress (load per area) further enhances the damage. Thus, in case of lesioned articular cartilage, the animal should be kept immobilized.

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The man who does not read good books has no advantage over the man who can't read them, . (Mark Twain)

References: Alexander R. McN. And Goldspink G. 1977. Mechanics and Energetics of Animal Locomotion. Ed. Chapman and Hall, London Altringham, j. D. and Young, i. S.1991. Power output and the frequency of Oscillatory work in mammalian diaphragm muscles: the effects of animal size. J. exp. Biol. 157, 381-389 Barnes H.A. 2000. A handbook of Elementary Rheology. Institute of Non- Newtonian Fluid Mechanics University of Wales Baxter G. M. 2011. Adams and Stashak's Lameness in Horses, 6th En. WileyBlackwell, Hoboken, New Jersey Bondi H. 1968. An Introduction to Relativity ed.L. Marder, Lngman Group Limited, London Davol A., Bingham M.S., Sah R. L. and Klisch S.M. 2008. A nonlinear finite element model of cartilage growth. Biomechanics and Modeling in Mechanobiology 7 (4): 295-307. (Available at: http://works.bepress.com/sklisch/11) Dubal. S.C. 1997, Comparative study on gross, histological, histochemical and biomechanical charateristics of locomotor apparatus of buffalo and ox. Ph.D. thsis submitted to Gujarat Agricultural University, Sardarkrushi Nagar, Dantiwada, Banaskantha, Gujarat, India Eschbach D. 2012. Equine Stay Apparatus. http://www. equinevetchiropractor. com/ horse-biomechanics/stay-apparatus Fletcher T.F. 2012. Anatomical Adaptations for Cursorial Locomotion and Impact of Diet. http://vanat.cvm.umn.edu/run Gaits (2012): http://vanat.cvm.umn.edu/gaits/index.html Heglund, N. C. and Taylor, C. R. 1988. Speed, stride frequency and energy cost per stride: how do they change with body size and gait? J. exp. Biol. 138, 301-318. Hildebrand M. 1987. The mechanics of horse legs. American Scientists, 75: 594 - 601 Hildebrand M.1995. Vertebrate Structural Analysis. Wiley John and Sons, New York Horse gait. (2012): http://en.wikipedia.org/wiki/Horse_gait Humphrey J.D. 2003.Review Paper: Continuum biomechanics of soft biological tissues. Proc. R.Soc. Lond. A. 459: 3-46 Humphrey J.D. and Delang Sherry L. 2004. An Introduction to Biomechanics: Solids and Fluids, Analysis and Design. Springer-Verlag, New York Kassab G.S. 2004. Y.C. Bert Fung: The Father of Modern iomechanics.MCB,1(1):522. Klisch S.M., Sah R.L. and Hoger A.A. 2005. Cartilage growth mixture model for infinitesimal strains: solutions of boundary-value problems related to in vitro growth experiments. Biomechanics and Modeling in Mechanobiology, 3: 209223 Klisch S.M., Asanbaeva Anna, Oungoulian S.R., Masuda K., Thonar E. JMA, Davol A., and Robert L. Sah R.L.. 2008. A cartilage growth mixture model with collagen Remodeling: validation protocols.J Biomech Eng. 2008 June ; 130 (3):
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Mow V.C., Ratcliffe A. 1997. Structure and function of articular cartilage and meniscus. In: Basic Orthopaedic Biomechanics. Eds. Mow V.C. and Hayes W.C., Raven Press, New York pp 113178 Nunamaker D. M. and Blauner P. D. 1985. Normal and Abnormal Gait. In: Textbook

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of Small Animal Orthopaedics, C. D. Newton and D. M. Nunamaker (Eds.) Publisher: International Veterinary Information Service (www.ivis.org), Ithaca, New York, USA Poincare H.1908. The future of mathematics. Revue generale des Sciences pures et appliquees, Paris, 19th year, No. 23, December, 1908 Poincare H.1921. The Value of Science and Method. In:The Foundations of Science Science and Hypothesis. The Science Press, New York Receprocal apparatus.(2010). http://cal.vet.upenn.edu/projects/grossanat/largemenu/ hplvlstrm.htm Ross M.W. and Dyson S.J. 2003. Diagnosis and Management of Lameness in the Horse.Elsevier Science (USA). Missouri Waite L. and Fine J. 2007. Applied Bio-fluid Mechanics. The McGraw-Hill Companies, New York .

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Arithmetic and geometry are the foundation and essence of all the sciences that treat of magnitude. But not only are they the foundation, they are also, so to speak, the capstone of these sciences .Joseph Louis Lagrange (25 January 1736 10 April 1813), the greatest of modern analysts

Appendix_I Angulation of Joints of Forelimb and Hind limb of the Ox, horse and Dog

Name of Joint Ox A. Forelimb 1. Synsarcosis 2. Shoulder 3. Elbow 4. Carpal 5. Fetlock B. hind limb 1. Sacro-ilic 2. Hip 3. Stifle 4. Tarsal 5. Fetlock

Articular angle (in Degree) Horse

Dog 45 90 135 160 30 110

140 130 172

120 - 130 150 140

30 168 155 169

30 115 150 150 145

Ox and hoese: 1. Sisson S.and Grossman J.D.. 1964. Anatomy of Domestic Animal. 4th Edn. Saunders and Co.,Philadelphia 2. Herlin A.H., Drevemo S. 1994. Effects of tie-stalls or cubicles on diary cows in grazing or zero-grazing situations - Studies on behaviour, locomotion, hygiene, health and performance. In: Biomechanics of dairy cows; Effects of different management systems. SLU, Institutionen fr husdjurens utfodring och vrd.:page 228 (http://www.vaxteko.nu/html/sll/slu/rapport_utf_vard/RHU228/RHU228E.H TM) Dog: 1. Seltzer J. 2012. Canine terminology - angulation Second in a series http://goldrushkennels.com/movement/movement.pdf 2. Canine terminology Angulation: http://outrageousshadowfilas.com/Documentss/ angulation.pdf

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