Question

Describe the fundamental characteristics of structural materials produced by animals and how these materials meet their functional requirements for tetrapod locomotion on land. Include in your answer: a. the definitions of (1) stress, (2) strain, (3) stiffness, (4) plastic, (5) elastic and (6) strength while comparing spider silk protein and synthetically produced nylon and rayon b. physical principles of terrestrial locomotion and the advantages conferred by various adaptations of legs and arrangements of tendons and muscles c. the differences in locomotory patterns of the cat, horse, giraffe, and human and how architecture contributes to the “spring in your step”

Answer 1

Animals have several characteristics that set them apart from other living things. Animals are eukaryotic and multicellular, unlike bacteria, which are prokaryotic, and unlike protists, which are eukaryotic but unicellular. Unlike plants and algae, which produce their own nutrients animals are heterotrophic, feeding on organic material and digesting it internally. With very few exceptions, animals respire aerobically. All animals are motile (able to spontaneously move their bodies) during at least part of their life cycle, but some animals, such as sponges, corals, mussels, and barnacles, later become sessile. The blastula is a stage in embryonic development that is unique to most animals, allowing cells to be differentiated into specialised tissues and organs.

Structural characteristics:

All animals are composed of cells, surrounded by a characteristic extracellular matrix composed of collagen and elastic glycoproteins. During development, the animal extracellular matrix forms a relatively flexible framework upon which cells can move about and be reorganised, making the formation of complex structures possible. This may be calcified, forming structures such as shells, bones, and spicules. In contrast, the cells of other multicellular organisms (primarily algae, plants, and fungi) are held in place by cell walls, and so develop by progressive growth. Animal cells uniquely possess the cell junctions called tight junctions, gap junctions, and desmosomes.

With few exceptions—in particular, the sponges and placozoans—animal bodies are differentiated into tissues. These include muscles, which enable locomotion, and nerve tissues, which transmit signals and coordinate the body. Typically, there is also an internal digestive chamber with either one opening (in Ctenophora, Cnidaria, and flatworms) or two openings (in most bilaterians).

a)

Fish-like ancestors of tetrapods did not need strong limb musculature because they inhabited waters and were practically imponderable. In the primitive tetrapods, principal function of the limbs was initially restricted to passive anchoring in the course of animal movements on the substrate by means of lateral bending of the body (undulation). However, progressive development of carrying function of tetrapod limbs lead to clearing the body off the substrate which reduced friction costs and made the tetrapods less dependent on the substrate properties. Along with this, the limbs became more important as the active locomotory organs. But at the beginning, this diminished locomotory speed as the momentum caused by undulation could no longer provide additional forward sliding. Locomotory function of the tetrapod limb could be carried out due to both retraction and pronation at the shoulder joint. Relatively short humerus of the primitive tetrapods made it indifferent which of these two particular actions lead to elongation of the steps. In most of the recent tetrapods with sprawling limbs (Urodela, Lacertilia Sphenodontia, Crocodilia), step elongation was carried out mainly by retraction at the shoulder joint. Contrary to this, in Tachyglossidae (Mammalia: Monotremata) retraction is absent while pronation at the shoulder joint becomes the most important component of step elongation. This made it possible to recognize two principal types, pronatory and retractory, of locomotion on the basis of the main movement in the phase of support. A mathematical model describing changes in step length during the phase of support in both of these types is elaborated. It takes into account relative sizes of stylopodium and zeugopodium, the angles of pronation and retraction at the shoulder joint, the angle of adduction at the elbow joint, and the angle of body undulation arc. It is shown on the basis of this model, varying of which of the above parameters is advantageous and which is disadvantageous in each of the locomotory types. In the pronatory locomotory type, adduction (lateral mobility) at the elbow joint is employed. It leads to special changes in morphology of the elbow joint due to which humeral condyle becomes spherical and promotes both adduction and rotation of the entire antebrachium. In the retractory locomotory type, amplification of pronation is to be limited in order to provide step elongation, so certain morphological adaptations occur in the elbow joint which prevent adduction at this joint. For step elongation, retraction at the shoulder joint is usually more advantageous than pronation, therefore historical emergence of the pronatory type could be considered as inadaptive. However, transversal horizontal axis of rotation at the shoulder joint appeared to be a prerequisite of the subsequent appearance of the most perfect locomotion in the therian mammals with their parasagittal limbs. Transition to the parasagittal limb construction was associated with adaptation to jumping asymmetric locomotion. It caused elongation of the shoulder bone downward which lead to widening of rotation cone of the humerus and, at the same time, to reduction of the coracoid portion of the glenoid fossa, the latter became horizontal rather than lateral. As a part of this process, the longitudinal axis of the scapula was displacing caudally with destruction of the suture-like articulation of the acromion process with the clavicle. The latter became articulated with the sternum directly or via much reduced interclavicle (or via procoracoid rudiment). This increases amortisatory function of the shoulder girdle during landing at the final stage of jump.

b)

Terrestrial locomotion has evolved as animals adapted from aquatic to terrestrial environments. Locomotion on land raises different problems than that in water, with reduced friction being replaced by the effects of gravity.

There are three basic forms of animal locomotion in the terrestrial environment:

• legged – moving by using appendages
• limbless locomotion – moving without legs, primarily using the body itself as a propulsive structure.
• rolling – rotating the body over the substrate

The legs of tetrapods, the main group of terrestrial vertebrates, have internal bones, with externally attached muscles for movement, and the basic form has three key joints: the shoulder joint, the knee joint, and the ankle joint, at which the foot is attached. Within this form there is much variation in structure and shape. An alternative form of vertebrate 'leg' to the tetrapod leg is the fins found on amphibious fish. Also a few tetrapods, such as the macropods, have adapted their tails as additional locomotory appendages.

The fundamental form of the vertebrate foot has five digits, however some animals have fused digits, giving them less, and some early tetrapods had more; Acanthostega had eight toes. Feet have evolved many forms depending on the animal's needs. One key variation is where on the foot the animal's weight is placed. Some vertebrates: amphibians, reptiles, and some mammals such as humans, bears, and rodents, are plantigrade. This means the weight of the body is placed on the heel of the foot, giving it strength and stability. Most mammals, such as cats and dogs are digitigrade, walking on their toes, giving them what many people mistake as a “backward knee”, which is really their ankle. The extension of the joint helps store momentum and acts as a spring, allowing digitigrade creatures more speed. Digitigrade mammals are also often adept at quiet movement. Birds are also digitigrade.^[7] Hooved mammals are known as ungulates, walking on the fused tips of their fingers and toes. This can vary from odd-toed ungulates, such as horses, pigs, and a few wild African ungulates, to even-toed ungulates, such as cows, deer, and goats. Mammals whose limbs have adapted to grab objects have what are called prehensile limbs. This term can be attributed to front limbs as well as tails for animals such as monkeys and some rodents. All animals that have prehensile front limbs are plantigrade, even if their ankle joint looks extended (squirrels are a good example).

Among terrestrial invertebrates there are a number of leg forms. The arthropod legs are jointed and supported by hard external armor, with the muscles attached to the internal surface of this exoskeleton. The other group of legged terrestrial invertebrates, the velvet worms, have soft stumpy legs supported by a hydrostatic skeleton. The prolegs that some caterpillars have in addition to their six more-standard arthropod legs have a similar form to those of velvet worms, and suggest a distant shared ancestry.

c)

The mechanics of mammalian walking and running can get quite complicated and involve the use of tendons and the back as energy storing springs to enhance efficiency. The modes of locomotion used by animals have been divided up into more than 30 different types and it is not unusual for an animal to change from one type of movement to another (i.e. from walking to jumping in a given period of locomotion).

Giraffes and a few other animals such as brown bears and camels move both legs on one side and then both legs on the other side. In some cases, the hind leg starts first so there is a slight lag. This is also called ‘Pacing’ and it is the first gait observed in young ‘colts’.

The trot is the same gait as the dog walk, but faster – so that there is a moment in each stride when all four legs are off the ground.

The most common four-legged locomotion which you will observe is the walk, sometimes called the diagonal walk. This is used by most hoofed animals, as well as cats and dogs. In this walk, the animal uses diagonally opposing legs, i.e. front left and right back legs to move forwards; then the front right and left back legs move, and so on.