LAB T O P I C 3 1 Investigating the Properties of Muscle and Skeletal Systems Supplies You should be able to describe in your own words the following concepts: Preparator's guide available on WWW at http://www.mhhe.com/dolphin Equipment Compound microscopes Intellitool Setups (Batavia, IL) Physiogrip and software Isolated, square wave stimulator Computer Sliding filament theory of muscle contraction How a nerve impulse causes a muscle to contract Basic structure of bone As a result of this review, you most likely have questions about terms, concepts, or how you will do the experiments included in this lab. Write these questions in the space below or in the margins of the pages of this lab topic. The lab experiments should help you answer these questions, or you can ask your instructor during the lab. Materials Fetal pig Dissection pans and instruments Millipedes, spiders, crabs, or insects Bird skeleton Human skeleton Frog skeleton Fresh beef long bones split longitudinally Dry long bones Prepared slides Skeletal muscle, longitudinal Smooth muscle section Cardiac muscle, longitudinal Bone Hyaline cartilage Tendon Earthworm, cross section Neuromuscular junction (demonstration) Student Prelab Preparation Before doing this lab, you should read the introduction and sections of the lab topic that have been scheduled by the instructor. You should use your textbook to review the definitions of the following terms: antagonistic muscles appendicular skeleton axial skeleton endoskeleton exoskeleton extensor flexor hydrostatic skeleton 31-1 pelvic and shoulder girdles sarcomere smooth, cardiac, and skeletal muscle summation tetany threshold effect twitch Objectives 1. To compare the microanatomy of skeletal muscle to that of smooth and cardiac muscle 2. To investigate the surface muscles in the fetal pig 3. To investigate the physiological concepts of threshold, recruitment, and temporal summation during muscle contraction 4. To view examples of hydrostatic skeletons and exoskeletons 5. To observe the microanatomy of bone 6. To compare the endoskeletons of several vertebrates Background Simple small organisms, such as bacteria, algae, protozoa, and sponges, are capable of movement without muscle systems. They use cilia or flagella to move through their aqueous environments. All higher animals, whether aquatic or terrestrial, depend on muscle systems for movement and on nervous systems to control that movement. These muscle systems are also intimately associated with the skeletal system, which converts muscular contraction into effective movement and locomotion. 417 Figure 31.1 Antagonistic muscle arrangements in human limb. Contraction of human biceps flexes the forearm, while contraction of the triceps extends the forearm. Origins Scapula Origins Flexors: Biceps brachii m. Extensors: Long head of triceps brachii m Brachialis m. Lateral head of triceps brachii m Medial head of triceps brachii m Insertion Creek Radius Ulna During contraction, actin and myosin filaments interact with each other, shortening the muscle in much the same manner as a deck of cards is aligned after it is shuffled (see fig. 31.4). The force for movement comes from the interaction of the filaments themselves rather than from a shortening of the filaments. Along with skeletal muscle, two other types of muscle tissue are found in animals. They function in movements within the body, rather than in movement of the body from one location to another. Smooth muscle tissue, also called involuntary muscle, is found in the gut, blood vessels, pupil, and reproductive system. Cardiac muscle is found in the heart. An integral part of any animal's locomotory system is its skeleton. While the word skeleton usually brings to mind the bony endoskeleton of vertebrates, other types of skeletons occur in the animal kingdom. Hydrostatic skeletons are found in earthworms and other soft-bodied invertebrates in which the pressure of internal fluids gives shape and allows the organism to move. Exoskeletons are found among such diverse animals as nematodes, arthropods, molluscs, and corals. This tough outer covering protects the organism and supports it in an upright position while allowing for muscle attachment and movement. Skeletons reflect adaptations to an organism's environment and lifestyle. Frogs and birds not only have limbs modified to fit their environments, but also have bones that show either thickening or shortening to fit their forms of locomotion. Consider the skeletal adaptations of a human, horse, bat, and seal to their respective forms of locomotion. Careful study of the skeleton by the trained observer can reveal much about an animal, because skeletons are excellent examples of the biological principle that form reflects function. Skeletons serve many other functions besides locomotion. They protect soft tissue, serve as a reservoir for calcium and phosphate, and are sites of blood cell formation. Skeletons are often the only parts of animals that survive from the past as fossils. Because the muscles attach to the skeleton, it is possible to see which muscles might have been highly developed and to speculate about how an extinct animal lived on a day-to-day basis. Muscles are nearly always found arranged as antagonistic pairs. As a result, contraction of one member of the pair causes an action, while contraction of the second member restores the body to its original position. In earthworms, the longitudinal and circular muscles in the body wall are antagonists. Contraction of the circular muscles elongates the segments by forcing the fluid of the hydrostatic skeleton forward and back in each segment, but contraction of the longitudinal muscles shortens the segments, leading to an increase in girth. In antagonistic pairs, neuronal activity is such that the contraction of one antagonist is usually accompanied by the relaxation of the second. Figure 31.1 shows the antagonistic arrangement of human muscles associated with an internal skeleton. Muscles that increase the angle between two bones at a joint are called extensors. Flexor muscles are antagonistic to extenwmmmmmmmmmffmmmmmt sors and decrease the angle between two bones. StraightenYou will study the anatomy and properties of ing your elbow is extension and bending it is flexion. and compare several types of skeletal systems About 80% of the mammalian body mass is muscle. Most of this is skeletal muscle (sometimes called striated or voluntary muscle). Skeletal muscle is characterized by a high degree of cellular organization. Muscle proteins, Muscular System consisting mostly of actin and myosin, occupy most of the cytoplasm of the cell. These proteins are arranged in a seMicroscopic Anatomy of Muscle ries of parallel filaments. Many cylindrical muscle cells make up a muscle, and the contractile proteins in these Ty Obtain prepared slides of smooth muscle, cardiac muscle, fibers are arranged in such a way that it gives a striated and a longitudinal section of skeletal muscle. Compare the microscopic pattern to the muscle. tissues on your slide to figure 31.2. 418 Investigating the Properties of Muscle and Skeletal Systems 31-2 Figure 31.2 Three Types of Muscle, (a) Smooth muscle has spindle-shaped cells, each with one nucleus (t) Skeletal muscle cells are cylindrical and have many nuclei, (c) Cardiac muscle is unique to the heart. Its striated cells join at connections called intercalated disks. The cylindrical cells branch. Cardiac and skeletal muscle cells are striated due to the orderly organization of contractile proteins. Smooth muscle (a) Smooth Muscle Observe the smooth muscle slide first. Note the shape of individual cells, the presence of nuclei, and the absence of filament organization in the cytoplasm. These muscle cells are spindle shaped, and their actin and myosin are not organized in parallel arrays. Smooth muscle is innervated by the autonomic nervous system and is not under voluntary control. It responds to various hormones. Its response is characterized by slow, rhythmic contractions. In some cases, contractions arise spontaneously in smooth muscles and are propagated along the length of the organ. Sketch two or three examples of smooth muscle cells. Nucleus -Muscle fiber H 10 jam Skeletal muscle (b) 10 nm Cardiac muscle (c) D- Skeletal Muscle Now examine the slide of skeletal muscle under the medium-power objective and compare its organization to that of smooth muscle. Note the substantial differences. Identify the individual cylindrical fibers that run the length of the tissue specimen on the slide. Find a very thin area of the section that is only one fiber thick and Nucleus examine it under high power. On the periphery of each fiber, you will see several nuclei. Each skeletal muscle Muscle fiber is multinucleated because it is formed by the fusion fiber of numerous smaller, uninucleated cells during embryonic development. Thus a muscle fiber is really a composite of several smaller cells. Though probably not visible, several mitochondria are also present in this peripheral cytoplasmic area. The central area of the cell consists of many parallel fibers called myofilaments running lengthwise in the cell. These fibers give a cross-banding appearance to the cytoplasm. Draw two or three of these cells. Nucleus 10 urn 31-3 Muscle fiber Intercalated disk Investigating the Properties of Muscle and Skeletal Systems 419 To understand how your drawing relates to the strucFigure 31.3 Scanning electron micrograph showing ture of a whole muscle, such as the biceps in your upper axon ending at the neuromuscular junction on a muscle cell. arm, look at figure 31.4. What most people call a muscle contains thousands of muscle fibers arranged parallel to one another and surrounded by a sheath of connective tissue. Within a single muscle cell are many myofibrils, which Axon consist of contractile units called sarcomeres joined end to end (fig. 31.4). Each sarcomere contains two types of filamentous proteins. Actin is the protein in thin filaments and myosin is the protein in thick filaments. The actin filaments at each end are anchored in the Z discs, which mark the ends of the sarcomere. The myosin filaments are suspended between and surrounded by actin filaments. It is the interdigitation of these filaments and the areas of overlap that create the alternating light and dark banding patterns that you saw on your slide of skeletal muscle. During contraction, the thin filaments slide toward the middle over the thick filaments, pulling the Z discs inward and shortening the overall length of the sarcomere. Because this process is repeated simultaneously in each sarcomere, Neuromuscular junctions muscle cells can dramatically shorten. In recent years, research has revealed the molecular mechanisms involved in muscle contractions. When a nerve impulse arrives at the neuromuscular junction, it causes the release of a chemical called acetylcholine (fig. 31.3). It depolarizes the muscle cell membrane and triggers an acLook at a slide of cardiac muscle through your comtion potential in the muscle cell. As the action potential pound microscope. Would you say that it is similar to moves along the muscle cell, it penetrates to the interior of the cell via the T tubule system (fig. 31.4). Membrane de- & smooth muscle or to striated muscle? Why? polarization at the Z disks causes the sarcoplasmic reticulum to release calcium ions into the sarcomere. The calcium ions allow the actin to react chemically with the myosin filaments. This pulls the actin filament over the surface of the myosin filament. The coordinated effect of the interaction between actin and myosin is the shortening of the muscle. Look at the demonstration slide of a neuromuscular A junction that is available in the lab. What do you find interV esting about this slide? Use your 43x objective and look closely at one muscle fiber. Move the slide so that you can follow a single fiber for some distance. You should find two things in cardiac muscle that are different from what you saw in striated muscle. What are these differences? Cardiac Musc/e 4>As the name implies, cardiac muscle is found in the heart, where it makes up the mass of the heart wall along with connective and nervous tissue. It is capable of rhythmic contraction which can be modulated by the pacemaker of the heart. The contraction of heart muscle is not under voluntary control. 420 The actin and myosin components in cardiac muscle are found in interdigitating linear arrays, as in skeletal muscle, but the fibers are capable of spontaneous contraction, as in smooth muscle. The muscle cells are shorter than in Investigating the Properties of Muscle and Skeletal Systems 31-4 Figure 31.4 Skeletal muscle structure and function. A muscle fiber or cell contains many myofibrils, consisting of sarcomeres. When the myofibrils of a muscle fiber contract, the sarcomere shortens as the actin filaments move toward the center of the sarcomere and Z discs move closer. bundle of muscle fibers T tubules nucleus sarcoplasmic reticulum calcium storage sites sarcoplasm skeletal muscle fiber one myofibril A muscle fiber has many myofibrils. sarcolemma A myofibril has many sarcomeres. cross-bridge myosin actin 31-5 Investigating the Properties of Muscle and Skeletal Systems 421 skeletal muscle, often branch, and are joined end to end by tight junctions, called intercalated disks, which electrically couple the cells and allow contraction to spread from cell to cell independent of the nervous system. 6 Starting at the head, identify the major muscles indicated in figure 31.5. 1. The latissimus dorsi is a broad muscle running obliquely around the sides of the thoracic region. It originates on the vertebrae and inserts on the proximal end of the humerus. The latissimus dorsi is involved in moving the foreleg. Fetal Pig Superficial Muscles 2. The trapezius originates on the occipital bone of the skull and from the first ten vertebrae and inserts on the scapula or shoulder blade. This broad muscle draws the scapula medially. When this muscle contracts, how does the leg move? This dissection may be done by students or by the instructor as a demonstration, depending on the time available and the emphasis of the course on gross anatomy. Three terms are used to describe the anatomical orientations of muscles: origin, belly, and insertion. The origin is the end attached to the less mobile portion of the skeleton. The belly is the central part of the muscle, and the insertion is the end of the muscle attached to the freely moving part of the skeleton. Tendons are fibrous connective tissues that connect muscles to the skeleton. If your fetal pig is not already skinned, skin it—being careful not to tear away muscle as you remove the skin. Use a probe or finger to separate the two. Under the skin, there may be adipose (fat) tissue, which should be removed to reveal the muscles. When the skin is removed, dry the carcass with paper towels to improve viewing. In young fetal pigs, the muscles are not fully developed and are often tightly connected to the skin so that they are torn during skinning. Identification will be difficult. Figure 31.5 The major muscles of the fetal pig. Deltoideus Latissimus dorsi Trapezius Tensor fasciae latae Gluteus medius Biceps femoris Position of gastrocnemius Brachiocephalic Sternocephalicus " ' Transversus Tendon to gastrocnemius Internal oblique' 422 Investigating the Properties of Muscle and Skeletal Systems 31-6 • §-*> « 3. The brachiocephalic muscle extends obliquely as a flat belt from the back of the skull (the occipital bone) to the foreleg (distal end of the humerus). It is involved in moving the leg anteriorly. 4. The sternocephalic is a long muscle below the brachiocephalic muscle. It controls the flexing of the head. It originates on the sternum and inserts on the mastoid process of the skull by means of a long tendon. Remove the parotid salivary gland to see the tendon. 5. The brachialis is a small muscle located in the angle formed by the flexed foreleg. It originates on the humerus and inserts on the ulna. What is its function? 13. The biceps femoris is a large muscle making up most of the back half of the thigh. It originates on the pelvis and inserts on the lower femur and upper part of the tibia. 14. The gastrocnemius is the large muscle of the calf, originating on the lower end of the femur and attaching to the heel by means of the Achilles tendon. Its action extends the foot. The soleus muscle lies close to the gastrocnemius and has a similar function. 15. The digital flexors and extensors originate on the tibia and fibula and insert on the metatarsals. What are their functions? 6. The deltoid is a broad shoulder muscle originating on the scapula and inserting on the humerus. It aids in flexing the humerus. 7. The extensors are several muscles in the lower foreleg that extend and rotate the wrist and digits. 8. The triceps (brachii) is a large muscle making up practically the entire outer surface of the forelimb. It originates on the humerus and inserts on the proximal end of the ulna. When it contracts, the forelimb extends. 9. The external oblique muscles make up the outer wall of the abdomen and run obliquely from the ribs to a ventral longitudinal ligament along the ventral midline. Contraction of these muscles constricts the abdomen. 10. The internal oblique fibers lie just under the external oblique muscle and run almost at right angles to them. Contraction of these fibers also results in abdominal constriction. 1 1 . The tensor fasciae latae is the most cranial of the thigh muscles. It originates on the pelvis and continues ventrally as a thin, triangular muscle until it becomes a sheet of connective tissue called the fasciae latae, which inserts on the kneecap and extends the leg. 12. The gluteus medius is a thick muscle covered by the tensor fasciae latae in the hip region. If the overlying muscle is removed, its origin on the hip and insertion on the femur can be seen. What action does it cause? 16. The peroneus muscles have origins and insertions similar to digital muscles. They are involved in moving the whole foot. 31-7 Physiology of Muscle This part of the lab may be performed by the instructor as a demonstration or by students working in groups of four or more, depending on the time and equipment available. Overview of Experiment In this section of the lab, you will investigate four properties of human muscles: threshold stimulus, recruitment, temporal summation, and tetany. You will use special equipment and software that allows a computer to collect and analyze data. The experimental subject(s) will be a volunteer member(s) of your class. Anyone who has heart problems or does not wish to participate directly in this experiment should not be a subject. They can operate the computer or record notes during the experiment. To do this experiment, you need to make adjustments to four major pieces of equipment—a stimulator, a transducer, an interface unit, and a computer running special software. A stimulator is an electronic device that produces a pulse of electricity that can be varied by volts (strength), duration of the pulse (length), and frequency (number of pulses per second). The transducer is a device that converts the movement of a trigger in a pistol handle into an electrical resistance that can be recorded. The interface is a box that connects various cables together into a single lead that is connected to a port of your computer. The software allows your computer to record raw data from the transducer and process the data, converting it into graphical displays. Before you start investigating the biology of muscle, this equipment must be calibrated. Investigating the Properties of Muscle and Skeletal Systems 423 the deflection of the trace on the computer screen as recruitment occurs? Use the same setup as before. Be sure to use NEW from the FILE menu to select a new data file. Start with the previously determined threshold voltage and record a twitch. Slowly increase the voltage and continue recording twitches. Continue this procedure until a maximum height of contraction is reached and further increments of stimulus intensity do not evoke stronger contrac& tions. What is the voltage at which no further increase in strength of contraction occurs?_ Turn off the stimulator. If excitable cells exhibit an all-or-none response when exposed to a threshold stimulus, explain why the height of contraction increased with stimulus strength above the threshold? (Hint: Consider the phenomenon of recruitment.) Temporal Summation and Tetany far in your investigations of muscle you have used a low frequency of stimulation. In this section, you will investigate the effects of increasing frequency of contraction on the strength of muscle contraction. With increasing frequency of contraction, there is insufficient time for the muscle to relax before the next stimulating impulse arrives. Consequently, the contractions begin to summate with a new contraction starting in a partially contracted muscle. This phenomenon is called temporal summation, and when the muscle is fully contracted and cannot relax between stimuli, tetany occurs. What would you hypothesize is the effect on the strength of muscle contraction? State your hypothesis. Once this base is established, gradually increase the frequency of stimulation over a 10-second interval. Can you see individual contractions at low frequencies? At higher frequencies? What happens to the strength of contraction as the frequency of stimulation increases? If the strength of contraction is so strong that the trigger reaches the end of its travel, stop the experiment and change the spring in the Physiogrip to a stronger one. Erase the previous trace from memory in the computer and repeat the experiment. When the strength of contraction no longer increases with increasing frequency of stimulation, stop the experiment and turn off the stimulator. Analyze the data that you have collected in the following way. First, print a copy of the screen by choosing PRINT from the FILE menu. This will provide a record for any report that you may have to write. From the FILE menu, select ANALYZE and then DISPLACEMENT. You should see a record of the contractions on the screen and at the top should be a series of hash marks indicating how often a stimulus was given. Using the ANALYZE cursor, determine the frequency of stimulation just before temporal summation started. What is the value in number per second? Now repeat this measurement at the point where tetanic contraction was reached. What is the frequency of stimulation that results in tetany? Measure the height of an individual twitch at the beginning of this experiment. What is it? Now measure the height of contraction reached in tetany. What is it? What is the percent change due to tetany? Skeletal Systems Skeletal systems support and protect an animal and make locomotion possible. In some animals, especially those with endoskeletons, the skeleton is a storage site for calcium and phosphate ions needed in metabolism. Three types of skeletal systems are found in the animal kingdom: hydrostatic (sometimes called hydraulic), exoskeletons, and endoskeletons. Types of Skeletons Hydrostatic Skeletons Test your hypothesis, using the following procedure. Choose NEW from the FILE menu to create a new data file. Turn on the stimulator. Have the subject grasp the Physiogrip and put a slight pressure on the trigger to raise the baseline a few millimeters. Set the stimulus parameters to 10 msec duration, frequency to 1 per sec in continuous mode, and a voltage that gives a single contraction. 426 Many soft-bodied invertebrates, such as roundworms, cnidarians, and annelids, do not appear to have specialized, differentiated skeletal systems. However, close examination reveals that the body wall muscles act on the incompressible fluids in the body cavity and intracellular spaces to facilitate very effective movement and support. Such skeletal systems are called hydrostatic skeletons. Investigating the Properties of Muscle and Skeletal Systems 31-10 Obtain a microscope slide of a cross section of an earthworm. Look at it first under the low-power objective and note the general anatomy of the animal by comparing the slide to figure 21.9. The external surface of the earthworm is covered by a highly flexible cuticle secreted by the underlying cells of the dermis. The fibrous proteins of the cuticle protect the worm and help maintain its form. Some cells in the dermis secrete mucus, which lubricates the passage of the earthworm through the soil and also maintains a moist surface through which respiratory gases are exchanged. Note that the body wall is made up of two layers of muscles: an outer circular set and a featherlike inner longitudinal set. Since the longitudinal set runs parallel to the long axis of the body, it will appear in cross section in the slide. Also note the well-developed body cavity, which is filled with fluid in live animals. What happens to the worm's shape when the circular muscles contract? (Consider the effects of both the muscles and internal fluids.) Exoskeletons Exoskeletons are characteristic of several animal phyla. In addition to supporting the animal, exoskeletons prevent water loss, protect the organs, and serve as points of attachment for muscles. Animals in the phylum Arthropoda, which includes crayfish, insects, spiders, and millipedes, have a well-developed exoskeleton composed of a complex polysaccharide called chitin combined with proteins to give flexibility. Several examples of these animals should be available in the laboratory. Look at an insect through your dissecting microscope and note the segmentation of the exoskeleton. The head and thorax are composed of several fused body segments. In insects, the thorax bears the walking legs and wings. The segments should be clearly visible in the abdomen. An insect's exoskeleton is made up of hardened plates called sclerites. In the head and thorax, the sclerites are rigidly joined to one another along suture lines. In the abdomen, the dorsal and ventral sclerites are enlarged and joined to one another by pleural membranes, which correspond to thin, reduced vestiges of the lateral sclerites. Similar membranous areas are found between adjacent dorsal or ventral sclerites. These thin, flexible areas allow the animal to expand, contract, and curl its abdomen. Flexible membranous areas are also found between the joints of the various appendages. What happens to the worm's shape when the longitudinal muscles contract? (Consider the effects of both the muscles and internal fluids.) The body cavity of the earthworm is divided into compartments, corresponding to each segment, by cross walls called septa. These prevent fluids from "sloshing" from one end of the worm to the other and allow earthworms to make a variety of movements with fine gradations. Although they are not part of the hydrostatic skeleton, you should note the spinelike setae in the body wall. Muscles attached to each seta extend or retract them. When extended they project into the soil and anchor the worm. Describe how you think an earthworm could crawl forward using its setae, circular muscles, and longitudinal muscles. Endoskeletons Defined as an internal supporting system of hardened material, endoskeletons are highly characteristic of vertebrates. Cartilage and bone make up the vertebrate endoskeleton. You will start your study of endoskeletons by examining the structure of bone. Bone Structure xamine a fresh beef femur that has been cut longitudinally in half. The central shaft of a long bone such as this one is called the bone's diaphysis, while the enlarged ends are the bone's epiphyses (fig. 31.8). Though difficult to observe in 31-11 Investigating the Properties of Muscle and Skeletal Systems 427 Figure 31.8 Structure of a bone: (a) a bone in partial longitudinal section; (b) scanning electron micrograph of cancellous (spongy) bone; (c) scanning electron micrograph of Haversian canal (HC) systems in compact bone. Canals allow blood vessels to pass into bone. Bone cells are located in the lacunae (La), (c) From R.G. Kessel and R.H. Kardon. Tissues and Organs: A Text-Atlas of Scanning Electron Microscopy. 1979. W.H. Freeman and Co. Epiphyseal disks Articular cartilage — Proximal epiphysis Spongy bone Space occupied by red marrow Compact bone Medullary cavity — Diaphysis Yellow marrow (b) Remnant of epiphyseal disk Spongy bone Compact bone Periosteum — - Distal epiphysis (a) Femur older bones, a narrow zone of cartilage, the epiphyseal disk (growth center) of the bone, separates the epiphysis from the diaphysis. The rounded projection from the upper end of the bone (fig. 31.8a) is called the head, and it articulates with the acetabulum, a depression where all three pelvic bones intersect. Condyles (rounded, articular prominences) are located on the lateral and medial sides of the distal end of the femur. With which bone(s) do they articulate? 428 Find the periosteum, the tough connective tissue layer surrounding the diaphysis. The surfaces of the condyles are covered by a glassy smooth articular cartilage, which, along with synovial fluid, reduces frictional erosion between articulating bones at the joints. Residues from the ligament that held the joint together may also be visible. On the outer surface of the bone, find the narrow ridges called crests and the small, round, elevated tubercles, where muscles attach by tendons to the bones. The surface of the bone is perforated by openings called foramina (sing, foramen) that allow blood vessels and nerves to penetrate the bone. The center of the bone is a hollow chamber called the medullary cavity. It is filled with marrow. Red marrow is where new red blood cells are formed, but in older animals much of the marrow is fatty yellow matter, which no longer produces blood cells. Investigating the Properties of Muscle and Skeletal Systems 31-12 Compare the structure of the bone in the diaphysis with that in epiphysis. Compact bone is very dense, whereas spongy bone has many interior cross braces. Spongy bone is less dense than compact bone. Microanatomy of Bone V4>Obtain a prepared slide of ground bone and examine it under low power. Using figure 31.8c as a guide, find a Haversian canal. These canals allow nerves, blood vessels, and lymphatics to penetrate bone. Note the chambers, called lacunae, arranged in concentric circles around the canal. Osteocytes, mature bone cells, line in these spaces and are responsible for depositing and removing calcium salts. Their activity is regulated by hormones. Note the canaliculi that radiate from the lacunae. They are transportation networks and allow osteocytes to extend their cytoplasm and form intimate contact with the bone matrix. Figure 31.9 (a) Skeleton of frog (ventral view), including (b) ventral view of pectoral girdle. Vomerine teeth Sphenethmoid Squamosal Maxillary Mandible Orbit Phalanges Metacarpals Carpal Episternum Suprascapula Scapula Radioulna Sacral vertebra Comparative Vertebrate Endoskeletons The vertebrate skeleton has two components: the axial skeleton, consisting of the skull and vertebral column; and the appendicular skeleton, which includes the pelvic and pectoral girdles as well as the bones of the appendages. Skeletal Comparisons the laboratory, you will observe the skeletons of three vertebrates: a frog, a bird, and a human (figs. 31.9-31.11). The task before you is to determine how these skeletons are similar and how they are different. For the differences, you should offer explanations of how these differences correlate with locomotion or life style. For each skeleton you should be able to quickly identify the axial and appendicular skeletons. Then you should be able to find and identify the bones listed below. Axial skeleton Skull cranium (fused bones encasing brain and sense organs) consisting of the following major bones: frontal parietal temporal occipital facial bones consisting of the following major bones: zygomatic maxilla mandible (lower jaw), the only movable bone in skull Vertebral column, which supports and protects the spinal cord. Cartilaginous discs are found between the vertebrae in living animals. The vertebral column is divided into five regions: cervical vertebrae (neck) thoracic vertebrae (on which the ribs articulate) lumbar vertebrae (abdominal region) 31-13 Tibiofibula Astragalus"! Calcaneumf Metatarsals Phalanges (a) Suprascapula Episternum Clavicle Scapula Glenoid fossa Coracoid Sternum Xiphoid process (b) Investigating the Properties of Muscle and Skeletal Systems 429 Figure 31.10 Skeleton of a bird. ** Parietal Supraoccipital Frontal Orbit Atlas Nasal Cervical vertebrae Maxilla Premaxillary Metacarpals Mandible Phalanges Hyoid apparatus Radius Ulna Humerus Scapula Rib Clavicle Coracoid Ilium Caudal vertebrae Ischium Pubis Tarsometatarsus Phalanges sacral vertebrae (enclosed by pelvic girdle) caudal vertebrae or tail (coccyx in human) Ribs (attached and floating) Sternum Xyphoid process Appendicular skeleton Pectoral girdle consisting of: clavicle (collarbone) scapula (shoulder blade) coracoid Forelimbs consisting of: humerus (long bone of lower limb) radius (long bone of lower limb, which forms a pivot joint with the ulna and is also part of the hinge joint of the elbow) ulna (other long bone of lower arm, forming hinge joint at elbow) 430 Investigating the Properties of Muscle and Skeletal Systems carpals (small bones of wrist) metacarpals (bones of palm) phalanges (finger bones) Pelvic girdle consisting of: ilium ischium pubis Hind limbs consisting of: femur (long bone of thigh) patella (kneecap) tibia (larger of two long bones of lower limb) fibula (smaller long bone of lower limb) tarsals (bones of ankle) calcaneus (heel bone) metatarsals (slender foot bones) phalanges (toe bones) 31-14 Figure 3 1 . T l Human skeleton. Occipital bone Skull— Shoulder girdle Clavicle -Scapula Costal cartilages Pelvic girdle Coxa— bone Phalanges 31-15 Investigating the Properties of Muscle and Skeletal Systems 431 Compare a bird skeleton and a human skeleton. List w List the skeletal differences between a frog and a five skeletal differences associated with locomotion and exhuman that are associated with locomotion. plain how each is involved in locomotory movement. Differences will include fusions, increase in size, reduction in size, but not the appearance of new bones. 3. Both recruitment and temporal summation lead to greater force of muscle contraction. Explain what is happening in both of these phenomena. 4. Describe the structure of a bone, such as the humerus, at the macroscopic and microscopic levels. 5. Describe the major common features of skeletons found in all vertebrates. 6. "Form reflects function" is a statement that applies to the skeletal system. How does the form of the vertebrate skeleton relate to the means of locomotion in a frog compared to a human? In a bird compared to a human? Learning Biology by Writing Write a lab report that describes your experiments with human muscles. It should have three sections in the results: Determining threshold Demonstrating motor unit recruitment Demonstrating temporal summation and tetany The discussion section should describe how recruitment and summation are important in graded responses such as are used in sports or playing a musical instrument. Your instructor may ask you to answer the Lab Summary and Critical Thinking Questions that follow. Critical Thinking Questions Internet Sources The disease rickets affects the human skeletal system. Use GOOGLE to search the WWW to learn what this disease is. Where is it common? Is it found in the U.S.? How can it be treated? Lab Summary Questions 1. What evidence do you have that smooth, cardiac, and skeletal muscle cell are different? Describe. Give examples of where you would find each in your body. 2. Describe the all-or-none response in muscle cells. How is it possible for us to have graded responses in our muscles if muscle cells respond in an all-or-none manner to stimuli? 432 Investigating the Properties of Muscle and Skeletal Systems 1. About 99% of the body's calcium is found as calcium phosphate salts in bone tissue. The calcium in plasma is in an ionic form, Ca++. Although the level of Ca++ in the blood and tissues is closely regulated by parathyroid hormone and calcitonin, imbalances in Ca++ can occur. What effect would an elevated Ca++ concentration have on muscle activity? What effect would a Ca++ deficiency have on muscle activity? 2. As can be seen in figure 31.10, the skeleton of a bird is modified for flight. Aside from the size and arrangement of the bones in the skeleton, what other feature of its bones might affect the ability of a bird to fly? 3. How do scientists deduce the appearance of hominids or dinosaurs when all they have to study are a few bone fragments? 31-16