Essentials of Human Anatomy and Physiology Nuclear enveloPe

Chromatin Nucleolus
Nuclear
Nucleus

pores Nucleus

Cytoplasm
Plasma membrane
(a) Rough ER

(b)

FEffiLSffiffis,"gAnatomyofthegeneralizedanimalcellnucleus. the generalized cell.

(a) orientation diagram: The ihree marn regions of (b) Structure of the nucleus'

large essential for life. In addition to containing constantly are cells amounts of water, all the body (something bathed in a dilute saltwater solution like seawater) called interstitial Jluid, which is decells rived from the blood. All exchanges between fluid' and blood are made through this Cells vary tremendously in length-ranging from 2 micrometers (1/12,000th of an inch) in the in smallest cells to over a meter (3 feet) or more toes' your wiggle to the nerye cells that cause you Furthermore, a cell's structure often reflects its function; this will become clear later in this chapter. Celis can have amazingly different shapes' have Some are disk-shaped (red blood cells), some like threadlike exlensions (nerve cells), others are toothpicks pointed at each end (smooth muscle
cells).

1. Hydrogen and nitrogen are

two of the four elements that make up the bulk of cell structure'

What are the other two?

K2. Why are electrocytes, particularly Na+ and ions, so imPortant to bodY function?
For answers. see APPendix
D

Anatomy of a Generalized Gell

Cells also vary dramatically in the functions' or roles, they play in the body' For example' white blood celL wander freely through the body bactetissues and protect the body by destroying cells Some ria and other foreign substances' other make hormones or chemicals that regulate body cells. Still others take patt in gas exchanges in tn" lungs or cleanse the blood (kidney tubule
cells).

Although no one cell type is exactly like- all others ar cells d.o have the same basic parts, and there u'-i certain functions common to all cells' Here we talk about tkre generalized cell, which demor strates these many typical features' In general, all cells have three main regions c purrr-"n nuclews (nu'kle-us), cytoplasm (si't< plazd'), and a plasma membrane (Figure 3'7a) Tf of the ce n -rcl".,s is usually located near the center It is surrounded by the semifluid cltoplasm, whic in turn is enclosed by the plasma membrane' whic forms the outer cell boundary' (Figure 3'4 on p'

Essentials of Human Anatomy and Physiology

Some proteins float freely in the lipid phase af the membrane whereas others are
anchored in specific locations. What could serve as anchortng structures tn the latter
case?

Extracellular fluid
(watery environment)

Glyco

Glycolipid $
Cholesterol

Sugar group
Po\ar heads o{

phospholipid
molecules Bimolecular lipid layer containing proteins Nonpolar tails of

phospholipid
molecules

Filaments of cytoskeleton

Cytoplasm (watery environmen:

P$#[,iffi

fi 3"tr

Structure of the plasma membrane.

torn. The hydrophobic makeup of the membrane interior makes the plasma membrane relatively impermeable to most water-so1uble molecules. The cholesterol heips keep the membrane fluid. The proteins scatterecl in the lipid bilayer are responsible for most of the specializecl functions of the membrane. Some proteins are enzymes. Many of the proteins protruding from the ce11 exterior are receptors for hormones or other chemical messengers or are binding sites for anchoring the cell to fibers or to otl-ier structures inside or outside the
cell. Most proteins that span the membrane are involved in transport filnctions. For example, some cluster together to form protein channels (tiny pores) through which s/ater and small watersoluble molecules or ions can move; others act as carriers that bind to a substance and move it through. the membrane. Branching sugar grollps are attached to most of the proteins abutting the extracellular space. Such "sugar-proteins" are called glycoproteizs, and because of their presence, the cell surface is a fuzzy, sticky, sugar-rich
'su tel,otd

area called the glycocal.1x (g1i-co-ka'liks). (You c., think of your cells as being sugar-coated.) Amor:-

other things, these glycoproteins cletermine )'o-.r blood type, act as receptors that certain baclelr. viruses, <l1' toxins c:rn bind to, and play a role r: cell-to-ceil inter:rctions. Definite changes in glr c, proteins occur in cells that are being tlansforuc into cancer cells. (Cancer is discttssed in "A Clos.: Look" on pp. 702-1.03.) Specializations of the Plasma Membrane Specializations of the plasrna 11smb1xns-such ,,. nticrouilli ancl membrane -functions-are con,monly displayed by the (epithelial) cells that for:: the linings of hollow body organs, such as the slr:r,intestine (Figure 3.3). Microvilli (mi"kro-vil'i; "litt . shaggy hairs") are tiny flngerlike projections t}'r:, llreatly increase the cell's surface area fot absorption so that the process occurs nore quickly. Membrane junctions vary structurally clcpending on their roles. o Tight junctions'are impermeable junctions thr,. bind cells together into leakproof sheets tfi.,. prevent substances from passing through th.

auerqwauJ ol

papelle

uo1e1e4so1Ac oLll

]o slueuteltj

Chapter 3: Cells and Tissues

69

:\Iracellular space between cells. In tight junc:ions. adjacent plasma membranes fuse together :ightl1. like a zipper. In the small intestine, for -rarnple, these junctions prevent digestive enz\-nies from seeping into the bloodstream.

Desmosomes (des'mo-somz) are anchoring 'unctions that prevent cells sr-rbjected to me:hanical stress (such as skin ce11s) from being :ulled apart. Structurally, these junctions are :rr-Lttonlike thickenings of adjacent plasma :lembranes (plaques), which are connected by ine protein fi1aments. Thicker protein fi1a:nents extend from the plaques inside the cells
:o the plaques on the ce11s' opposite sides, thus lorming an internal system of strong guy wires.

Desmosome (ancnoflng junction)

;Plasma 'membranes of adjacent cells
',:t,:l:rfi

Gap junctions, commonly seen in the heart :rnd belween embryonic cells, function mainly to allow communication. Chemical molecules. such as nLltrients or ions, can pass directly from one cell to another through them. In gap junctions, the neighboring cells are connected by connexons, which are hollow cylinders composed of proteins that span the entire width of the abutting membranes.

.a;:,

Connexon

Underlying basement membrane

P g f.$

spacebetween (communicating)

cells

junction

6. Why do phospholipids (which form the bulk of

cell membranes) organize into a bilayer, tail to tail, in an aqueous environment?

7. The external faces of some membrane proteins have sugar groups attached to them. What are three roles these sugar-coated proteins play in the life of a cell? 8. What is the special function of gap junctions? Of tight junctions?
For answers, see Appendix D.

# ffi # 3 " 3 Cell junctions. An epithelial cell is shown joined to adjacent cells by the three common types of cell junctions: tight junctions, desmosomes, and gap junctions. Also illustrated are microvilli (seen projecting from the free cell
surface).

The Cytoplasm
T1-re

cytoplasm is the cellular material outside the

rr,rcleus and inside the plasma membrane. It is the site of most cellular activities, so you might think of he cytoplasm as the "factory area" of the ce1l. Al-

ilrough early scientists believed that the cltoplasm rl-as a stn:ctureless gel, the electron microscope has revealed that it has three malor elements: the c_ytosol, organelles, and inclusions. The cytosol is semitransparent fluid that suspends the other elenents. Dissolved in the cytosol, which is iargely

water, are nutrients and a variety of other solutes (dissolved substances). The organelles (or"gah-nelz''), described in detail shortly, are the metabolic machinery of the cell. Each type of organelle is specialized to carry out a specific ftrnction for the cell as a whole. Some synthesize proteins, others package those proteins, and so on. Inclusions are chemical substances that may or may not be present, depending on the speciflc cell type. Most inclusions are stored nutrients or cell products. They include the lipid droplets common in fat cells, glycogen granules abundant in liver and muscle cel1s, pigments such as melanin in skin and hair cells, mucus and other secretory products, and various kinds of crysta1s.

70

Essentials of Human Anatomy and Physiology

Which nuclear component contains vour genes?

ffi
Chromat\n Nucleolus

Nuc\ear enve\ope
Nucleus Plasma

Smooth endoplasmic reticulum Cytosol Lysosome M\tochondr\on

inembrane

Rough endop\asm\c reticu\um Ribosomes

Centrioles

Microvilli

Go\giapParatus

Microfilament

li,r.tt.

"r'tii-Secretlon
Microtubule
Peroxisome

"

being released from cell by exocYtosis

lntermediate

filamenis

Structure of the generalized cell' No cell is illustrates exactly like this one, but this generalrzed cell drawing cells' human features common to many

F$#q-$ffiH

$.4

Cytoplasmic Organelles The c1'toplasmic organelles, literally "little organs"'

(Figure

3'4)' are specralized cellular compartments of life the maintain each performing its own iob to mema by the cill. Many organelles are bounded membrane similar to the plasma membrane' The to them allow organelles brane boundaries of such

maintain an internal environment quite different

from that of the surrounding cytosol' This corp partmentali zation is crucial to their ability to perus ?orm their specialized functions for the cell' Let ol workshops the of each in consider what goes on our cellular factory' Mitochondria Mitochondria (mi"to-kon'dre-ah ;;;-;"11y depicted as tinv threacllike (mitos = (see Figun thread) or sausage-shaped organelles lengthen' arx 3.4), but in living cells they squirm,

&

uteworqc

Chapter 3: Cells and Tissues

71

O n. the protein is synthesized on the

ribosome, it migrates into the rough
ER cistern.

Rough ER

tn tf,e cistern, the protein folds into its functional shape Short sugar

chains may be attached to the protein (forming a glycoprotein).

@ ffre protein is packaged

vesicle.

in a tiny

membranous sac called a transport

@ fne transport vesicle buds from the

rough ER and travels to the Golgi apparatus for further processing or goes directly to the plasma membrane where its contents are
secreted.

Protein inside transport vesicle

F

lG ei ffiffi S.

Synthesis and export of a protein bythe rough

ER.

-1ange shape almost continuously. The mitochon::ia1 wall consists of a double membrane, equal to ,:i'o p\asma membranes, placed side by side. The lter membrane is smooth and featureless, but the :r1er membrane has shelflike protrusions called :rlstae (kris'te; "crests"). Enzymes dissolved in the .--,id within the mitochondria, as well as enzymes ----it form part of the cristae membranes, carry out --e reactions in which oxTgen is used to break , -'rr-n foods. As the foods are broken down, energy .: :eleased. Much of this energy escapes as heat, - *r some is captured and used to form ATp mole,:,,Ies. ATP provides the energy for all cellular work, ,:d every living cell requires a constant supply of l3 for its many activities. Because the mitochon,::r supply most of this ATP, they are referred to as "powerhouses" of the cell. --:r ,\Ietabolically "busy" cells, like liver and muscle -:,-s. use huge amounts of ATP and have hundreds i nritochondria. By contrast, cells that are rela'.'elr- inactive (an unfertilized egg, for instance) --'.'e just a few.

Some ribosomes float free in the cytoplasm, where they manufacture proteins that function in the cytoplasm. Others attach to membranes, and the whole ribosome-membrane combination is called the rougb endoplasmic reticulwm.

Endoplasnic Beticulum The endoplasmic reticulum (en"do-plas' mik r6-tik'u-lum; "network within the cyoplasm") (ER) is a system of fluid-filled cisterns (tubules, or canals) that coil and twist through the cyoplasm. It accounts for about half of a cell,s membranes. It serves as a minicirculatory system for the cell because it provides a network of channels for carrying substances (primarily proteins) from one par-t of the cell to another. There are ttvo forms of ER; a p rLicular cell may have both forms or only one, depending on its specific functions. The rough ER is so called because it is studded with ribosomes. Because essentially all of the building materials of cellular membranes are formed either in it or on it, the rough ER can be thought of as the cell's membrane factory. The proteins made on its ribosomes migrate into the tubules of the rough ER, where they fold into their functional threedimensional shapes and then are dispatched to other areas of the cel1 in transport vesicles (Figure 3.5). Rough ER is especially abundant in cells that make

fiibosomes Ribosomes (ri'bo-somz) are tiny, - -rbed, dark bodies made of proteins and one va*:rof RNA called ribosomal Rl/,4. Ribosomes are --- actual sites of protein synthesis in the cell.

72

and Physiology Essentials of Human Anatomy

Rough ER Proteins in cisterna Membrane Lysosome fuses with

Cisterna

ingested substances

Golgi vesicle contatnrng digestive enzymes becomes a lYsosome

t.T?:

'
Golgi vesicle containing ptotein. to be secreted " becomes a secretorY vesicle

slo'Jto'Y vesicles

Golgi vesicle containing membrane comPonents fuses with the Plasma membrane Plasma membrane Extracellular fluid

F{Sl-}ffifi

tn" Role of the Golsi apparatus T:,11*:n pinch vesicles Protein-contalning transport products of the it As apparatus 'ougi'Ln' f use,with the Golgi (and off the rough ER uno'*igi;," to sorted is product the protein oasses through ,r.r" coini upparatus, packaged within vesicles' which thm is slightly modified). ff'te p"'oJ'ct various destrnations (pathways 'leave the Golgi appar-ai" unO head for 1-3), as shown.

*,s

li

example' pancreas ancl expott protein products-for to be deliv."fft, -n,.n produie digestive enzymes that catalyze enzymes ered to the small intestinl. The reside on the exlipids in" ,yr"t n"ris of membrane the needed buildternal face of the rough ER, where ing blocks are readily

vesicles lt is generassociated with swarms of tiny ;ilr"""; close to the nucleus and is the principal

;;;;ff,.

- ---^l with the smooth ER communicates Although "variety, it plays no role in protein synthe rough metabolism in"rir."i"r,eacl tt f.,,-r.tior-rr in lipid ancl breakdown)' ancl C.f..i"r,"r"1 ancl fat synthesis it is )"i""tn."ion of dntgs and pesticides ,Hence of are chock-fr-rll not surprising that the liver cells that produce cells smooth ER. So too are body instance' cells of the steroid-based hormones-for testosterone' male testes that manufacture (gol'je) apparatus Golgi Apparatus The Golgi of flattened membranous sacs' ;;;"tt'u, n

available'

Its major clirector" for cellular proteins' packege proreins (sent r.rr"r.iior_, is to mociity encl in spethe rough ER vie trunspolt r-esicles) ;;;y -*"Vt, depencling on their fina1 destination .tni /, (Ftgttrc J.t-r t. ' 'A, in proteins "taggec1" for export accumulate Then their the Golgi nppn,n"t-', the sacs su'ell' off and pinch swollen ends, filied with protein' travel which (ves' i -kuls)' form secretory vesicles the vesicles reach to the plasma membrane Vhen it' the mem;";*t"" membrane, they fuse with the sac are of brane rlrptLlres, and the contents 1 in (pathway tt the outsicle of the cell are as this wayt "1".t"a n'ig.rr" 3.6). Mucus is packaged cells' digestive enzymes -nJ" by pancreas

'tutk

Essentials of Human Anatomy and Physiology Actin subunit

@]znm
(b) Microfilament
Fibrous subunits

(c) lntermediate filament

$
.t

ild

Tubulin subunits

{'

:l:
1fi

;,!

i{
ti, .ri{

1,.,.'
(a)

(d) Microtubule

The cytoskeleton. {a) ln this light micrograph of the cytoskeleton of a nerve cell, the microtubules appear green; the microfilaments are blue. lntermedrate filaments form most of the rest of the network. (b-d) Diagrammatic views of cytoskeletal elements.

ErCffiUffiffi

ffi.?

9. How do 10.

the cytosol and the cytoplasm differ?

Which two organelles are sacs of enzymes and what is the function of each of these organelles?

11.

Which organelle is the major site of

synthesis?

ATP

12. Name the three types of protein structures that make up the cytoskeleton. Which type helps form desmosomes? Which type is involved in cell motility?
For answers, see Appendix D.

relative numbers of the various organelles thelcontain relate to specialized cell functions. Let's take a look at some of these cell specialists. L. Cells that connect body parts: . Fibroblast. The elongated shape of this cell lies along the cable-like fibers that it secretes. It has an abundant rough ER and a large Golgi apparatus to make and secrete the protein building blocks of these
fibers.

Eryttbrocyte (red blood cell). This cell carries oxygen in the bloodstream. Its concave disk shape provides extra surface area for the uptake of oxrygen and streamlines the

Gell Diversity
in this chapter, we have focused on an average human cell. However, the trillions of cells in the human body are made up of some 200 different cell types that vary greatly in size, shape, and function. They include sphere-shaped fat cells, disk-shaped red blood cells, branching nerue cells, and cube-shaped cells of kidney tubules. Figure 3.8 illustrates how the shapes of cells and the
So far

cell so it flows easily through the bloodstream. So much oxygen-carrying pigment is packed in erythroc),tes that all other organelles have been shed to make room.

Cell that covers and lines body organs! o Epithelial cell. The hexagonal shape of this cell is exactly like a "cell" in a honeycomb of a beehive. This shape allows epithelial

Chapter 3: Cells and Tissues

75

cells to pack together in sheets. An epithelial cell has abundant intermediate filaments that resist tearing when the epithelium is rubbed or pulled.
3. Cells

h ER and golgi

that move organs and body parts: Skeletal muscle and smootb muscle cells. These cells are elongated and filled with
abundant contractile filaments, so they can shorten forcefully and move the bones or change the size of internal organs.

::
Epithelialcells

%
Nucleus Nuclei

Erythrocytes

(a) Cells that connect body parts

Cell that stores nutrients: . Fat cell. The huge spherical shape of a fat cell is produced by a large lipid droplet in its c1'toplasm. Cell that fights disease: . Macrophage (a phagocytic cell). Thts cell extends long pseudopods ("false feet") to crawl through tissue to reach infection sites. The many lysosomes within the cell digest the infectious microorganisms it takes up. 6. Cell that gathers information and controls body functions: o lVetue cell (neuron). This cell has long pfocesses for receiving messages and transmitting them to other structures in the body The processes are covered with an extensive plasma membrane, and a plentiful rough ER is present to synthesize membrane components. Cells of reproduction: Oocyte (female). The largest cell in the body, this egg cell contains several copies of all organelles, for distribution to the daughter cells that arise when the fertilized egg divides to become an embryo. Sperm (male), This cell is long and streamlined, built for swimming to the egg for fertilization.Its flagellum acts as a motile whip to propel the sperm.

lntermediate filaments

(b) Cells that cover and line body organs Skeletal

muscle cell
Contractile filaments

Smooth muscle cells

(c) Cells that move organs and body parts

Fat cell

Lipid droplet
Pseudo-

pods
Nucleus

(d) Cellthat stores

(e) Cell that fights

nutrients

disease

,r,-t^to""aa_":
Nerve cell
Nucleus

(f) Cellthat gathers information and controls OoOy

functions

Cell diversity. The shape of human cells and the relative abundances of their various organelles relate to their function in the
F
aJ

lG

ffi

S.

Nucleus,*F{_Flagellum

'ffifutoq,***/
Sperm (g) Cell of reproduction

body.

78

and Physiology Essentials of Human Anatomy

what "facilitates" facilitated diffusion?

ru
Extracellular fluid

Lipidinsoluble solutes

' .,7-LiPid\ 'I soluble solutes \J

Water molecules
ii-:

.--.1

-:_

.%.

s* @* @
CytoPlasm (a) SimPle diffusion directlY through

E-fl in_

LY-: !
ci

:.-

'.{} (d) Osmosis, diffusion of water through a

specific channel protein (aquaPorin) or through the liPid bilayer

the phosPholiPid
bilayer

(c) Channel-mediated (b) Carrier-mediated facilitated facilitated diffusion carrier protetn [itrtion via through a channel t"t one chemical;

Jp".it" niJing of substrate

shape change

causes transport in

protein

protern; mostly ions t."l:t]:d on basis of size and charge

the plasma membrane' (a) ln i: i,; * iq fl 3 ' : i: Diffusion through dltfrr" directly through the lipid simple diffusion, tuuroruoiu ,-r-,olecules (b) ln in which they can dissolve bilayer of the plasma il'"tttun"' mole.lles
rarge, lipid-insoluble faciritated diff usron ,ri"g pr"i"in'"urri"r., via a transport proteln membrane (e.g., glucos") ur" tou"i across the

(c)lnfacilitateddiffusro"n-uiun.."*n'unechannels,smallpolarorcharged constructed by channel particles diffuse tf,rough t"tn'un" channels the plasma membrane via across proteins. (d) ln osmosis, water moves directly through the lipid portion specific channels taq,]aporinst or diffuses of the membrane. to pass in the clesired clirection by drl: i"tl"o They may be too large to pas's through men.tlack special probrane channels, the membrane may
r-rsually unable

into the kidney solutes fllter or-rt of the capillaries the capiilarres in ..-'r,l-,r"' because tlre bloocl pfessure fluicl pressure in the tr-rbules' Pafi ir-gt"o,"t than the
becomes flltrate formed in this way eventually most the For "rin" selective' very urine. Filtration is not large too molecules f"n, ""LV bloocl cells and protein back' held are ti-tr.r.tgh the membrane pores

teincat'riersfortheirtranspofi,theymaynotbeable

il;

Active TransPort Processes

supply to move Vhenever a cell uses some of its AIP the process is le,ubrror"t.", 2lcross the memllrane' actively are ferrecl lo as actiue substzrnces rnovecl

have to move to dissolve in the fat core, or they may i"pntn" r;tgaimst their concentration graclients The membrane two most impofiant mechanisms of active transpofi' tfanspofi are active transpofi and vesicular solrrte punxpActive Transport Sometimes called ing, active transport is similar to the carrierdescribed earlier in that -Jaat"a facilitatecl diffusion that combine both processes require protein carriers with the ,t'b'tu'-tt"t to be transported

,*"rliUfy

,&

,,"rrn

qc utelo'd to surcloLd tattte3

Chapter 3: Cells and Tissues

79

Extracellular fluid

Na+

O-@
.ATP .,..'.] @

_/ \
ADP

einOing of cytoplasmic Na" to the pump protein stimulates phosphorylation by ATP, which causes the pump protern to change its shape.

@ fne

shape change outside Extracellular K* binds causing release of the phosphate group.
Na+ to the

expels

Loss of phosphate restores the original conformation of the pump protein. K* is released to the cytoplasm and Na* sites are ready to bind Na' again: the cycle repeats. Cytoplasm

- ii EJ lt *; $ . 't 'l Operation of the sodium-potassium pump, a solute pump.ATP provides the energy for a "pump" protein to move -hree sodium ions out of the cell and two potassium ions into the cell. Both ions are moved against their concentration gradients"

rcross the membrane. However, facilitated diffr-rsion is driven by the kinetic energy of the diffusing molecules, whereas active transpofi uses ATP to energize its protein carriers, which are called solute pumps. Amino acids, some sugars, and most ions are transported by solute pumps, and in lnost cases these substances move against concentration (or electrical) gradients. This is opposite to the direction in which substances would naturally flow by diffusion, which explains the need for energy in the form of ATP. Amino acids are needed to build cellular proteins but are too large to pass through the membrane channels and are not lipidsoluble. The sodium-potassium pump that simultaneously carries sodium ions (Na+) out of and potassium ions into the cell is absolr-rtely necessary for normal transmission of impulses by nerue cells

(Figure 3.11). There are more sodium ions outside the cells than inside, so they tend to remain in the ce1l unless the cell uses ATP to force, or "pump," them out. Likewise, there are relatively more potassium ions inside cells than in the interstitial (extracellular) fluid, ancl potassium ions that leak out of cells must be actively pumped back inside. Because each of the pumps in the plasma membrane transports only specific substances, active transport provides a way for the cell to be very selective in cases where substances cannot pass by diffusion. (No pump-no transport.)
Vesicular Transport Some substances cannot get through the plasma membrane by passive trans-

port or by active transport. Vesicular tfanspoft, which involves help from ATP, moves substances

Essentials of Human Anatomy and Physiology

rl
1

l
.;

Key:

W
'**r.d

= Adenine

W
@il

visible barlike bodies called chromosomes (.chromo: colored; sorna: body) appear. Because DNA has already been replicated, each chromosome rs actually madetrp of rwo strands, each called a chromatid (kro'mah-tid), held together by a small buttonlike body called a centromere (sen'tro-mOr). The centrioles separate from each other and begin to move toward opposite sides of the cell, directing the assembly of a mitotic spindle (composed of microtubules) between them as they move' The spinclle provides a scaffolding for the attachment and movement of the chromosomes during the later mitotic stages. By the end of
prophase, the nuclear envelope and the nucleoli nurr" broken down and disappeared' and the chromosomes have attached randomly to the spindle fibers by their centromeres' Metaphase (met'ah-faz). In this short stage, the chromosomes cluster and become aligned at the metapbase plate (the center of the spindle midway between the centrioles) so that a straight line of chromosomes is seen. Anaphase Qn'ahfaz). During anaphase, the centromeres that have held the chromatids together split. The chromatids (now called chromosomes again) begin to move slowly apafi, drawn toward opposite ends of the cell' The chromosomes seem to be pulled by their halfcentromeres, with their "arms" dangling behind them. Anaphase is over when chromosome movement ends. r Telophase (tel'o-faz). Telophase is essentially prophase in reverse. The chromosomes at opposite ends of the ce1l uncoil to become threadiike chromatin again. The spindle breaks down and disappears, a nuclear envelope forms around each chromatin mass, and nucleoli appear in each of the daughter nuclei' Mitosis is basically the same in all animal cells' Depending on the type of tissue, it takes from 5 minutes to several hours to complete, but rypically it lasts about 2 hours. Centriole rephcation is deferred until late interphase of the nert ce1l cycle, when DNA replication begins before the onset of mitosis'

= Cytosine = Guanine

ii
ii
rt

;w ry:

ffi \- - x
ord

(template) synthesized

New\ \ (temPlate)r strand Old (template)

formino strand -'-"DNA of one chromatid
l

strand

strand

t'""""'v

F$ffi#ffiffi 3"$4

RePlicationof theDNA molecule during interphase. The DNA helix unwinds (center), and its nucleotide strands are separated. Each strand then acts as a template for building a new complementary strand. As a result, two helixes, each identical to the original DNA helix, are formed.

Cytokinesis Cy'tokinesis, or the division of the cytoplasm, usually begins during late anaphase and completes cluring telophase. A contractile ring made
of microfilaments fot'ms a cleavage

furrow

over the

Chapter 3: Cells and Tissues

Chromatin

Forming mitotic

Centromere

Spindle microtubules

Centromere

spindle

Chromosome, consisting of two sister chromatids

rflilerplhase

Fragments of nuclear envelope

Spindle
pole

Early prophase

Late prophase
Nucleolus

lr:rrrrrrrrf

Metapha plate

forming

l
l
Daughter chromosomes

-slka Zn"R

/ffi,,,,i *ioo\o\o
furrow .,w #.\'*ff W* = \. '/' ,/ # .,*f
Nuclear
envelope

cleavage/ *-o

' #

,l

'"ol

chromatids

$0ffiphasej,R= *."$$

Anaphase

forming Telophase and cytokinesis

Stagesof mitosis.

L:,- oi the spindle, and it eventually squeezes or cltoplasmic mass into lwo pafis. .r- -.: the end of cell division, two daughter cells -; =-,.h is smaller and has less cltoplasm than the ,.':: .e11. but it is genetically identical to it. The . --.:-. ce11s grow and carry out normal cell activi''. - r-r:' it is their turn to divide.

' -:: Il-ie original

Mitosis and division of the cyoplasm usually go hand in hand, but in some cases the cytoplasm is not divided. This condition leads to the formation of binwcleale (two nuclei) or multinucleate cells. This is fairly common in the liver. As mentioned earlier, 'mitosis provides the

"new" cells for body growth in youth and

is

Nucleus (site of transcil ption)

Cytoplasm (site of translation )

Amino acids

mRNA leaves nucleus and attaches to ribosome, and translation

Correct amino acid attached to each species of IRNA by

an enzyme

Nuclear pore Nuclear membrane

Growing

begins

the growing

Released tRN reenters the

Peptide bond

cytoplasmic pool, ready to be recharged with a new amino acid

complementary mRNA codon calling for its amino acid by binding via its anticodon to the codon
IRNA "head" bearing

anticodon Large ribosomal subunit

Portion of mRNA already translated

Small ribosomal

subunit

Directron of ribosome advance; ribosome moves the mRNA strand along sequentially as each codon

b read

LJ*# S.1#

Protein synthesis. (@) Transcription. (@-@) Translation

87

Chapter

3: Cells and

91

\
Air sacs of lungs Nucleus of squamous epithelial cell Nuclei of squamous epithelial

cells
Basement membrane

Photomicrograph : Simple squamous

epithelium forming part of the alveolar
(air sac)walls (100x).

(a) Diagram: Simple squamous

Basement membrape

Simple

cuboidal
epithelial cells Basement membrane Connective tissue

Nucleus of simple

cuboidal
epithelial
nall

(b) Diagram: Simple cuboidal

Photomicrograph : Simple cuboidal epithelium in kidney tubules (a00x).

Nucleus of simple columnar epithelial cell

Simple

columnar
epithelial cell

Basement membrane

Basement membrane

Photomicrograph : Simple columnar

epithelium of the stomach lining (900x) (c) Diagram: Simple columnar

FIGUfl fi 3. n #

Types of epithelia and their common locations in the bodV. (Continued on page 92.)

Essentials.of Human Anatomy and Physiology

t,
ll l

Pseudostratified epithelial layer

Cilia Mucus of goblet cell Pseudostratified epithelial layer Basement membrane Connective tissue

Basement membrane

(d) Diagram: Pseudostratified (ciliated) columnar

Photomicrograph : Pseudostratif ied

ciliated columnar epithelium lining the human trachea (7OOx).

Stratified squamous epithelium Basement membrane Basement membrane Connective tissue

Photomicrograph : Stratif ied

(e) Diagram: Stratified squamous squamous epithelium lining of the esophagus (200x).

Basement membrane

Basement membrane

Transitional epithelium
Connective tissue

Photomicrograph: Transitional (f) Diagram : Transitional

epithelium lining of the bladder, relaxed state (300x); note the rounded appearance of the cells at the surface; these cells flatten and elongate when the bladder fills with urine Types of epithelia and their

FeffiUffiffi 3" ,E * f**re*ff?sjedj

common locations in the body.

Essentials of Human Anatomy and physiology ?nonomers, of these fibers are made by the connective tissue cells and secreted into the ground substance in the extracellular space, where they join together to form the various fiber types. Because of its extracellular matrix, connective tissue is able to form a soft packing tissue around other organs, to bear weight, and to withstand stretching and other abuses, such as abrasion, that no other tissue could endure. But there is variation. At one extreme, fat tissue is composed mostly of cells, and the matrix is soft. At the opposite extreme, bone and cafi1lage have very few cells and large amounts of hard matrix, which makes them extremely strong. Find the various types of connective tissues in Figure 3.19 as you read their descriptions, which follow.

layers of a very hard matrix that contains calcium salts in addition to large numbers of collagen fibers (Figure 3.79a). Because of its rocklike hardness. bone has an exceptional ability to protecr and support other body organs (for example, the Jkuil protects the brain).

Types of Connective Tissue

As noted above, all connective tissues consist of liv-

ing cells surrounded by a matrix. Their major differences reflect fiber type and the number of fibers in the matrix. From most rigid to softest, the major connective tissue classes are bone, cartilage, dense connectiue tisswe, Ioose connectiue tissue, and blood.

Bone Bone, sometimes called osseous (os,e-us) tissue, is composed of bone cells sitting in cavities called lacunae (lah-ku'ne; "pits") and surrounded by

Caftilage Cafiilage is less hard and more flexible than bone. It is found in only a few places in the body. Most widespread is hyaline (hi' ahlin) cartilage, which has abundant collagen fibers hidden by a rubbery matrix with a glassy (hyalin : glass), blue-white appearance (Figure 3.19b).It forms the supporting structures of the larynx, or voice box, attaches the ribs to the breastbone, and covers the ends of many bones, where they form joints. The skeleton of a fetus is made largely of hyaline car-tilage; but, by the time the baby is born, most of that carrilage has been replaced by bone. Although hyaline cartilage is the mosr abun_ dant type of cartilage in the body, there are others. Highly compressible fibrocartilage forms the cushionlike disks between the vertebrae of the spinal column (Figure 3.79c). Elastic cartilage is found where a structure with elasticity is desired. For example, it supports the external ear. (Elastic cartilage is not illustrated in Figure 3.I9.)
(Continues on page 97-)

Bone cells in lacunae Central canal

Lacunae Lamella

(a) Diagram: Bone

Photomicrograph: Cross_sectional view of ground bone (250x).

FPffi UFqffi #" locations.

$*

Gonnective tissues and their common body

Chapter

3: Cells and Tissues

95

',,$ry/,,'

W
Lacunae

Chonflrocyte
in lacu'na

Matrix

(b) Diagram: Hyaline cartilage

Photomicrograph: Hyaline cartilage from the trachea (a00x).

Chondrocytes
in lacunae
Cf,tondro-

ri(es in
lacunae Collagen Collagen fiber

fbers
(c) Diagram: Fibrocartilage

Photomicrograph: Fibrocartilage of an intervertebral disc (200x).

Ligament
Tendon

Collagen

toers
*broblasts

Nuclei of

fibroblasts

Nucleiof

(d) Diagram: Dense fibrous

Photomicrograph: Dense fibrous connective tissue from a tendon (500x). (Continues on page 96 )

Mucosa epithelium Lamina

propria

Elastic ibers

Collagen

fibers
Fibers of matrix Nuclei oj Fibroblast nuclei

fibroblasts

(e) Diagram: Areolar

Photomicrograph: Areolar connective tissue, soft packaging tissue of the body (330x).

Nuclei of fat cells

Nuclei of

fat cells
Vacuole Vacuole

containing fat droplet (f) Diagram: Adipose

containing fat droplet

Photomicrograph: Adipose tissue from the subcutaneous layer beneath the skin (330x).

White blood cell Reticular cell Blood cell Reticular

(lymphocyte)
Reticular fibers

fibers (g) Diagram: Reticular

Photomicrograph: Dark-stainin g network

of reticular connective tissue
(aOOx).

r:&u*F S."ES
96

f**raf'ssaaieCj Connective tissues and their common body locations. (e, f, and g are subclasses of loose connective tissues.)

Chapter

3: Cells and Tissues

Blood cells in capillary

%
ffiffiffi

Neutrophil (white blood cell) Red blood cells Monocyte (white blood cell)

cell

: J cells

(h) Diagram: Blood

Photomicrograph: Smear of human blood (13OOx)

" 1 .1

-,,r,sTiJ

3L

-:-

lcrrse Connective Tissue 0trm-se connective tissue, also called dense 5fr'r'ous tissue, has collagen fibers as its main
"r-i -j'- element (Figure 3.19d). Crowded belween

-i:igen fibers are rows of Jibrctbla.s/s (fibercells) that manufacrure the building blocks '= ibers. ' Dense connective tissue forms strong, ' -n:.-..e structures such as tendons ancl liga'r,:''--: Tfldons attach skeletal muscles to bones; eg"-ents connect bones to bones at ioints. Liga:rr:"- : lre more stretchy and contain more elastic T'r:- :han tendons. Dense connective tissue also 'r-.i !-:> up the lower layers of the skin (dermis), ;, ' : : li is arranged in sheets.

'-" :-l

:-

-o@se Connective Tissue

fluid matrix conrains all types of fibers, which form a loose network. In fact, when viewed through a microscope, most of the matrix appears to be empty space, which explains the name of this tis_ slre type (.areola: small open space). Because of its loose and fluid nature, areolar connective tissue provides a reseloir of water and salts for the sur_ rounding tissues, and essentially ali body cells obtain their nutrients from and release their wastes into this "tissue fluid." \7hen a body region is inflamed, the areolar tissue in the area soaks up tire excess fluid like a sponge, and the area swells and becomes puffy, a condition called edema. Many types of phagocytes wander through rhis tis_ sue, scavenging for bacteria, dead cells, and other debris, which they destroy.
Adipose Tissue Adipose (ad,i-p6s) tissue is com_ monly caIled fat. Basically, it is an areolar tissue in

r .. --" clr- speaking, loose connective tissues are ', r:- -rrtcl have more cells and fewer fibers than iir ':,er connective tissue type except blood. *mlar Tissue Areolar (ah-re'o-lar) tissue, the -: :..'16lsly distributed connective tissue variety in r -, .,h-. is a soft, pliable, ,,cobwebby,' tissue that
,

rrr -.:r positions. A soft layer of areolar connective - - c:rlled the lamina propria (lah,mi-nah ," -i;-rrh) undedies all mucous membranes. Its

.- : -rnd connective tissue "glue" because it helps ,: the internal organs together and in their

' -'-:c 3.19). It functions as a universal packing

which fat cells predominate (Figure 3.I9D. A glis_ tening droplet of oil occupies most of a fat cell,s voiume and compresses the nucleus, clisplacing it to one side. Becarrse the oil-containing region looks empty and the thin rim of cltoplasm con_ taining the bulging nucleus looks like a ring with a seai, fat ceils are sometimes called signet ring cells. Adipose tissue forms the subcutaneous tissue beneath the skin, where it insulates the body and protects it from bumps and extremes of both heat
and cold. Adipose tissue also protects some organs

Cell division typically yrelds two daughter cells, each with one nucleus. How is the multinuclear condition of skeletal muscle explained?

Part of muscle

fiber

(a) Diagram: Skeletal muscle

Photomicrograph: Skeletal muscle (approx. 250x).

lntercalated

discs

Nucleus

(b) Diagram: Cardiac muscle

Photomicrograph: Cardiac muscle (BOOx).

Smooth

muscle cell
Nuclei

(c) Diagram: Smooth muscle

Photomicrograph: Sheet of smooth muscle (approx. 250x).

FIGURK S.H#
&
,,r"r,
olAc Aq

'types of muscle tissue and their common locations in the body.

paruedwocreun s*ortLu oorcpun fipaleadeL srpo apsnLt) telap4s

It
100
Essentials of Human Anatomy and Physiology
Brain

Nuc Spinal

supc:
cel
s

:'

cord

Y'
Nuclei of Ce I c::

supporting
cells

of ne-.':

Cell body
of neuron Neuron

processes

Diagram: Nervous tissue

Photomicrograph : Neurons (200x)

Fi#tiffiffi fr.f'1 Nervous tissue. Neurons and supporting cells form the brain, spinal cord, and nerves.

Tissue Repair
{Woqrnd *flealing}
The body has many techniques for protecting itself from uninvited guests or injury. Intact physical barriers such as the skin and mucous membranes, cilia, and the strong acid produced by stomach glands are just three examples of body defenses exerted at the iocal tissue level. When tissue injury does occur, it stimulates the body's inflammatory and immune responses, and the healing process begins almost immediately. Inflammation is a generalized (nonspecific) body response that attempts to prevent further injury. The immune response, in contrast, is extremely specific and mounts a vigorous attack against recognized invaders (bacteria, viruses, toxins). These protective responses afe considered in detail in Chapter 12. Here we will concentrate on the process of tissue repair itself. Tissue repair, or wound healing, occurs in lwo major ways: by regeneration and by fibrosis. Regeneration is the replacement of destroyed tissue by the same kind of cells, whereas fibrosis involves repair by dense (fibrous) connective tissue, that is, by the formation of scar tissue. \7hich occurs depends on (1) the type of tissue damaged

and

(.2')

the severity of the injury. Generaliy speaksuc-

ing, clean cuts (incisions) heal much more

cessfully than ragged tears of the tissue. Tissue injury sets a series of events into motion:

The capillaries become very permeatrle. This allows fluid rich in clotting proteins and other substances to seep into the injured area from the bloodstream. Then leaked clotting
proteins construct a clot, which stops the loss of blood, holds the edges of the s-ound together, and walls off the injured area. prevent, ing bacteria or other harmful sr-rbsr..rnces from spreading to surrounding tissue:. \\ here the clot is exposed to air, it quickh --r':c< rtnd hardens, forming a scab. Granulation tissue forms.'''. t'.;. ;',,', t I i (.t t.t tissue is a delicate pink rissuc , :-,-,:- :cl largely of new capillaries that 91' r. -- - . : _.-.r:-rged area from undamagecl 1- - :-,: - r--:-:ibr'. These capillaries ale fi':ts = .'.- -. --:c,\-. as when a scab is pickc* -, .-1n wound. Granulatior-r lrr- --: evenrualh .-:-: i , .-, -lot and connecti\etti>-": ::hltsynthesize
.

ti-ie bur.::.:-,Sller l i-:1

-::- iiters
.: ::lp.

(scar tis-

_:-&4.l+ffi