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    Human Anatomy and Physiology SECOND EDITION Elaine N. Marieb, R.N., Ph.D. Holyoke Community College LC) The Benjamin/Cummings Publishing Company, Inc. Redwood City, California + Menlo Park, California Reading, Massachusetts + New York + Don Mills, Ontario Wokingham, U.K. + Amsterdam + Bonn + Sydney Singapore + Tokyo + Madrid + San Juan SwMieng |iSSue: The Living Fabric oo nt Chapter Outline and Student Objectives erent ees aren ic) Epithelial Tissue (pp, 104-113) 4, List several characteristics that typify epithelial tissue, 2 Describe the criteria used to classify epithelia structurally. 3, Name and describe the various types of epithelia; also, indicate their chief function(s) and location(s) 4. Compare endocrine and exocrine glands relative to thoir general structure, product(s), and mode of secretion, 5, Describe how multicellular exocrine glands are classi- fied structurally and functionally. Connective Tissue (pp. 114-126) 6, Describe common characteristics of connective tissue. 7, List the structural elements of connective tissue and describe each element. 8. Explain the bases for classification of connective tissue. 9, Describe the types of connective tissue found in the body, and indicate their characteristic functions. Epithelial Membranes (pp. 126-127) 10, Contrast a sheet of covering or lining epithelium with an epithelial membrane, Name and describe the three varieties of epithelial membranes. Muscle Tissue (p. 129) 11, Compare and contrast the structures and body locations of the three types of muscle tissue, Nervous Tissue (pp. 129-130) 412, Note the general characteristics and functions of ner: vous tissue, Tissue Repair (pp. 130-132) 13. Describe the process of tissue repair involved in the normal healing of a superficial wound. Developmental Aspects of Tissues (pp. 132-133) 14, Name the three primary germ layers and indicate the primary tissue classes arising from each. 15. Briefly mention tissue changes that occur with age. Cite possible causes of such changes, ( ells that exist as isolated unicellular (one-cell) organisms, such as amoebas, are rugged indi- vidualists, They alone obtain and digest their food, carry out gas exchange, and perform all the other activities necessary to keep themselves alive and healthy, But in the multicellular human body, cells do not operate independently. Instead, they form tight cell communities that function cooperatively. Indi- vidual cells are specialized, with each type perform- ing specific functions that help maintain homeostasis and benefit the body as a whole. The specialization of our cells is obvious: Muscle cells look and act quite differently from skin cells, which in turn are easily distinguished from cells of the brain. Cell specialization allows the various parts of the body to function in very sophisticated ways, but this division of labor also has certain hazards. When a par- ticular group of cells is indispensable, its loss can severely disable or even destroy the body. Groups of closely associated cells that are similar in structure and perform a common function are called tissues (ssw = woven). Four primary tissue types interweave to form the “fabric” of the body: epi- thelial (ep"i-the’le-ul), connective, muscle, and ner- vous tissue, and each has numerous subclasses or varieties. If we had to assign a single term to each pri- mary tissue type that would best describe its overall role, the terms would most likely be covering (epithe- lial), support (connective), movement (muscle), and contro! (nervous). However, these terms reflect only a fraction of the functions that each tissue performs. As explained in Chapter 1, tissues are organized into organs such as the kidneys and the heart, Most, organs contain representatives of all four tissue types, 103 404 Umitone Opponization ofthe Body. and the arrangement of the tissues determines the ongan’s structure and capabilities. The study of tis- sues, or histology, complements the study of gross anatomy. Together they provide the structural basis for understanding organ physiology. Furthermore, learning the characteristic patterns of each tissue type will allow you tu predict the function of an organ once its structure is known, and vice versa. TR Epithelial Tissue Epithelial (epithe ~ laid on, covering) tissue, or epi- thelium, occurs in the body as (1) covering and lining epithelium and (2) glandular epithelium, Covering and lining epithelium is found on all free surfaces of the body such as the outer layer of the skin, dipping into and lining the open cavities of the digestive and respiratory systems, lining blood vessels and the heart, and covering the walls and organs of the closed ventral body cavity. Since epithelium forms the inter- faces or boundaries that mark us off from the outside world, nearly all substances received or given off by the body must pass through epithelium. Glandular epithelium fashions the glands of the body. Epithelium is highly specialized to accomplish many functions, including (1) protection, (2) absorp- tion, (3) filtration, (4) excretion, and (5) secretion. Each of these functions is described in detail as we consider various types of epithelium, but briefly, the epithelium of the skin protects underlying tissues from mechanical and chemical injury and bacterial invasion; that lining the digestive tract is specialized to absorb substances; and that found in the kidneys performs nearly the whole functional “menu” — absorption, secretion, and filtration. Secretion is the specialty of glands. Special Characteristics of Epithelium Specie See eee Epithelial tissues have many characteristics that dis- tinguish them from other tissue types. 1. Cellularity. Epithelial tissue is composed almost entirely of cells, The cells are close and only a tiny amount of extracellular material lies in the narrow spaces between them, 2. Specialized contacts. Epithelial cells fit close together to form continuous sheets. Adjacent cells are bound together at many points by lateral con- tacts, including tight junctions and desmosomes (see Chapter 3) . Polarity. Epithelium always has one free surface, 3. PoltTt the apical surface, which is exposed to the body exterior or the cavity of an internal organ Some exposed plasma membrane surfaces are gmooth and slick; others exhibit cell surface mod. fications such as microvilli ot cilia. Microvilli, fingerlike extensions of the plasma membrane, are found on most epithelial surfaces and tremen. dously increase the exposed surface area. In epi- thelia that absorb or secrete substances (intestinal lining and kidney tubules), the microvilli are often so dense that the cell apices have a fuzzy appear- ‘nce referred to as a brush border. Cilia projecting from the epithelial lining of the trachea and certain other internal tracts propel substances along the epithelial surface. “Avascularity (a-vas’ku-lar'I-te). Although epithe- ium may be well supplied by nerve fibers, it is avascular (contains no blood vessels). Epithelial cells are nourished by substances diffusing from ‘blood vessels in the underlying connective tissue. 5. Basement membrane. The lower or basal surface of an epithelium rests on a thin supporting basal lamina (ba’sal lam‘i-nah), which separates it from the underlying connective tissue. The basal lamina is a nonliving, adhesive material formed largely of glycoproteins secreted by the epithelial cells superficial to it. The connective tissue cells, just deep to the basal lamina, secrete a similar extracel- lular material containing fine collagenous fibers; this layer is called the reticular (ré-tik'u-lar) lam- ina. The basal lamina of the epithelium and the reticular lamina of the connective tissue together form the basement membrane. The basement membrane reinforces the epithelial sheet, helping it to resist stretching and tearing forces, and defines the space that may be occupied by the epi- thelial cells. ‘An important characteristic of cancerous epi- thelial cells is their failure to respect this boundary, which they penetrate to invade the tis sues beneath. m 6. Regeneration, Epithelium has a high regenerative capacity. Some epithelia are exposed to friction, and their surface cells tend to abrade off. Others are damaged by hostile substances (bacteria, acids, smoke) in the extemal environment and die. As long as epithelial cells receive adequate nutrition, they can replace lost cells rapidly by cell division Classification of Epithelia ‘The many types of epithelia are identified structurally according to two criteria: the number of cell layer? @ Figure 4.1 Classification of epithelia. (a) Cisssiication on the basis of numberof layers. (b)Classifcaton on the bass of cell shape. For each category, @ whole celis shown onthe let and a longtucinal sections shown onthe right present and the shape of the cells (Figure 4.1). Based on the number of cell layers, there are two major vari- eties of epithelium. Simple epithelia, composed of a single layer of cells, are typically found where absorp- tion and filtration occur and where the thinness of the barrier helps to speed these processes. Stratified epi- thelia, consisting of multiple cell layers stacked one on top of the other, are common in areas of high abra- sion where protection is important, such as the skin surface and the lining of the mouth, ‘Three-dimensionally, all epithelial cells have six (somewhat irregular) sides. This polyhedral shape allows the cells to be so closely packed that a cross section of an epithelial sheet has the appearance of a honeycomb. However, epithelial cells vary in cell vol- ume and, consequently, in height. On the basis of height, there are three common shapes of epithelial cells, Squamous (skwa'mus) cells are flattened and scalelike (squam = scale); cuboidal (ku-boi’dahl) cells are approximately as tall as they are wide; and columnar (k6-lum'nar) cells are tall and column shaped. In each case, the shape of the nucleus con- forms to that of the cell. The nucleus of a squamous cell is disc shaped; that of a cuboidal cell is spherical; and a columnar cell nucleus is elongated from top to bottom and is usually located close to the cell base. Keep nuclear shape in mind when you attempt to identify epithelial types. The forms denoting shape and arrangement of epithelial cells are then combined to classify epithelia fully, as shown in Figure 4.1. The simple epithelia are easy to classify by cell shape, because all the cells in the layer have the same shape. There are four major Chapters Tissue; The Living Fabric 105 ‘Squamous Cuboidal Columnar () classes of simple epithelia: simple squamous, simple cuboidal, simple columner, and a highly modified simple epithelium called pseudostratified (pseudo = false). ‘There are also four major classes of stratified epi- thelia: stratified squamous, stratified cuboidal, strati- fied columnar, and a modified stratified squamous epithelium called transitional epithelium. In terms of body distribution and abundance, only stratified squamous and transitional epithelia are significant. In stratified epithelia, cell shape varies in the different layers. Thus, to avoid ambiguity, stratified epithelia are named according to the shape of the cells at the free surface, not according to deeper cell types. For example, the apical cells of a stratified squamous epi- thelium are squamous cells, but its basal cells are cuboidal or columnar. ‘As you read about the individual epithelial clas- ses and study their illustrations in Figure 4.2, keep in mind that tissues are three-dimensional, but that their structure is studied using stained tissue sections viewed through the microscope, Therefore, a cross- sectional view differs from a longitudinal view of the same tissue. Depending on the precise plane of the cut made when tissue slides are prepared, the nucleus of a particular cell may or may not be visible, and (frus- tratingly) the plasma membrane boundaries between epithelial cells are often indistinct. 106 Unit One Organization of the Body ‘Simple Epithelia ‘The simple epithelia are most concemed with absorp tion, secretion, and filtration. Because they are usu- ally very thin, protection is not one of their “specialties.” Simple Squamous Epithelium. The cells ofa simple squamous epithelium are flattened laterally, and their cytoplasm is sparse (Figure 4.2a). In a surface view, the close-fitting cells resemble a tiled floor; when cut perpendicular to their free surface, the cells resemble fried eggs seen from the side, with their cytoplasm wisping out from the slightly bulging nucleus. Thin and often permeable, this epithelium is found where filtration or the exchange of substances by rapid dif- fusion is a priority. In the kidneys, simple squamous epithelium forms part of the filtration membrane; in the lungs, it forms the walls of the air sacs across which gas exchange occurs. ‘Two simple squamous epithelia in the body have special names. Endothelium (en’do-the’le-um) pro- vides a slick, friction-reducing lining in lymphati vessels as well as in all hollow organs of the cardio- vascular system—blood vessels and the heart. The walls of capillaries consist exclusively of endothe- lium, where its exceptional thinness encourages the efficient exchange of nutrients and wastes between the bloodstream and surrounding tissue cells, Meso- thelium (mez’o-the'le-um) is the epithelium found in serous membranes lining the ventral body cavity and covering its organs. (The composition of the serous membranes is covered in more detail on p. 126.) Simple Cuboidal Epithelium. Simple cuboidal ¢ thelium consists of a single layer of cubical cells (Fi ure 4.2b). The spherical nuclei stain darkly, causing the layer to look like a string of beads when viewed ‘microscopically. Important functions of simple cuboi- dal epithelium are secretion and absorption. In glands, it forms both the secretory portions and the ducts that deliver secretions to their destinations. Some simple cuboidal epithelium in the kidney tubules has dense microvilli, betraying its active role in absorption. ‘Simple Columnar Epithelium. Simple columnar epithelium is seen as a single layer of tall, closely packed cells, aligned like soldiers in a row (Figure 4.2c), This epithelial type lines the digestive tract from the stomach to the rectum. Columnar cells are most associated with absorption and secretion, and the digestive tract lining has two distinct modifica- tions that reflect that dual function: (1) dense micro- villi on the apical surface of absorptive cells and (2) goblet cells that secrete a protective lubricating mucus. The goblet cells, named for their “goblet” shape, contain “cups” of mucus that occupy most of the apical cell volume (see Figure 4.3) ‘Some simple columnar epithelia display cilia on thoir free surfaces. This more unusual variety, called simple ciliated columnar epithelium, lines the uter ine tubes and limited areas of the respiratory tract Pseudostratified Columnar Epithelium. ‘The cells of pseudostratified (s00°do-stra'ti-fid) columnar epi- thelium are varied (Figure 4.2d). All ofits cells rest on the basement membrane, but some are shorter than others and, as seen in the figure, may not reach the surface of the coll layer. Their nuclei vary in shape and are located at different levels above the basement membrane, giving the false (pseudo) impression that Several cell layers are present, hence “pseudostrati- fied.” This epithelium, like simple columnar variety, functions in secretion or absorption. A ciliated ver- sion containing goblet cells (more precisely called. pseudostratified ciliated columnar epithelium) lines Inost of the respiratory tract. The mucus produced by the cells traps inhaled dust and other debris, and the motile cilia act to propel it superiorly away from the Tungs. Stratified Epithelia Stratified epithelia consist of two or more cell layers. Considerably more durable than the simple epithelia, their major (but not their only) function is protection. Stratified Squamous Epithelium. Stratified squa- ‘mous epithelium is the most widespread stratified epithelium (Figure 4.2e). Composed of several layers, itis thick and well suited for its protective role in the body. Its free surface cells are squamous; cells of the deeper layers are cuboidal or, less commonly, colum- nar, This epithelium is found in areas subjected to wear and tear, and its surface cells are constantly being rubbed away and replaced by mitotic division of the cells of its basal layer. Since epithelium depends on diffusion of nutrients from a deeper con- nective tissue layer, the epithelial cells farther from the basement membrane are less viable and those at the free apical surface are commonly flattened and atrophied. Stratified squamous epithelium forms the exter- nal part of the skin and extends a short distance into every body opening directly continuous with the skin, Therefore, it also covers the tongue and lines the mouth, pharynx, esophagus, anal canal, and vagina, The outer layer, or epidermis, of the skin is said tobe a keratinized (ker/ah-tin'tzd) stratified squamous epithelium because its surface sells contain keratin, & tough waterproofing protein. (The epidermis, which provides a protective covering for the body surface, discussed in Chapter 5.) The other stratified squ® ‘mous epithelia of the body are unkeratinized. Text continues np 111 Figure4.2. Epithelialtissues. Simple epithelia (a and). {a) Simple squamous epithelium Description: Single layer ol flationed cols with disk'shaped central ‘uclel and sparse cyto plasm: the simplest of the epithelia. Location: Air sacs of lungs: Kidney glomeruli lining of heart, blood vessels, and lymphatic vessels; lining of ventral body cavity (Serosae). Function: Allows passage of materials by dit- fusion and fitration in sites where protection, is not important; secretes lubricating substances in serosae. Photomicrograp squamous SaaS Eal la saan te a hee Nucleus Simple squamous epithelial cell aca alata a (©) Simple cuboidal epithelium Description: Single layer of cuboiike cells ‘with large, spherical ‘central nucle Location: Kidney tubules: ‘ducts and secretory portions ‘of small glands: ovary surface. Function: Secretion and absorption. Photomicrograph: Simple cuboidal epithelium in kidney tubules (260%) Simple — cuboidal epithelial cols Basement — membrane i BE ce t - o >, onnectve ac ; tue KLKO) 108 cnitOne Onganization of the Body Figure 4.2. (continued) Simple epithelia (c and ¢). {) Simple columnar epithelium eure See. eee ie Herein gal oat sgatlacder and excretory ‘ducts of some glands; cltsted ‘riety lines small bronchi, seo, ard se regions of the Fupeton: Absorption: secaon of mucus, eneymes, And other 8 patina cs (orrprocueive cots by ety acon. /TOe Siege & My (280%). Figure4.2 (continued) Stratified epithelia (e and). (6) Stratified squamous epithelium Description: Thick ‘membrane composed of Soveral call ayers: basal Cells ate cuboidal or Columnar and metabolically actve; surtace cells are tlattened (squamous); in the keratnized ‘ype, the surtace cells are full of keratin and dead; basal cells are ‘active in mitosis and produce the els ofthe more superficial layers. Location: Nonkeratinized type: forms the moist linings of the esophagus, mouth, and vagina; keratinized variety forms the epidermis of the skin, a dry membrane, Function: Protects underlying tissues in areas sub- jected to abrasion, Photomicrograph: Stratied squamous epthelim fining ofthe ssophagus (173) i Ghaplerd Tissue: The Living Fabric 108 ( Stratified cuboidal epithelium Description: Generally two layers of cube: tke cals, Location: Largest ducts of sweat glands, mammary glands, y ‘and salivary glands. Function: Protection, Photomicrograph: Stratified cuboidal epithelium forming a salivary gland duct (400°). 110 cit One Organization of the Body Figure 4.2 (continued) Stratified epithelia (g and h).. (6) Stratified columnar epithelium Description: Several cll lay: core Basal cols usualy ‘cuboidal, superficial elongated and columnar. Location: Rare in the body: small amounts in male Urethra and in large ducts ‘of some glands, Function: Protection; secretion, Photomicrograph: Stratiied columnar epithelium lining of the male urethra (360 x). Basement {membrane (n) Teansitional epithelium eee Sea ee aeeeian ae Location: Lines the ureters, blader, and part of the urethra Function: Stretches readily and permits dis- tension of urinary organ by contained urine Photomicrograph: Tanstlonal epithelium ining of the bladder, tolaxed state (170) note the Bubous, of ‘Dunded, appearance of he col at te surface, these els laten and become slongated when the bladder ed wih urine, a Ta et Bet cohen! tissue ‘examples of unicellular exocrine Glands.” (a) Photomicrograph of simple cournnar (testing) cP thetum with goblet cols (approx. 200%). 2) Diagram ofthe foe ‘Sructure of gobet cel. Notice the abundant ough endoplas is rotclum (ER) responsible for synthesizing the proten mucin, and the large Golgi apparatus, which s involved in packaging the muon seeretionin vesicles. Stratified Cuboidal Epithelium. Generally formed of only two cell layers, stratified cuboidal epithelium has a very limited distribution in ‘the body siento 4.20; tis found primarily in the ducts of sweat glands and other large glands. Chapters Tissue: The Living Fabric AA Stratified Columnar Epithelium. True stratified columnar epithelium is rare. Its specific locations are listed in Figure 4.2g. Its free surface cells are colum- nar; those in deeper layers are small and vary in shape. Transitional Epithelium. ‘Transitional epithelium forms the lining of urinary organs, which are sub- jected to considerable stretching and varying internal pressure as they fill with urine (Figure 4.2h). Cells of its basal layer are cuboidal or columnar, The apical cells vary in appearance, depending on the degree of distension of the organ. When the organ is not stretched, the membrane is many-layered and the superficial cells are rounded and domelike. When the organ is distended with urine, the epithelium thins (undergoes a transition) from about six cell layers to three, and its apical cells flatten and become squa- mouslike. The ability of transitional cells to slide past one another and change their shape accommodates the flow of a greater volume of urine through a tube- like organ; in the bladder, it allows more urine to be stored. Glandular Epithelia Cae ee ‘A gland consists of one or more cells that make and secrete a particular product. This product, called a ‘secretion, is an aqueous (water-based) fluid, typically containing proteins. Secretion is an active process whereby glandular cells obtain needed substances from the blood and transform them chemically into the cells’ secretory product, which is then discharged. Notice that the term secretion can refer to both the gland’s product and the process of making and re~ Teasing that product. Glands are classified as endocrine (en'do-krin) ot exocrine (ek’so-krin), depending on their route of, secretion, and as unicellular (“one-celled”) or mult- cellular (‘many-celled”) on the basis of their struc- ture, During embryonic development, most multicel- ular epithelial glands form by invagination from an. epithelial sheet and, at least initially, have ducts con- necting them to the epithelial sheet Endocrine Glands Because endocrine glands eventually lose their ducts, they are often called ductless glands. They produce regulatory chemicals called hormones, which they secrete directly into the extracellular space, The hor- mones then enter the blood or lymphatic fluid, Since not all endocrine glands are epithelial derivatives, we defer consideration of their structure and function to Chapter 17. 112 Unit One Organization of the Body (9) Simple tubular Example. intestinal ‘lands a simple duct structure (auct does ‘not branch) . gee Figure 44 Typesof multicellular exocrine glands. | Muttokar ands ore coset! {Gimple or compound) and the structix of ther secretory Units (Ubu according to duct type lar, alveolar, or tubuoalvecka). Exocrine Glands Exocrine glands are far more numerous than endo- crine glands, and many of their products are familiar ‘ones, The multicellular glands secrete their products ‘a duct onto body surfaces or into body cavi- ties, Exocrine glands are a diverse lot. They include sweat and oil glands, salivary glands, the liver (which secretes bile), the pancreas (which synthesizes diges- through tive enzymes), mammary (milk) glands, mucous glands, and many others. Unicellular Exocrine Glands. Unicellular exocrine glands are single cells interposed in an epithelium between cells with other functions. They have 22 ducts. In humans, all such glands produce mucit (mu’sin), a complex glycoprotein that dissolves (a) Merocrine gland Figure 4.5 Modes of secretion in exocrine glands. (a) Merocrine glands secrete their products by exooytosis. (b) In holocrine glands, the entire secretory cel ruptures, releasing secretions and dead cell fragments. (e) In apocrine glands, the apex of each secretory cal pinches off and releases its secretions. water. Once dissolved, mucin forms mucus, a slimy coating that both protects and lubricates surfaces. The only important unicellular glands in humans are the goblet cells found scattered among the columnar epi- thelial cells lining the intestinal and respiratory tracts (Figure 4.3). Although unicellular glands probably outnumber multicellular glands, unicellular glands are the less well known of the two gland types. Multicellular Exocrine Glands. Multicellular exo- crine glands have two common structural elements: an epithelium-derived duct and a secretory unit con- sisting of secretory cells. Furthermore, in all but the simplest glands, supportive connective tissue sur. founds the secretory unit and supplies it with blood vessels and nerve fibers. Often, the cunnective tissue forms a fibrous capsule that extends into the gland proper and divides the gland into lobes. P Multicellular glands are divided into two major categories on the basis of their duct structures. Simple glands have a single unbranched duct, whereas com- pound glands have a branching duct. The glands can be further described according to ae art sre secretory parts as (1) tubular, with the secretory cells. fone tube; (2) alveolar (al-ve'o-lar), with the secretory cells forming small, flasklike sacs (a/veo/us = small hollow cavity); and (3) tubuloalveolar, con- taining both tubular au alyeole: ae use, ‘Terms denoting duct and secretory part structure combined to “describe the gland fully (Figure 4.4). It should be noted that the term acinar (as‘i-nar; “ber- rylike”) is used interchangeably with alveolar. ‘Since multicellular glands secrete their products in different ways, they can also be classified function- ally, according to their secretory behavior. Most exo- crine glands are merocrine (mer‘o-krin) glands, which secrete their products by exocytosis shortly after the products are produced. The secretory cells are not altered in any way. The pancreas, most sweat glands, and salivary glands belong to this class (Figure 45a). Secretory cells of holocrine (hol’o-krin) glands accumulate their products within them until they rupture, (They are replaced by the division of under- ying cells.) Since holocrine gland secretions include the synthesized product plus dead cell fragments (holas = all), you could say that their cells “die for their cause.” Sebaceous (oil) glands of the skin are the only true example of holocrine glands (Figure 4.5b). Apocrine (ap’o-krin) glands also accumulate their products, but in this case accumulation occurs only just beneath the free surface. Eventually, the apex of the cell pinches off (apo = from, off) and the secretion is released. The cell repairs its damage and repeats the process again and again, (Figure 4.5¢). ‘There is some controversy over whether humans have apocrine glands, but apocrine glands are definitely present in other animals. The only possibilities in humans are the mammary glands but, since eytoplas- mic loss is slight, many histologists classify mammary glands as merocrine glands. 14 UuitOne Organization ofthe Body ie Soe Fe SN. s | Ferobiast chondisbast_ oslo _ Bn . =o os 1 | pees moons Chontoarte See cee | | pee ) LJ | | soll Class of J connective Se Jee igs as } ‘om resulting: Coomectiyp tative: Cartilage ‘Osseous (bone) Blood Du | ee | eldeatassy (pene ive 1. compact *eood cel latin one vonige Sone and dtorariaton we le oobi | Tos oar armocnese Stag atsme md soe Crethou oe Reticular 3. Elastic bone Chapter 18. j Era | 2 Dente rac | tae Tyoee: Rua eae | oes | Figure 4,6 Major classes of connective tissue. ‘common embryonic tissue type (mesenchyme) lo So Connective Tissue me Connective tissue is found everywhere in the body. It is the most abundant and widely distributed of the primary tissues, but its amount in particular organs Varies greatly. For example, skin consists primarily of connective tissue, while the brain contains very little. ‘Connective tissue does much more than connect. body parts; it has many forms and many functions. Its chief subclasses are connective tissue proper, carti- lage, bone, and blood. Its major functions include (1) binding and support, (2) protection, (3) insulation, and, as blood, (4) éransporfation of substances within the body. For example, cordlike connective tissue structures connect muscle to bone (tendons), and fine, resilient connective tissue invades soft organs and supports and binds their cells together. Bone and car- tilage support and protect body organs by providing hard “underpinnings”; fat cushions insulate and pro- tect body organs as well as providing reserve energy fuel. Al these classes arise from the same Common Characteristics of Connective Tissue | Despite their multiple and varied functions in the body, connective tissues have certain propetis i common that set them apart from other primary tissues: 1, Common origin. All connective tissues aise fom mesenchyme, an embryonic tissue derived ‘rom the mesoderm germ layer, and hence have a common kit: ship (Figure 4.6), The embryonic germ layers *° described later in this chapter (p. 132). 2, Degrees of vascularity. Unlike epithelium, iis avascular, and muscle and nervous tissue, which bi? a rich vascular supply, connective tissues UP the entire gamut of vascularity. Cartilage is avast dense connective tissue is poorly vascularize the other types have a rich supply of blood vess"* 3. Extracellular matrix. Whereas all thor primes} sues are composed mainly of cells, connective tr are composed largely of nonliving extracell™ Snel ix, which separates, often widel 4 ates, often widely, the living cells of the tissue. Because of this matrix, connective tis is able to bear weight, withstand gre: endure abuses, such rat tension, physical trauma and abra: that no other tissue could withstand. Structural Elements of Connective Tissue In any typo of connective tissue, three elements must be considered: ground substance, fibers, and cells. The ground substance and fibers make up the extra: cellular matrix, (Note that some authors use the term matrix to indicate the ground substance only.) The properties of the cells and the composition and arrangement of extracellular matrix elements vary tre- mendously, giving rise to an amazing diversity of con- noctive tissues, each uniquely adapted to perform its specific function in the body. For example, the matrix can be delicate and fragile to form a soft “packing” around an organ, or it can form “ropes” (tendons and ligaments) of incredible strength. Ground Substance Ground substance is an amorphous (unstructured) material that fills the space between the cells and con- tains the fibers. Itis composed of interstitial fluid, and proteoglycans (pro'te-o-gli’kanz). The proteoglycans consist of a protein “core” to which g/rcosaminogly- ‘cans (gli"kos-ah-me"no-gli’kanz) (GAGs) are attached. ‘The GAGs, a diverse group of large, negatively charged polysaccharides, stick out from the core protein like the fibers of a bottle brush. The strandlike GAGs coil, intertwine, and trap water, forming a substance that varies from a fluid to a somistiff hydrated gel. One type of GAG, hyaluronic (hi’ab-lu-ron‘ik) acid, is found in virtually all connective tissues, and its rela- tive amount helps determine the viscosity and per- meability of the ground substance. ‘The ground substance functions as a molecular sgieve"” or medium, through which nutrionts.and. other dissolved substances can diffuse between the blood capillaries and the cells. The fibers in the matrix impede diffusion somewhat and make the ground substance less pliable. Fibers TThrwe types of fibers are found in the matrix of con- nective tissue: collagen, elestic, and reticular fibers. Of these, collagen fibers aro by far the most abundant, Collagen fibers are constructed primarily of the fibrous protein collagen. Collagen molecules, ore secreted Into the extracellular space, where they assemble spontaneously into fibers. Collagen fibers Qhaptert) Tissue: The Living Fabric 118 are extremely tough and provide high tensile strength (that is, the ability to resist longitudinal stress) to the matrix. Indeod, stress testing has shown collagen fibers to be stronger than steel fibers of the same size! When fresh, they have a glistening white appearance; they are therefore also called white fibers. Elastic fibers are formed largely from another fibrous protein, e/astin, Elastin has a randomly coiled structure that allows it to stretch and recoil like a rubber band. The presence of elastin in the matrix gives it a rubbery, orresilient, quality. Collagen fibers, always found in the same tissuo, stretch a bit and then lock” in full extension, which limits the extent of stretch and prevents the tissue from tearing, Elastic fibers then snap the connective tissue back to its nor- ‘mal length when the tension lets up, Elastic fibers are found where greater elasticity is needed, for example, in the skin, lungs, and blood vessel walls. Since fresh clastic fibers appear yellow, they are sometimes called yellow fibers. Reticular fibers are fine collagenous fibers (with a slightly different chemistry and form) and are con- tinuous with collagen fibers. They branch exten- sively, forming dolicate networks (reticu/ = network) that surround small blood vessels and support the soft tissue of organs, They are particularly abundant at junctions between connective tissue and other tissue types, for example, in the basement membrane of epi- thelial tissues Cells Each major class of connective tissue has a fundamen- tal coll type that exists in immature and mature forms (see Figure 4.6). The undifferentiated cells, indicated by the suffix B/ast (literally, “bud,” or “sprout,” but meaning “forming”), are actively mitotic cells that secrete both the ground substance and the fibers char- acteristic of their particular matrix. The primary blast cell types by connective tissue class are (1) connective tissue proper: fibroblast; (2) cartilage: chondroblast (Kon’dro-blast); (3) bone: osteoblast (os'te-o-blast’); and (4) blood: hemocytoblast (he’mo-si’to-blast) or hematopoietic stem cell. Once the matrix has been synthesized, the blast colls assume their less active, mature mode, indicated by the suffix cyte (see Figure 4.6). The mature cells are responsible for maintaining the matrix in a healthy state. However, if the matrix is injured, the mature cells can easily revert to theirmore active state to make repairs and regenerate the matrix. (Note that the hemocytoblast, the stem cell of bone marrow, always remains actively mitotic.) ‘Additionally, connective tissue proper, espe- cially the loose connective tissue type called areolar, is “home” to an assortment of other cell types, such as nutrient-storing fat cells and mobile cells that migrate into the connective tissue matrix from the blood- 116 Unit One Organization of the Body stream, The latter include white blood cells (neutro- phils, eosinophils, lymphocytes) and other cell types concerned with tissue response to injury, such as mast cells, macrophages (mak'ro-fajz), and antibody-pro- ducing plasma cells Although all of these accessory cell types are described in lator chapters, the mast cells and macro- phages aro so significant to overall body defense that they deserve a brief mention here, The oval mast cells aro typically found clustered in tissue spaces deep to an epithelium or along blood vessels. These cells act as ‘‘sensitive sontinels” to detect foreign substances (og., bacteria, fungi) and initiate local inflammatory responses against them. In the mast cell cytoplasm are conspicuous secretory granules containing (1) Aepa- vin (hepah-rin) and (2) Aistamine (his'tah-mén). Although heparin is known to be an anticoagulant (a chemical that helps prevent blood clotting) when free in the bloodstream, its significance in human mast colls is still uncertain, Histamine, which is released during inflammatory reactions, makes the capillaries leaky. (The inflammatory process is discussed in Chapter 22.) Macrophages (:2acro ~ large: phago = eat) are largo, irregularly shaped cells that avidly phagocytize both foreign matter that has managed to invade the body and dying or dead tissue cells. They are also cen- tral actors in the immune system, In connective tis- sues, they may bo fixed (attached to the connective tissue fibers) or they may migrate freely through the matrix, However, macrophages are not limited to con~ nective tissue, In fact, their body distribution is so broad and their numbers so vast that they are often referred to collectively as the macrophage system. ‘Macrophages are peppered throughout loose con- nective tissue, bone marrow, lymphatic tissue, the spleen, and the mesentery that suspends the abdomi- nal viscera. Those in certain sites are given specific names; they are called Aistiocytes (his'te-o-sitz’) in loose connective tissue, Kupffer (KO6p' fer) ced/sin the liver. and microglial (mi-ktog’le-ul) ced/s inthe brain. Although all these cells are phagocytes, some have selective appetites. For example, the macrophages of the spleen function primarily to engulf aging red blood cells; but they will not tum down other “deli- cacies” that come their way. Types of Connective Tissue ‘As noted, all classes of connective tissue consist of living cells surrounded by a matrix, Their major dif- ferences reflect cell type, fiber type, and proportion of the matrix contributed by fibers. Collectively, these throe factors determine not only major connective tis- but also their subclasses and types, The sue classes described in this section are sue classe connectiv {llustrated in Figure 4.7. Since the matu tissues arise from a common embryonic tissue, appropriate to describe this hore as well, Embryonic Connective Tissue: Mesenchyme Mesenchyme (mes‘eng-kim), or mesenchymal tissue, is the first definitive tissue formed from the meso. derm gorm layer. It arises during the early weeks of embryonic development and eventually differentiates (specializes) into all other connective tissues. Mesen- ‘chyme is composed of star-shaped mesonchymal cells and a fluid ground substance containing fine fibrils (Figure 4.7a). 7 Mucous connective tissue is a temporary tissue, derived from mesenchyme and similar to it, that appears in the fetus in very limited amounts. Whar- ton’ jelly, which supports the umbilical cord, is the best representative of this scant embryonic tissue. Connective Tissue Proper Connective tissue proper has two subclasses: the loose connective tissues (areolar, adipose, and reticu- lar) and dense connective tissues (dense regular, dense irregular, and elastic). Except for bone, carti- lage, and blood, all mature connective tissues belong to this class. Areolar Connective Tissue. Areolar (ah-re’o-lar) connective tissue has a semifluid ground substance formed primarily of hyaluronicacid in which all three fiber types are loosely dispersed (Figure 4.7b). Areolar connective tissue can be viewed as the prototype, ot model, for the connective tissues proper, because in terms of types (but not actual number) of matrix ele~ ‘ments, all other subclasses are simy sof this common tissue type. <0 Se il Fibroblasts, flat, branchi dle shaped in profile, are the predominant cell type of thie tissue, Numerous macrophages are also seen and present a formidable barrier to invading microorgan- isms, but other cell types are scattered throughout. Fat colls appear singly orin small clusters, and occasional mast cells are identified easily by the large, darkly stained cytoplasmic granules that often obscure their nuclei Perhaps the most obvious structural feature of this tissue is the loose arrangement of its supportive fibers, which account for only small portions of trix. The rest of the matrix, occupied by fluid ground substance, appears to be empty space when viewed through the microscope; in fact, the Latin term areola means ‘‘a small open space.” Essentially all body cells obtain their nutrients from and release their wastes into this “tissue fluid,” Because of its Figure 4.7 Connectivettissues, Embryonic connective tissue (@) Mesenchyme Deserpion: Embyonic somectve sue ike ground substance containing fine toes; areneped mesenchyiel cota. i Location: Primariy in embryo, Function: Gives rise to all other connective tissue types, tissue, an tissue (475 x); the clear-appearing ‘substance of the, tibers. Ghopter4 Teste: The Living Fabric WT wd ‘Connective tissue proper: Loose connective tissue (b to d) (©) Areolar connective tissue Location: Widely distributed under epithelia of body, eg, forms lamina propria of =~ mucous membranes; packages ‘organs surrounds capibaies, Immportant role in inflammation, and Areca a soft packaging tissue ofthe body (170%). 118 nit One Organization of the Body Figure 4.7. (continued) i tt (©) Adipose tissue Description: Mains asin areolar, but very sparse, closely packed adipocytes, of lat cells, have nucleus pushed to tne side by large Tat droplet. Location: Under skin; around kidneys and eyeballs; in bones and within abdomen: in breasts Funetion: Provides reserve food fuel insulates Function: Fibers form a soft interna skeleton ‘against heat loss: supports and protects organs ‘hat suppons other cel ypes. Photomicrograph: Adpose tissue fom the sub- Photomicrograph: Dark staining network of Claneous aperunder te sr (300%) Tesla eonracive tase bre torming thenterrastleton ofthe spin (625%). Figure 4.7. (continued) Connective tissue proper: Dense connective tissue (¢ og) (©) Dense regular connective tissue Description: Pri ‘afew olastin tbe fibroblast. ly parallel collagen fibers; ‘major cal type isthe Location: Tendons, most ligaments, aponeuroses. Shoulder joint Ligament Tendon "Photomicrograph: Dens reuar connective tasve ~ trom a tendon (200%). Chapters ssue: The Living Fabric 119 (1) Dense irregular connective tissue Description: Primarly irregularly arranged ‘colagen fibers; some elastic fers; major Coll type is the fibrobast. Location: Dermis of the skin; submucosa of — ‘digestive tract fibrous ‘capauies of organs and of jpints. Function: Able to withstand tension exerted in many directions; provides structural strength. Photomic \: Dense irregular connective tissue from ‘of the skin (475%). ———<=

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