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OpenStax-CNX module: m47742 1

Histology Module 4A and 4B:

Connective Tissue*

Donna Browne

Based on Connective Tissue Supports and Protects� by

OpenStax

This work is produced by OpenStax-CNX and licensed under the

Creative Commons Attribution License 3.0�

Abstract

By the end of this section, you will be able to:

• Identify and distinguish between the types of connective tissue: proper, supportive, and �uid• Explain the functions of connective tissues

As may be obvious from its name, one of the major functions of connective tissue is to connect tissues andorgans. Unlike epithelial tissue, which is composed of cells closely packed with little or no space in between,connective tissue cells are spread out in a matrix. The matrix plays a major role in the functioning of thistissue. The major component of the matrix is protein �bers. The rest of the matrix is usually a �uid, but itcan also be mineralized and solid, as in bones. Connective tissues come in a vast variety of forms, yet theytypically have in common three characteristic components: cells, matrix and protein �bers. The amountand structure of each component correlates with the function of the tissue, from the rigid ground substancein bones supporting the body to the inclusion of specialized cells; for example, a cell that destroy moleculesthat can make you sick.

1 Functions of Connective Tissues

Connective tissues perform many functions in the body, but most importantly, they support and connectother tissues; from the connective tissue covering that surrounds muscle cells, to the tendons that attachmuscles to bones, and to the skeleton that supports the positions of the body. Protection is another majorfunction of connective tissue, in the form of capsules and bones that protect delicate organs and, of course,the skeletal system. Specialized cells in connective tissue defend the body from microorganisms that enter thebody. Transport of �uid, nutrients, waste, and hormones is ensured by specialized �uid connective tissues,such as blood and lymph. Adipose cells store surplus energy in the form of fat and contribute to the thermalinsulation of the body.

The Functions of Connective Tissue

*Version 1.2: Feb 25, 2014 10:10 am -0600�http://cnx.org/content/m46049/1.4/�http://creativecommons.org/licenses/by/3.0/

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• . Encloses and separates. Connective tissue covers each muscle separating it from the other musclesaround it.

• Connects tissues. Straps of connective tissue connect muscle to bone• Supports and moves. Bone and cartilage add strength and support to the nose and joints.• Stores. Adipose stores fat. Bone stores minerals.• Cushions and insulates. Adipose tissue cushions your kidneys and eyeballs, it also serves as insulate

your body against heat loss.• Transports. Blood transports many substance throughout the entire body.• Protects. Bone protects the brain and heart.

2 Classi�cation of Connective Tissues

The three broad categories of connective tissue are classi�ed according to the characteristics of their matrixand the types of �bers found within the matrix (Table 1). Connective tissue proper includes loose orareolar connective tissue and dense connective tissue. Both tissues have a variety of cell types andprotein �bers suspended in a thick liquid matrix. Dense connective tissue is reinforced by bundles of �bersthat provide strength, elasticity, and protection. In loose connective tissue, the �bers are loosely organized,leaving large spaces in between. Supportive connective tissue�bone and cartilage�provide structureand strength to the body and protect soft tissues. A few distinct cell types and densely packed �bers ina matrix characterize these tissues. In bone, the matrix is rigid and described as calci�ed because of thedeposited calcium salts. In �uid connective tissue, in other words, lymph and blood, various specializedcells circulate in a watery �uid containing salts, nutrients, and dissolved proteins.

Connective Tissue Examples

Connective tissue proper Supportive connective tissue Fluid connective tissue

Loose connective tissue

• Areolar• Adipose• Reticular

Cartilage

• Hyaline• Fibrocartilage• Elastic

Blood

Dense connective tissue

• Regular elastic• Irregular elastic

Bones

• Compact bone• Cancellous bone

Lymph

Table 1

3 Connective Tissue Proper

Fibroblasts are present in all connective tissue proper (Figure 1 (Connective Tissue Proper )). Fibroblastsand adipocytes are �xed cells, which means they remain within the connective tissue. Other cells, such asmast cells move in and out of the connective tissue in response to chemical signals.

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Connective Tissue Proper

Figure 1: Fibroblasts produce this �brous tissue. Connective tissue proper includes the �xed cells�brocytes, adipocytes, and mesenchymal cells. LM × 400. (Micrograph provided by the Regents ofUniversity of Michigan Medical School©2012)

3.1 Cell Types

The most abundant cell in connective tissue proper is the �broblast.Proteoglycans and proteins secretedby �broblasts combine with extra-cellular �uids to produce a viscous ground substance that, with embedded�brous proteins, forms the extra-cellular matrix. As you might expect, a �brocyte, a less active form of�broblast, is the second most common cell type in connective tissue proper.

3.2 Connective Tissue Fibers and Ground Substance

Three main types of �bers are secreted by �broblasts: collagen �bers, elastic �bers, and reticular �bers.Collagen �bers, while �exible, have great tensile strength, resist stretching, and give ligaments and tendonstheir characteristic resilience and strength. These �bers hold connective tissues together, even during themovement of the body.

Elastic �ber contains the protein elastin. The main property of elastic �bers are that after beingstretched or compressed, it will return to its original shape. Elastic �bers are prominent in elastic tissuesfound in skin and the elastic ligaments of the vertebral column and the outer ear.

Reticular �ber is also formed from the same protein subunits as collagen �bers; however, these �bersform a branching network. They are found throughout the body, but are most abundant in the reticulartissue of soft organs, such as liver and spleen, where they anchor and provide structural support

3.3 Loose Connective Tissue

Loose connective tissue is found between many organs where it acts both to absorb shock and bind tissuestogether. It allows water, salts, and various nutrients to di�use through to adjacent or imbedded cells andtissues.

Adipose tissue consists mostly of fat storage cells, with little extracellular matrix (Figure 2 (AdiposeTissue )). White fat contributes mostly to lipid storage and can serve as insulation from cold temperaturesand mechanical injuries. Adipose tissue can be found protecting the kidneys and cushioning the back of theeye. It is also found directly beneath the skin.

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Adipose Tissue

Figure 2: This is a loose connective tissue that consists of fat cells with little extracellular matrix.It stores fat for energy and provides insulation. LM × 800. (Micrograph provided by the Regents ofUniversity of Michigan Medical School©2012)

Areolar tissue shows little specialization. It contains all the cell types and �bers previously describedand is distributed in a random, web-like fashion. It �lls the spaces between muscle �bers, surrounds bloodand lymph vessels, and supports organs in the abdominal cavity. Areolar tissue underlies most epithelia andrepresents the connective tissue component of epithelial membranes, which are described further in a latersection.

Reticular tissue is a mesh-like, supportive framework for soft organs such as lymphatic tissue, thespleen, and the liver (Figure 3 (Reticular Tissue )). Reticular cells produce the reticular �bers that form thenetwork onto which other cells attach. It derives its name from the Latin reticulus, which means �little net.�

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Reticular Tissue

Figure 3: This is a loose connective tissue made up of a network of reticular �bers that provides asupportive framework for soft organs. LM × 1600. (Micrograph provided by the Regents of Universityof Michigan Medical School ©2012)

3.4 Module 4B begins here ##########################################################Dense

Connective Tissue

Dense connective tissue contains more collagen �bers than does loose connective tissue. As a consequence, itdisplays greater resistance to stretching. There are two major categories of dense connective tissue: regularand irregular. Dense regular connective tissue �bers are parallel to each other, enhancing tensile strength andresistance to stretching in the direction of the �ber orientations. Ligaments and tendons are made of denseregular connective tissue, but in ligaments not all �bers are parallel. Dense regular elastic tissue containselastin �bers in addition to collagen �bers, which allows the ligament to return to its original length afterstretching. The ligaments in the vocal folds and between the vertebrae in the vertebral column are elastic.

In dense irregular connective tissue, the direction of �bers is random. This arrangement gives the tissuegreater strength in all directions and less strength in one particular direction. In some tissues, �bers crisscrossand form a mesh. In other tissues, stretching in several directions is achieved by alternating layers where�bers run in the same orientation in each layer, and it is the layers themselves that are stacked at anangle. The dermis of the skin is an example of dense irregular connective tissue rich in collagen �bers. Denseirregular elastic tissues give arterial walls the strength and the ability to regain original shape after stretching(Figure 4 (Dense Connective Tissue )).

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Dense Connective Tissue

Figure 4: (a) Dense regular connective tissue consists of collagenous �bers packed into parallel bundles.(b) Dense irregular connective tissue consists of collagenous �bers interwoven into a mesh-like network.From top, LM × 1000, LM × 200. (Micrographs provided by the Regents of University of MichiganMedical School©2012)

: Connective Tissue: Tendinitis

Your opponent stands ready as you prepare to hit the serve, but you are con�dent that you willsmash the ball past your opponent. As you toss the ball high in the air, a burning pain shootsacross your wrist and you drop the tennis racket. That dull ache in the wrist that you ignoredthrough the summer is now an unbearable pain. The game is over for now.

After examining your swollen wrist, the doctor in the emergency room announces that you havedeveloped wrist tendinitis. She recommends icing the tender area, taking non-steroidal anti-in�ammatory medication to ease the pain and to reduce swelling, and complete rest for a fewweeks. She interrupts your protests that you cannot stop playing. She issues a stern warning aboutthe risk of aggravating the condition and the possibility of surgery. She consoles you by mentioningthat well known tennis players such as Venus and Serena Williams and Rafael Nadal have alsosu�ered from tendinitis related injuries.

What is tendinitis and how did it happen? Tendinitis is the in�ammation of a tendon, the thick

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band of �brous connective tissue that attaches a muscle to a bone. The condition causes painand tenderness in the area around a joint. On rare occasions, a sudden serious injury will causetendinitis. Most often, the condition results from repetitive motions over time that strain thetendons needed to perform the tasks.

Persons whose jobs and hobbies involve performing the same movements over and over again areoften at the greatest risk of tendinitis. You hear of tennis and golfer's elbow, jumper's knee, andswimmer's shoulder. In all cases, overuse of the joint causes a microtrauma that initiates thein�ammatory response. Tendinitis is routinely diagnosed through a clinical examination. In caseof severe pain, X-rays can be examined to rule out the possibility of a bone injury. Severe cases oftendinitis can even tear loose a tendon. Surgical repair of a tendon is painful. Connective tissue inthe tendon does not have abundant blood supply and heals slowly.

:

Watch this animation1 to learn more about tendonitis, a painful condition caused by swollen or

1http://openstaxcollege.org/l/tendonitis

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injured tendons.

4 Supportive Connective Tissues

Two major forms of supportive connective tissue, cartilage and bone, allow the body to maintain its postureand protect internal organs.

4.1 Cartilage

The distinctive appearance of cartilage is due to the semisolid matrix. Embedded within the cartilage matrixare chondrocytes, or cartilage cells, and the space they occupy are called lacunae (singular = lacuna).Cartilaginous tissue is avascular (without blood), thus all nutrients need to di�use through the matrix toreach the chondrocytes. This is a factor contributing to the very slow healing of cartilaginous tissues.

The three main types of cartilage tissue are hyaline cartilage, �brocartilage, and elastic cartilage (Figure 5(Types of Cartilage )). Hyaline cartilage, the most common type of cartilage in the body, consists of shortand dispersed collagen �bers and contains large amounts of proteoglycans. Under the microscope, tissuesamples appear clear. The surface of hyaline cartilage is smooth. Both strong and �exible, it is found in therib cage and nose and covers bones where they meet to form moveable joints. It makes up the embryonicskeleton before bone formation. A plate of hyaline cartilage at the ends of bone allows continued growthuntil adulthood. Fibrocartilage is tough because it has thick bundles of collagen �bers dispersed throughits matrix. The knee and jaw joints and the the intervertebral discs are examples of �brocartilage. Elasticcartilage contains elastic �bers as well as collagen and proteoglycans. This tissue gives rigid support aswell as elasticity. Tug gently at your ear lobes, and notice that the lobes return to their initial shape. Theexternal ear contains elastic cartilage.

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Types of Cartilage

Figure 5: Cartilage is a connective tissue consisting of collagenous �bers embedded in a �rm matrixof chondroitin sulfates. (a) Hyaline cartilage provides support with some �exibility. The example isfrom dog tissue. (b) Fibrocartilage provides some compressibility and can absorb pressure. (c) Elasticcartilage provides �rm but elastic support. From top, LM × 300, LM × 1200, LM × 1016. (Micrographsprovided by the Regents of University of Michigan Medical School ©2012)

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4.2 Bone

Bone is the hardest connective tissue. It provides protection to internal organs and supports the body. Bone'srigid extracellular matrix contains mostly collagen �bers embedded in a mineralized matrix containing aform of calcium phosphate. Both components of the matrix, �bers and minerals, contribute to the unusualproperties of bone. Without collagen, bones would be brittle and shatter easily. Without mineral crystals,bones would �ex and provide little support. Osteocytes, bone cells like chondrocytes, are located withinlacunae. Bone is a highly vascularized tissue. Unlike cartilage, bone tissue can recover from injuries in arelatively short time.

5 Fluid Connective Tissue

Blood and lymph are �uid connective tissues. Cells circulate in a liquid extracellular matrix. The formedelements circulating in blood are all derived from stem cells located in bone marrow (Figure 6 (Blood: AFluid Connective Tissue)). Erythrocytes, red blood cells, transport oxygen and some carbon dioxide.Leukocytes, white blood cells, are responsible for defending against potentially harmful microorganisms ormolecules. Platelets are cell fragments involved in blood clotting. Nutrients, salts, and wastes are dissolvedin the liquid matrix and transported through the body.

Lymph contains a liquid matrix and white blood cells. Lymphatic capillaries are extremely permeable,allowing larger molecules and excess �uid from interstitial spaces to enter the lymphatic vessels. Lymphdrains into blood vessels, delivering molecules to the blood that could not otherwise directly enter thebloodstream. In this way, specialized lymphatic capillaries transport absorbed fats away from the intestineand deliver these molecules to the blood.

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Blood: A Fluid Connective Tissue

Figure 6: Blood is a �uid connective tissue containing erythrocytes and various types of leukocytesthat circulate in a liquid extracellular matrix. LM × 1600. (Micrograph provided by the Regents ofUniversity of Michigan Medical School©2012)

6 Chapter Review

Connective tissue is a heterogeneous tissue with many cell shapes and tissue architecture. Structurally, allconnective tissues contain cells that are embedded in an extracellular matrix stabilized by proteins. Thechemical nature and physical layout of the extracellular matrix and proteins vary enormously among tissues,re�ecting the variety of functions that connective tissue ful�lls in the body. Connective tissues separateand cushion organs, protecting them from shifting or traumatic injury. Connect tissues provide supportand assist movement, store and transport energy molecules, protect against infections, and contribute totemperature homeostasis.

Many di�erent cells contribute to the formation of connective tissues. Fibroblasts are the most abundantand secrete many protein �bers, adipocytes specialize in fat storage, stem cells from the bone marrow giverise to all the blood cells, chondrocytes form cartilage, and osteocytes form bone. The extracellular matrixcontains �uid, proteins, polysaccharide derivatives, and, in the case of bone, mineral crystals. Protein �bersfall into three major groups: collagen �bers that are thick, strong, �exible, and resist stretch; reticular �bersthat are thin and form a supportive mesh; and elastin �bers that are thin and elastic.

The major types of connective tissue are connective tissue proper, supportive tissue, and �uid tissue.Loose connective tissue proper includes adipose tissue, areolar tissue, and reticular tissue. These serve tohold organs and other tissues in place and, in the case of adipose tissue, isolate and store energy reserves. Thematrix is the most abundant feature for loose tissue although adipose tissue does not have much extracellularmatrix. Dense connective tissue proper is richer in �bers and may be regular, with �bers oriented in parallel as

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in ligaments and tendons, or irregular, with �bers oriented in several directions. Organ capsules (collagenoustype) and walls of arteries (elastic type) contain dense irregular connective tissue. Cartilage and bone aresupportive tissue. Cartilage contains chondrocytes and is somewhat �exible. Hyaline cartilage is smooth andclear, covers joints, and is found in the growing portion of bones. Fibrocartilage is tough because of extracollagen �bers and forms, among other things, the intervertebral discs. Elastic cartilage can stretch andrecoil to its original shape because of its high content of elastic �bers. The matrix contains very few bloodvessels. Bones are made of a rigid, mineralized matrix containing calcium salts, crystals, and osteocyteslodged in lacunae. Bone tissue is highly vascularized. Cancellous bone is spongy and less solid than compactbone. Fluid tissue, for example blood and lymph, is characterized by a liquid matrix and no supporting�bers.

Glossary

De�nition 6: adipocytes

lipid storage cells

De�nition 6: adipose tissue

specialized areolar tissue rich in stored fat

De�nition 6: areolar tissue

(also, loose connective tissue) a type of connective tissue proper that shows little specialization withcells dispersed in the matrix

De�nition 6: chondrocytes

cells of the cartilage

De�nition 6: collagen �ber

�exible �brous proteins that give connective tissue tensile strength

De�nition 6: connective tissue proper

connective tissue containing a viscous matrix, �bers, and cells.

De�nition 6: dense connective tissue

connective tissue proper that contains many �bers that provide both elasticity and protection

De�nition 6: elastic cartilage

type of cartilage, with elastin as the major protein, characterized by rigid support as well as elasticity

De�nition 6: elastic �ber

�brous protein within connective tissue that contains a high percentage of the protein elastin thatallows the �bers to stretch and return to original size

De�nition 6: �broblast

most abundant cell type in connective tissue, secretes protein �bers and matrix into the extracellularspace

De�nition 6: �brocartilage

tough form of cartilage, made of thick bundles of collagen �bers embedded in chondroitin sulfateground substance

De�nition 6: �brocyte

less active form of �broblast

De�nition 6: �uid connective tissue

specialized cells that circulate in a watery �uid containing salts, nutrients, and dissolved proteins

De�nition 6: ground substance

�uid or semi-�uid portion of the matrix

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De�nition 6: hyaline cartilage

most common type of cartilage, smooth and made of short collagen �bers embedded in a chondroitinsulfate ground substance

De�nition 6: lacunae

(singular = lacuna) small spaces in bone or cartilage tissue that cells occupy

De�nition 6: loose connective tissue

(also, areolar tissue) type of connective tissue proper that shows little specialization with cellsdispersed in the matrix

De�nition 6: matrix

extracellular material which is produced by the cells embedded in it, containing ground substanceand �bers

De�nition 6: mesenchymal cell

adult stem cell from which most connective tissue cells are derived

De�nition 6: mesenchyme

embryonic tissue from which connective tissue cells derive

De�nition 6: mucous connective tissue

specialized loose connective tissue present in the umbilical cord

De�nition 6: parenchyma

functional cells of a gland or organ, in contrast with the supportive or connective tissue of a glandor organ

De�nition 6: reticular �ber

�ne �brous protein, made of collagen subunits, which cross-link to form supporting �nets� withinconnective tissue

De�nition 6: reticular tissue

type of loose connective tissue that provides a supportive framework to soft organs, such as lym-phatic tissue, spleen, and the liver

De�nition 6: supportive connective tissue

type of connective tissue that provides strength to the body and protects soft tissue

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