17

The Special Senses

An Introduction to the Special Senses 550
Olfaction 550
Olfactory Receptors 551
Olfactory Pathways 551
Olfactory Discrimination 551
Gustation 552
Taste Receptors 553
Gustatory Pathways 553
Gustatory Discrimination 553
Key 55
Vision 55
!ccessory Structures of the "ye 55
The "ye 557
Key 5##
$isual Physiolo%y 5##
The $isual Pathway 571
Equilibrium and Hearing 573
!natomy of the "ar 573
"&uili'rium 57#
(earin% 57)
Key 5*#
Chapter e!ie" 5*#
Clinical #otes
Dia'etic Retinopathy 55)
Detache+ Retina 5#1
Glaucoma 5#2
!ccommo+ation Pro'lems 5#
248927081.doc 1
An Introduction to the Special Senses
Our ,nowle+%e of the worl+ aroun+ us is limite+ to those characteristics that stimulate our
sensory receptors- !lthou%h we may not reali.e it/ our picture of the en0ironment is
incomplete- 1olors we cannot +istin%uish %ui+e insects to flowers2 soun+s we cannot hear
an+ smells we cannot +etect pro0i+e +olphins/ +o%s/ an+ cats with important information
a'out their surroun+in%s-
3hat we do percei0e 0aries consi+era'ly with the state of our ner0ous systems- 4or
e5ample/ +urin% sympathetic acti0ation/ we e5perience a hei%htene+ awareness of sensory
information an+ hear soun+s that woul+ normally escape our notice- 6et/ when concentratin%
on a +ifficult pro'lem/ we may remain unaware of relati0ely lou+ noises- 4inally/ our
perception of any stimulus reflects acti0ity in the cere'ral corte5/ an+ that acti0ity can 'e
inappropriate- 7n cases of phantom lim' pain/ for e5ample/ a person feels pain in a missin%
lim'/ an+ +urin% an epileptic sei.ure/ an in+i0i+ual may e5perience si%hts/ soun+s/ or smells
that ha0e no physical 'asis-
Our +iscussion of the %eneral senses an+ sensory pathways in 1hapter 15 intro+uce+ 'asic
principles of receptor function an+ sensory processin%- 3e now turn our attention to the fi0e
special senses: olfaction/ %ustation/ 0ision/ e&uili'rium/ an+ hearin%- !lthou%h the sense
or%ans in0ol0e+ are structurally more comple5 than those of the %eneral senses/ the same
'asic principles of receptor function apply- !T8!S9 "m'ryolo%y Summary 139 The
De0elopment of Special Sense Or%ans
Olfaction
Objectives
: Descri'e the sensory or%ans of smell an+ trace the olfactory pathways to their +estinations
in the 'rain-
: "5plain what is meant 'y olfactory +iscrimination an+ 'riefly +escri'e the physiolo%y
in0ol0e+-
The sense of smell/ more precisely calle+ olfaction/ is pro0i+e+ 'y paire+ olfactory organs-
These or%ans are locate+ in the nasal ca0ity on either si+e of the nasal septum ;4i%ure 17<
1a:=- The olfactory or%ans are ma+e up of two layers9 the olfactory epithelium an+ the
248927081.doc 2
lamina propria- The olfactory epithelium ;4i%ure 17<1':= contains the olfactory receptor
cells/ supportin% cells/ an+ re%enerati0e basal cells ;stem cells=- 7t co0ers the inferior surface
of the cri'riform plate/ the superior portion of the perpen+icu
lar plate/ an+ the superior nasal conchae of the ethmoi+- lp$ %&' The un+erlyin% lamina
propria consists of areolar tissue/ numerous 'loo+ 0essels/ an+ ner0es- This layer also
contains olfactory glands/ or Bowmans glands/ whose secretions a'sor' water an+ form a
thic,/ pi%mente+ mucus-
3hen you inhale throu%h your nose/ the air swirls an+ e++ies within the nasal ca0ity/ an+ this
tur'ulence 'rin%s air'orne compoun+s to your olfactory or%ans- ! normal/ rela5e+ inhalation
carries a small sample of the inhale+ air ;a'out 2 percent= to the olfactory or%ans- Sniffin%
repeate+ly increases the flow of air across the olfactory epithelium/ intensifyin% the
stimulation of the olfactory receptors- (owe0er/ those receptors can 'e stimulate+ only 'y
water>solu'le an+ lipi+>solu'le materials that can +iffuse into the o0erlyin% mucus-
Olfactory eceptors
Olfactory receptors are hi%hly mo+ifie+ neurons- The e5pose+ tip of each receptor cell forms
a prominent ,no' that pro?ects 'eyon+ the epithelial surface ;see 4i%ure 17<1':=- The ,no'
pro0i+es a 'ase for up to 20 cilia that e5ten+ into the surroun+in% mucus- These cilia lie
parallel to the epithelial surface/ e5posin% their consi+era'le surface area to +issol0e+
compoun+s-
Olfactory reception occurs on the surfaces of the olfactory cilia as +issol0e+ chemicals
interact with receptors/ calle+ odorant-binding proteins/ on the mem'rane surface-
Odorants are chemicals that stimulate olfactory receptors- 7n %eneral/ o+orants are small
or%anic molecules2 the stron%est smells are associate+ with molecules of hi%h solu'ility 'oth
in water an+ in lipi+s- The receptors in0ol0e+ are G proteins2 'in+in% of an o+orant to its
receptor lea+s to the acti0ation of a+enylate cyclase/ the en.yme that con0erts
!TP to cyclic>!@P ;c!@P=- lp$ (&& The c!@P then opens so+ium channels in the
mem'rane/ resultin% in a locali.e+ +epolari.ation- 7f sufficient +epolari.ation occurs/ an
action potential is tri%%ere+ in the a5on/ an+ the information is relaye+ to the 1AS-
Between 10 an+ 20 million olfactory receptors are pac,e+ into an area of rou%hly 5 cm
2
;0-*
in-
2
= 7f we ta,e into account the e5pose+ ciliary surfaces/ the actual sensory area pro'a'ly
248927081.doc 3
approaches that of the entire 'o+y surface- Ae0ertheless/ our olfactory sensiti0ities cannot
compare with those of other 0erte'rates such as +o%s/ cats/ or fishes- ! German shepher+
+o% sniffin% for smu%%le+ +ru%s or e5plosi0es has an olfactory receptor surface 72 times
%reater than that of the near'y customs inspectorC
Olfactory )ath"ays
The olfactory system is 0ery sensiti0e- !s few as four o+orant molecules can acti0ate an
olfactory receptor- (owe0er/ the acti0ation of an afferent fi'er +oes not %uarantee an
awareness of the stimulus- 1onsi+era'le con0er%ence occurs alon% the olfactory pathway/
an+ inhi'ition at the inter0enin% synapses can pre0ent the sensations from reachin% the
olfactory cortex of the cere'ral hemispheres- lp$ (*( The olfactory receptors themsel0es
a+apt 0ery little to a persistent stimulus- Rather/ it is central a+aptation which ensures that
you &uic,ly lose awareness of a new smell 'ut retain sensiti0ity to others-
!5ons lea0in% the olfactory epithelium collect into 20 or more 'un+les that penetrate the
cri'riform plate of the ethmoi+ 'one to reach the olfactory bulbs of the cere'rum ;see
4i%ure 17<1:=/ where the first synapse occurs- "fferent fi'ers from nuclei elsewhere in the
'rain also inner0ate neurons of the olfactory 'ul's- This arran%ement pro0i+es a mechanism
for central a+aptation or facilitation of olfactory sensiti0ity- !5ons lea0in% the olfactory 'ul'
tra0el alon% the olfactory tract to reach the olfactory corte5/ the hypothalamus/ an+ portions
of the lim'ic system-
Olfactory stimulation is the only type of sensory information that reaches the cere'ral corte5
+irectly2 all other sensations are relaye+ from processin% centers in the thalamus- The parallel
+istri'ution of olfactory information to the lim'ic system an+ hypothalamus e5plains the
profoun+ emotional an+ 'eha0ioral responses/ as well as the memories/ that can 'e tri%%ere+
'y certain smells- The perfume in+ustry/ which un+erstan+s the practical implications of
these connections/ e5pen+s consi+era'le effort to +e0elop o+ors that tri%%er se5ual
responses-
Olfactory +iscrimination
The olfactory system can ma,e su'tle +istinctions amon% 2000<000 chemical stimuli- Ao
apparent structural +ifferences e5ist amon% the olfactory cells/ 'ut the epithelium as a whole
248927081.doc 4
contains receptor populations with +istinct sensiti0ities- !t least 50 Dprimary smellsE are
,nown/ an+ it is almost impossi'le to +escri'e these sensory impressions effecti0ely- 7t
appears li,ely that the 1AS interprets each smell on the 'asis of the o0erall pattern of
receptor acti0ity-
!lthou%h the human olfactory or%ans can +iscriminate amon% many smells/ acuity 0aries
wi+ely/ +epen+in% on the nature of the o+orant- @any o+orants are +etecte+ in ama.in%ly
small concentrations- One e5ample is 'eta>mercaptan/ an o+orant commonly a++e+ to
natural %as/ propane/ an+ 'utane/ which are otherwise o+orless- Because we can smell 'eta>
mercaptan in e5tremely low concentrations ;a few parts per 'illion=/ its a++ition ena'les us
to +etect a %as lea, almost at once an+ ta,e steps to pre0ent an e5plosion-
!%in% an+ Olfactory Sensiti0ity
The olfactory receptor population un+er%oes consi+era'le turno0er2 new receptor cells are
pro+uce+ 'y the +i0ision an+ +ifferentiation of 'asal cells in the epithelium- This turno0er is
one of the few e5amples of neuronal replacement in a+ult humans- Despite this process/ the
total num'er of receptors +eclines with a%e/ an+ the remainin% receptors 'ecome less
sensiti0e- !s a result/ el+erly in+i0i+uals ha0e +ifficulty +etectin% o+ors in low
concentrations- This +ecline in the num'er of receptors accounts for Gran+motherFs
ten+ency to use too much perfume an+ e5plains why Gran+fatherFs aftersha0e seems so
stron%9 They must apply more to 'e a'le to smell it-
Gustation
Objectives
: Descri'e the sensory or%ans of taste an+ trace the %ustatory pathways to their +estinations
in the 'rain-
: "5plain what is meant 'y %ustatory +iscrimination an+ 'riefly +escri'e the physiolo%ic
processes in0ol0e+-
Ge
248927081.doc 5
Gustation/ or taste/ pro0i+es information a'out the foo+s an+ li&ui+s we consume- ,aste
receptors/ or gustatory ;GHS>ta>tor>= receptors/ are +istri'ute+ o0er the superior surface of
the ton%ue an+ a+?acent portions of the pharyn5 an+ laryn5- The most important taste
receptors are on the ton%ue2 'y the time we reach a+ulthoo+/ the taste receptors on the
pharyn5/ laryn5/ an+ epi%lottis ha0e +ecrease+ in importance an+ a'un+ance- Taste receptors
an+ speciali.e+ epithelial cells form sensory structures calle+ taste buds- !n a+ult has a'out
3000 taste 'u+s-
The superior surface of the ton%ue 'ears epithelial pro?ections calle+ lingual papillae ;pa>
P78>
2
Ge
papilla/ a nipple>shape+
moun+=- The human ton%ue 'ears three types of lin%ual papillae ;4i%ure 17<2:=9 ;1= filiform
;filum/ threa+= papillae/ ;2= fungif-
Ga
orm ;fungus/ mushroom= papillae/ an+ ;3= circum!allate ;sir>,um>$!8>t= papillae
;circum>/ aroun+ tri'ution of these lin%ual papillae 0aries 'y re%ion- 4iliform papillae pro0i+e
friction that helps the ton%ue mo0e o'?ects aroun+ in the mouth/ 'ut +o not contain taste
'u+s- "ach small fun%iform papilla contains a'out fi0e taste 'u+s2 each lar%e circum0allate
papilla contains as many as 100 taste 'u+s- The circum0allate papillae form a $ near the
posterior mar%in of the ton%ue-
,aste eceptors
Taste 'u+s are recesse+ into the surroun+in% epithelium/ isolate+ from the relati0ely
unprocesse+ contents of the mouth- "ach taste 'u+ ;4i%ure 17<2'/c:= contains a'out 0
slen+er/ spin+le>shape+ cells of at least four +ifferent types- .asal cells appear to 'e stem
cells- These cells +i0i+e to pro+uce +au%hter cells that mature in sta%es2 the cells of the last
sta%e are calle+ gustatory cells- "ach %ustatory cell e5ten+s slen+er micro0illi/ sometimes
calle+ taste hairs/ into the surroun+in% flui+s throu%h the taste pore/ a narrow openin%-
Despite this relati0ely protecte+ position/ itFs still a har+ life9 ! typical %ustatory cell sur0i0es
for only a'out 10 +ays 'efore it is replace+- !lthou%h e0eryone a%rees that %ustatory cells
248927081.doc 6
are taste receptors/ it is not clear whether the cells at earlier sta%es of +e0elopment also
pro0i+e taste information- ;1ells at all three sta%es are inner0ate+ 'y sensory neurons-=
I vallum/ wall=- The +is
Gustatory )ath"ays
Taste 'u+s are monitore+ 'y cranial ner0es $77 ;facial=/ 7J ;%lossopharyn%eal=/ an+ J
;0a%us=- The facial ner0e monitors all the taste 'u+s locate+ on the anterior two>thir+s of the
ton%ue/ from the tip to the line of circum0allate papillae- The circum0allate papillae an+ the
posterior one>thir+ of the ton%ue are inner0ate+ 'y the %lossopharyn%eal ner0e- The 0a%us
ner0e inner0ates taste 'u+s scattere+ on the surface of the epi%lottis- The sensory afferents
carrie+ 'y these cranial ner0es synapse in the solitary nucleus of the me+ulla o'lon%ata/ an+
the a5ons of the postsynaptic neurons enter the me+ial lemniscus- There/ the neurons ?oin
a5ons that carry somatic sensory information on touch/ pressure/ an+ proprioception- !fter
another synapse in the thalamus/ the information is pro?ecte+ to the appropriate portions of
the primary sensory corte5-
! conscious perception of taste is pro+uce+ as the information recei0e+ from the taste 'u+s
is correlate+ with other sensory +ata- 7nformation a'out the te5ture of foo+/ alon% with
taste>relate+ sensations such as DpepperyE or D'urnin% hot/E is pro0i+e+ 'y sensory afferents
in the tri%eminal ner0e ;$=- 7n a++ition/ the le0el of stimulation from the olfactory receptors
plays an o0erwhelmin% role in taste perception- Thus/ you are se0eral thousan+ times more
sensiti0e to DtastesE when your olfactory or%ans are fully functional- By contrast/ when you
ha0e a col+ an+ your nose is stuffe+ up/ air'orne molecules cannot reach your olfactory
receptors/ so meals taste +ull an+ unappealin%- This re+uction in taste perception occurs
e0en thou%h the taste 'u+s may 'e respon+in% normally-
Gustatory +iscrimination
6ou are pro'a'ly alrea+y familiar with the four primary taste sensations9 sweet/ salty/ sour/
an+ 'itter- There is some e0i+ence for +ifferences in sensiti0ity to tastes alon% the a5is of the
ton%ue/ with %reatest sensiti0ity to salty<sweet anteriorly an+ sour<'itter posteriorly-
(owe0er/ there are no +ifferences in the structure of the taste 'u+s/ an+ taste 'u+s in all
portions of the ton%ue pro0i+e all four primary taste sensations-
248927081.doc 7
(umans ha0e two a++itional taste sensations that are less wi+ely ,nown9
: /mami$ /mami ;oo>@!(>m = is a pleasant taste that is characteristic of 'eef 'roth/
chic,en 'roth/ an+ parmesan cheese- This
eGtaste is +etecte+ 'y receptors sensiti0e to the presence of amino aci+s ;especially
%lutamate=/ small pepti+es/ an+ nucleoti+es- The +istri'ution of these receptors is not ,nown
in +etail/ 'ut they are present in taste 'u+s of the circum0allate papillae-
: 0ater$ @ost people say that water has no fla0or- (owe0er/ research on humans an+
other 0erte'rates has +emonstrate+ the presence of "ater receptors/ especially in the
pharyn5- The sensory output of these receptors is processe+ in the hypothalamus an+ affects
se0eral systems that affect water 'alance an+ the re%ulation of 'loo+ 0olume- 4or e5ample/
minor re+uctions in !D( secretion occur each time you ta,e a lon% +rin,-
The mechanism 'ehin+ %ustatory reception resem'les that of olfaction- Dissol0e+ chemicals
contactin% the taste hairs 'in+ to receptor proteins of the %ustatory cell- The +ifferent tastes
in0ol0e +ifferent receptor mechanisms- Salt receptors an+ sour receptors are chemically
%ate+ ion channels whose stimulation pro+uces +epolari.ation of the cell- Receptors
respon+in% to stimuli that pro+uce sweet/ 'itter/ an+ umami sensations are G proteins calle+
gustducins ;GHST>+oos>in.=Kprotein comple5es that use secon+ messen%ers to pro+uce
their effects- The en+ result of taste receptor stimulation is the release of neurotransmitters
'y the receptor cell- The +en+rites of the sensory afferents are ti%htly wrappe+ 'y fol+s of
the receptor cell mem'rane/ an+ neurotransmitter release lea+s to the %eneration of action
potentials in the afferent fi'er- Taste receptors a+apt slowly/ 'ut central a+aptation &uic,ly
re+uces your sensiti0ity to a new taste-
The threshol+ for receptor stimulation 0aries for each of the primary taste sensations/ an+
the taste receptors respon+ more rea+ily to unpleasant than to pleasant stimuli- 4or e5ample/
we are almost a thousan+ times more sensiti0e to aci+s/ which taste sour/ than to either
sweet or salty chemicals/ an+ we are a hun+re+ times more sensiti0e to 'itter compoun+s
than to aci+s- This sensiti0ity has sur0i0al 0alue/ 'ecause aci+s can +ama%e the mucous
mem'ranes of the mouth an+ pharyn5/ an+ many potent 'iolo%ical to5ins ha0e an e5tremely
'itter taste-
248927081.doc 8
Taste sensiti0ity +iffers si%nificantly amon% in+i0i+uals- @any con+itions relate+ to taste
sensiti0ity are inherite+- The 'est>,nown e5ample in0ol0es sensiti0ity to the compoun+
phenylthiourea/ also ,nown as phenylthiocarbamide/ or ),C- Rou%hly 70 percent of
1aucasians can taste this su'stance2 the other 30 percent are una'le to +etect it-
!%in% an+ Gustatory Sensiti0ity
Our tastin% a'ilities chan%e with a%e- 3e 'e%in life with more than 10/000 taste 'u+s/ 'ut
the num'er 'e%ins +eclinin% +ramatically 'y a%e 50- The sensory loss 'ecomes especially
si%nificant 'ecause/ as we ha0e alrea+y note+/ a%in% in+i0i+uals also e5perience a +ecline in
the num'er of olfactory receptors- !s a result/ many el+erly people fin+ that their foo+ tastes
'lan+ an+ unappeti.in%/ whereas chil+ren ten+ to fin+ the same foo+s too spicy-
&11 Keys 2 Olfactory information is route+ +irectly to the cere'rum/ an+ olfactory stimuli
ha0e powerful effects on moo+ an+ 'eha0ior- Gustatory sensations are stron%est an+ clearest
when inte%rate+ with olfactory sensations-
Concept Chec3
3hen you first enter the !LP la' for +issection/ you are 0ery aware of the o+or of
preser0ati0es- By the en+ of the la' perio+/ the smell +oesnFt seem to 'e nearly as stron%-
3hyM
7f you completely +ry the surface of your ton%ue an+ then place salt or su%ar crystals on
it/ you canFt taste them- 3hy notM
6our %ran+father canFt un+erstan+ why foo+s he use+ to en?oy ?ust +onFt taste the same
anymore- (ow woul+ you e5plain this to himM
Ans"ers begin on p$ A4&
Vision
Objectives
: 7+entify the accessory structures of the eye an+ e5plain their functions-
: Descri'e the internal structures of the eye an+ e5plain their functions-
: "5plain how we are a'le to +istin%uish colors an+ percei0e +epth-
248927081.doc 9
: "5plain how li%ht stimulates the pro+uction of ner0e impulses an+ trace the 0isual
pathways to their +estinations in the 'rain-
3e rely more on 0ision than on any other special sense- Our 0isual receptors are containe+
in the eyes/ ela'orate structures that ena'le us not only to +etect li%ht/ 'ut also to create
+etaile+ 0isual ima%es- 3e will 'e%in our +iscussion of these fascinatin% or%ans 'y
consi+erin% the accessory structures of the eye/ which pro0i+e protection/ lu'rication/ an+
support-
Accessory Structures of the Eye
The accessory structures of the eye inclu+e the eyeli+s an+ the superficial epithelium of the
eye/ an+ the structures associate+ with the pro+uction/ secretion/ an+ remo0al of tears-
4i%ure 17<3: shows the superficial anatomy of the eye an+ its accessory structures-
"yeli+s an+ Superficial "pithelium of the "ye
The eyeli+s/ or palpebrae ;pal>P
G"
>'r
Ge
=/ are a continuation of the s,in- Their continual 'lin,in% ,eeps the surface of the eye lu
'ricate+/ an+ they act li,e win+shiel+ wipers/ remo0in% +ust an+ +e'ris- The eyeli+s can also
close firmly to protect the +elicate surface of the eye- The palpebral fissure is the %ap that
separates the free mar%ins of the upper an+ lower eyeli+s- The two eyeli+s are connecte+/
howe0er/ at the medial canthus ;N!A>thus= an+ the lateral canthus ;4i%ure 17<3a:=- The
eyelashes/ alon% the mar%ins of the eyeli+s/ are 0ery ro'ust hairs that help pre0ent forei%n
matter ;inclu+in% insects= from reachin% the surface of the eye-
The eyelashes are associate+ with unusually lar%e se'aceous %lan+s- !lon% the inner mar%in
of the li+/ mo+ifie+ se'aceous
GO
G
248927081.doc 10
%lan+s calle+ tarsal glands/ or Meibomian ;m >B
O stic,in% to%ether- ;These %lan+s are too small to 'e seen in 4i%ure 17<3:-= !t the me+ial
canthus/ the lacrimal caruncle ;N!R>un%>,ul=/ a mass of soft tissue/ contains %lan+s
pro+ucin% the thic, secretions that contri'ute to the %ritty +eposits that sometimes appear
after a %oo+ ni%htFs sleep- These 0arious %lan+s are su'?ect to occasional in0asion an+
infection 'y 'acteria- ! chalazion
Ge
>an= glands/ secrete a lipi+>rich pro+uct that helps ,eep the eyeli+s from >m
;,ah>8!G
>.
Ge
>on2 small lump=/ or cyst/ %enerally results from the infection of a tarsal %lan+- !n infection
in a se'aceous %lan+ of
one of the eyelashes/ a tarsal %lan+/ or one of the many sweat %lan+s that open to the surface
'etween the follicles pro+uces a painful locali.e+ swellin% ,nown as a sty-
The s,in co0erin% the 0isi'le surface of the eyeli+ is 0ery thin- Deep to the s,in lie the
muscle fi'ers of the orbicularis oculi an+ levator palpebrae superioris muscles- lp$ 556
These s,eletal muscles are responsi'le for closin% the eyeli+s an+ raisin% the upper eyeli+/
respecti0ely-
The epithelium co0erin% the inner surfaces of the eyeli+s an+ the outer surface of the eye is
calle+ the con7uncti!a ;,on>?un,>
G7
T >0uh=- 7t is a mucous mem'rane co0ere+ 'y a speciali.e+ stratifie+ s&uamous epithelium-
The palpebral con7uncti!a co0ers the inner surface of the eyeli+s/ an+ the ocular
con7uncti!a/ or bulbar conjunctiva/ co0ers the anterior surface of the eye
Ge
;4i%ure 17<3':=- The ocular con?uncti0a e5ten+s to the e+%es of the cornea ;NOR>n >uh=/ a
transparent part of the outer fi'rous layer of the eye- The cornea is co0ere+ 'y a 0ery
+elicate s&uamous corneal epithelium/ fi0e to se0en cells thic,/ that is continuous with the
ocular con?uncti0a- ! constant supply of flui+ washes o0er the surface of the eye'all/
,eepin% the ocular con?uncti0a an+ cornea moist an+ clean- Go'let cells in the epithelium
248927081.doc 11
assist the accessory %lan+s in pro0i+in% a superficial lu'ricant that pre0ents friction an+
+ryin% of the opposin% con?uncti0al surfaces-
Con7uncti!itis/ or pin,eye/ results from +ama%e to/ an+ irritation of/ the con?uncti0al
surface- The most o'0ious si%n/ re+ness/ is +ue to the +ilation of 'loo+ 0essels +eep to the
con?uncti0al epithelium- This con+ition may 'e cause+ 'y patho%enic infection or 'y
physical/ aller%ic/ or chemical irritation of the con?uncti0al surface-
The 8acrimal !pparatus
! constant flow of tears ,eeps con?uncti0al surfaces moist an+ clean- Tears re+uce friction/
remo0e +e'ris/ pre0ent 'acterial infection/ an+ pro0i+e nutrients an+ o5y%en to portions of
the con?uncti0al epithelium- The lacrimal apparatus pro+uces/ +istri'utes/ an+ remo0es
tears- The lacrimal apparatus of each eye consists of ;1= a lacrimal gland with associate+
+ucts/ ;2= paire+ lacrimal
canaliculi/ ;3= a lacrimal sac/ an+ ;= a nasolacrimal duct ;see 4i%ure 17<3':=-
The poc,et create+ where the palpe'ral con?uncti0a 'ecomes continuous with the ocular
con?uncti0a is ,nown as the forni8 of the eye ;4i%ure 17<a:=- The lateral portion of the
superior forni5 recei0es 10<12 +ucts from the lacrimal gland/ or tear %lan+ ;see 4i%ure 17<
3':=- This %lan+ is a'out the si.e an+ shape of an almon+/ measurin% rou%hly 12<20 mm
;0-5<0-75 in-=- 7t nestles
within a +epression in the frontal 'one/ ?ust insi+e the or'it an+ superior an+ lateral to the
eye'all- lp$ %&5 The lacrimal %lan+ normally pro0i+es the ,ey in%re+ients an+ most of the
0olume of the tears that 'athe the con?uncti0al surfaces- The nutrient an+ o5y%en +eman+s
of the corneal cells are supplie+ 'y +iffusion from the lacrimal secretions/ which are watery
an+ sli%htly al,aline- They contain the anti'acterial en.yme lyso9yme an+ anti'o+ies that
attac, patho%ens 'efore they enter the 'o+y-
The lacrimal %lan+ pro+uces a'out 1 ml of tears each +ay- Once the lacrimal secretions ha0e
reache+ the ocular surface/ they mi5 with the pro+ucts of accessory %lan+s an+ the oily
secretions of the tarsal %lan+s- The result is a superficial Doil slic,E that assists in lu'rication
an+ slows e0aporation-
Blin,in% sweeps the tears across the ocular surface/ an+ they accumulate at the me+ial
canthus in an area ,nown as the lacrimal lae ;lacus lacrimalis=/ or Dla,e of tears-E The
248927081.doc 12
lacrimal la,e co0ers the lacrimal caruncle/ which 'ul%es anteriorly- The lacrimal puncta
;sin%ular/ punctum=/ two small pores/ +rain the lacrimal la,e- They empty into the lacrimal
canaliculi/ small canals that
in turn lea+ to the lacrimal sac ;see 4i%ure 17<3':=/ which nestles within the lacrimal sulcus
of the or'it- lp$ %&6 4rom the inferior portion of the lacrimal sac/ the nasolacrimal duct
passes throu%h the nasolacrimal canal/ forme+ 'y the lacrimal 'one an+ the ma5illary 'one-
The nasolacrimal +uct +eli0ers tears to the nasal ca0ity on that si+e- The +uct empties into
the inferior meatus/ a narrow passa%eway inferior an+ lateral to the inferior nasal concha-
3hen a person cries/ tears rushin% into the nasal ca0ity pro+uce a runny nose/ an+ if the
lacrimal puncta canFt pro0i+e enou%h +raina%e/ the lacrimal la,e o0erflows an+ tears stream
across the face-
,he Eye
The eyes are e5tremely sophisticate+ 0isual instrumentsKmore 0ersatile an+ a+apta'le than
the most e5pensi0e cameras/ yet compact an+ +ura'le- "ach eye is a sli%htly irre%ular
spheroi+ with an a0era%e +iameter of 2 mm ;almost 1 in-/ a little smaller than a Pin%>Pon%
'all= an+ a wei%ht of a'out * % ;0-2* o.=- 3ithin the or'it/ the eye'all shares space with the
e5trinsic eye muscles/ the lacrimal %lan+/ an+ the cranial ner0es an+ 'loo+ 0essels that supply
the eye an+ a+?acent portions of the or'it an+ face- Orbital fat cushions an+ insulates the
eye ;see 4i%ures 17<3' an+ 17<c:=-
The wall of the eye contains three +istinct layers/ or tunics ;4i%ure 17<':=9 ;1= an outer
fibrous tunic/ ;2= an interme+iate vascular tunic/ an+ ;3= an inner neural tunic !retina"- The
0isual receptors/ or photoreceptors/ are locate+ in the neural tunic- The eye'all is hollow2 its
interior can 'e +i0i+e+ into two ca0ities ;4i%ure 17<c:=- The lar%e posterior ca!ity is also
calle+ the vitreous chamber/ 'ecause it contains the %elatinous vitreous body ;vitreo-/
%lassy=- The smaller anterior ca!ity is su'+i0i+e+ into the anterior an+ posterior chambers-
The shape of the eye is sta'ili.e+ in part 'y the 0itreous 'o+y an+ the clear a#ueous humor/
which fills the anterior ca0ity-
The 4i'rous Tunic
248927081.doc 13
The fibrous tunic/ the outermost layer of the eye/ consists of the sclera ;SN8"R>uh= an+
the cornea- The fi'rous tunic ;1= pro0i+es mechanical support an+ some +e%ree of physical
protection/ ;2= ser0es as an attachment site for the e5trinsic eye muscles/ an+
;3= contains structures that assist in the focusin% process-
@ost of the ocular surface is co0ere+ 'y the sclera ;see 4i%ure 17<'/c:=/ or Dwhite of the
eye/E which consists of a +ense fi'rous connecti0e tissue containin% 'oth colla%en an+ elastic
fi'ers- This layer is thic,est o0er the posterior surface of the eye/ near the e5it of the optic
ner0e/ an+ thinnest o0er the anterior surface- The si5 e5trinsic eye muscles insert on the
sclera/ 'len+in% their
colla%en fi'ers with those of the fi'rous tunic- lp$ 55'
The surface of the sclera contains small 'loo+ 0essels an+ ner0es that penetrate the sclera to
reach internal structures- The networ, of small 0essels interior to the ocular con?uncti0a
%enerally +oes not carry enou%h 'loo+ to len+ an o'0ious color to the sclera/ 'ut on close
inspection/ the 0essels are 0isi'le as re+ lines a%ainst the white 'ac,%roun+ of colla%en
fi'ers-
The transparent cornea is structurally continuous with the sclera2 the 'or+er 'etween the
two is calle+ the limbus ;see 4i%ures 17<3a/ 17<a/c:=- Deep to the +elicate corneal
epithelium/ the cornea consists primarily of a +ense matri5 containin% multiple layers of
colla%en fi'ers/ or%ani.e+ so as not to interfere with the passa%e of li%ht- The cornea has no
'loo+ 0essels2 the superficial epithelial cells must o'tain o5y%en an+ nutrients from the tears
that flow across their free surfaces- The cornea also has numerous free ner0e en+in%s/ an+ it
is the most sensiti0e portion of the eye-
1orneal +ama%e may cause 'lin+ness e0en thou%h the functional components of the eyeK
inclu+in% the photoreceptorsK are perfectly normal- The cornea has a 0ery restricte+ a'ility
to repair itself/ so corneal in?uries must 'e treate+ imme+iately to pre0ent serious 0ision
losses- Restorin% 0ision after corneal scarrin% %enerally re&uires the replacement of the
cornea throu%h a corneal transplant- 1orneal replacement is pro'a'ly the most common
form of transplant sur%ery- Such transplants can 'e performe+ 'etween unrelate+ in+i0i+uals/
'ecause there are no 'loo+ 0essels to carry white 'loo+ cells/ which attac, forei%n tissues/
into the area- 1orneal %rafts are o'taine+ from the eyes of +onors who ha0e +ie+ from illness
248927081.doc 14
or acci+ent- 4or 'est results/ the tissues must 'e remo0e+ within fi0e hours after the +onorFs
+eath-
The $ascular Tunic ;H0ea=
Ge
GH
The !ascular tunic/ or u!ea ;>0 >uh=/ contains numerous 'loo+ 0essels/ lymphatic 0essels/
an+ the intrinsic ;smooth= muscles of the eye ;see 4i%ure 17<'/c:=- The functions of this
mi++le layer inclu+e ;1= pro0i+in% a route for 'loo+ 0essels an+ lymphatics that supply
tissues of the eye2 ;2= re%ulatin% the amount of li%ht that enters the eye2 ;3= secretin% an+
rea'sor'in% the a#ueous humor that circulates within the cham'ers of the eye2 an+ ;=
controllin% the shape of the lens/ an essential part of the focusin% process- The 0ascular tunic
inclu+es the iris/ the ciliary body/ an+ the choroid-
The 7ris The iris/ which is 0isi'le throu%h the transparent corneal surface/ contains 'loo+
0essels/ pi%ment cells/ an+ two layers of smooth muscle fi'ers calle+ pupillary muscles
;4i%ure 17<5:=- 3hen these muscles contract/ they chan%e the +iameter of the pupil/ or
central openin% of the iris- One %roup of smooth muscle fi'ers/ the pupillary constrictor
muscles/ forms a series of concentric circles aroun+ the pupil- 3hen these sphincter muscles
contract/ the +iameter of the pupil +ecreases- ! secon+ %roup of smooth muscles/ the
pupillary dilator muscles/ e5ten+s ra+ially away from the e+%e of the pupil- 1ontraction of
these muscles enlar%es the pupil- Both muscle %roups are controlle+ 'y the autonomic
ner0ous system- 4or e5ample/ parasympathetic acti0ation in response to 'ri%ht li%ht causes
the pupils to constrict ;the consensual light reflex=/ an+ sympathetic acti0ation in response
to +im li%ht causes the pupils to +ilate-
The 'o+y of the iris consists of a hi%hly 0ascular/ pi%mente+/ loose connecti0e tissue- The
anterior surface has no epithelial co0erin%2 instea+/ it has an incomplete layer of fi'ro'lasts
an+ melanocytes- @elanocytes are also scattere+ within the 'o+y of the iris- The posterior
surface is co0ere+ 'y a pi%mente+ epithelium that is part of the neural tunic an+ contains
melanin %ranules- "ye color is +etermine+ 'y %enes that influence the +ensity an+
+istri'ution of melanocytes on the anterior surface an+ interior of the iris/ as well as 'y the
+ensity of the pi%mente+ epithelium- 3hen the connecti0e tissue of the iris contains few
248927081.doc 15
melanocytes/ li%ht passes throu%h it an+ 'ounces off the pi%mente+ epithelium- The eye then
appears 'lue- 7n+i0i+uals with %reen/ 'rown/ or 'lac, eyes ha0e increasin% num'ers of
melanocytes in the 'o+y an+ surface of the iris- The eyes of human al'inos appear a 0ery
pale %ray or 'lue>%ray-
The 1iliary Bo+y !t its periphery/ the iris attaches to the anterior portion of the ciliary
body/ a thic,ene+ re%ion that 'e%ins +eep to the ?unction 'etween the cornea an+ the sclera-
The ciliary 'o+y e5ten+s posteriorly to the le0el of the ora serrata ;O>ra
G
ser>R!>tuh2 serrate+ mouth=/ the serrate+ anterior e+%e of the thic,/ inner portion of the
neural tunic ;see 4i%ure 17<a/c:=- The 'ul, of the ciliary 'o+y consists of the ciliary
muscle/ a smooth muscular rin% that pro?ects into the interior of the eye- The epithelium
co0erin% this muscle is thrown into numerous fol+s calle+ ciliary processes- The
suspensory ligaments of the lens attach to the tips of these processes- The connecti0e>
tissue fi'ers of these li%aments hol+ the lens posterior to the iris an+ centere+ on the pupil-
!s a result/ any li%ht passin% throu%h the pupil will also pass throu%h the lens-
The 1horoi+ The choroid is a 0ascular layer that separates the fi'rous an+ neural tunics
posterior to the ora serrata ;see 4i%ure 17<c:=- 1o0ere+ 'y the sclera an+ attache+ to the
outermost layer of the retina/ the choroi+ contains an e5tensi0e capillary networ, that
+eli0ers o5y%en an+ nutrients to the retina- The choroi+ also contains melanocytes/ which
are especially numerous near the sclera-
The Aeural Tunic ;Retina=
The neural tunic/ or retina/ is the innermost layer of the eye- 7t consists of a thin/ outer
layer calle+ the pigmented part/ an+ a thic, inner layer calle+ the neural part- The pi%mente+
part of the retina a'sor's li%ht that passes throu%h the neural part/ pre0entin% li%ht from
'ouncin% 'ac, throu%h the neural part an+ pro+ucin% 0isual Dechoes-E The pi%ment cells also
ha0e important 'iochemical interactions with the retinaFs li%ht receptors/ which are locate+
in the neural part of the retina- 7n a++ition to li%ht receptors/ the neural part of the retina
contains supportin% cells an+ neurons that perform preliminary processin% an+ inte%ration of
0isual information-
248927081.doc 16
The two layers of the retina are normally 0ery close to%ether/ 'ut not ti%htly interconnecte+-
The pi%mente+ part of the retina continues o0er the ciliary 'o+y an+ iris2 the neural part
e5ten+s anteriorly only as far as the ora serrata- The neural part of the retina thus forms a
cup that esta'lishes the posterior an+ lateral 'oun+aries of the posterior ca0ity ;see 4i%ure
17<'/c:=-
Clinical #ote
! retinopathy is a +isease of the retina- +iabetic retinopathy +e0elops in many in+i0i+uals
with diabetes mellitus/ an en+ocrine +isor+er
that interferes primarily with %lucose meta'olism- @any systems are affecte+ 'y +ia'etes/ 'ut
serious car+io0ascular pro'lems are
particularly common- Dia'etic retinopathy/ which +e0elops o0er a perio+ of years/ results
from the +e%eneration/ rupture/ an+ e5ces
si0e %rowth of a'normal 'loo+ 0essels that in0a+e the retina an+ e5ten+ into the space
'etween the pi%ment layer an+ the neural
layer- $isual acuity is %ra+ually lost throu%h +ama%e to photoreceptors ;which are +epri0e+
of o5y%en an+ nutrients=/ lea,a%e of
'loo+ into the posterior cham'er/ an+ the o0er%rowth of 'loo+ 0essels- 8aser therapy can
seal lea,in% 0essels an+ 'loc, new 0essel
%rowth- The posterior cham'er can 'e +raine+ an+ the clou+y flui+ replace+ 'y a suita'ly
clear su'stitute- This proce+ure is calle+ a
vitrectomy- (owe0er/ these are only temporary fi5es that must 'e perio+ically repeate+/
'ecause they fail to correct the un+erlyin%
meta'olic pro'lems-
Or%ani.ation of the Retina 7n sectional 0iew/ the retina is seen to contain se0eral layers of
cells ;4i%ure 17<#a:=- The outermost layer/ closest to the pi%mente+ part of the retina/
contains the photoreceptors/ or cells that +etect li%ht-
The eye has two types of photoreceptors9 ro+s an+ cones- ods +o not +iscriminate amon%
colors of li%ht- (i%hly sensiti0e to li%ht/ they ena'le us to see in +imly lit rooms/ at twili%ht/
an+ in pale moonli%ht- Cones pro0i+e us with color 0ision- Three types of cones are present/
an+ their stimulation in 0arious com'inations pro0i+es the perception of +ifferent colors-
248927081.doc 17
1ones %i0e us sharper/ clearer ima%es than ro+s +o/ 'ut cones re&uire more intense li%ht- 7f
you sit outsi+e at sunset with your te5t'oo, open to a colorful illustration/ you can +etect
the %ra+ual shift in your 0isual system from cone>'ase+ 0ision ;a clear ima%e in full color= to
ro+>'ase+ 0ision ;a relati0ely %rainy ima%e in 'lac, an+ white=-
Ro+s an+ cones are not e0enly +istri'ute+ across the outer surface of the retina-
!ppro5imately 125 million ro+s form a 'roa+ 'an+ aroun+ the periphery of the retina2 as
you mo0e away from the periphery/ towar+ the center of the retina/ the +ensity of ro+s
%ra+ually +ecreases- 7n contrast/ most of the rou%hly # milion cones are concentrate+ in the
area where a 0isual ima%e arri0es after
it passes throu%h the cornea an+ lens- This re%ion/ which is ,nown as the macula lutea
;@!N>
Gu
>luh 8OO>t
Ge
>uh2 yellow spot=/
contains no ro+s- The 0ery hi%hest concentration of cones occurs at the center of the macula
lutea/ an area calle+ the fo!ea ;4
GO
Ge
>uh2 shallow +epression=/ or fovea centralis ;4i%ure 17<#c:=- The fo0ea is the site of sharpest
0ision9 3hen you loo, +irectly at
an o'?ect/ its ima%e falls on this portion of the retina- !n ima%inary line +rawn from the
center of that o'?ect throu%h the center of the lens to the fo0ea esta'lishes the !isual a8is of
the eye ;see 4i%ure 17<c:=-
6ou are pro'a'ly alrea+y aware of the 0isual conse&uences of this +istri'ution of
photoreceptors- 3hen you loo, +irectly at an o'?ect/ you are placin% its ima%e on the fo0ea/
the center of color 0ision- 6ou see a 0ery %oo+ ima%e as lon% as there is enou%h li%ht to
stimulate the cones- But in 0ery +im li%ht/ cones cannot function- That is why you canFt see a
+im star if you stare +irectly at it/ 'ut you can see it if you shift your %a.e to one si+e or the
other- Shiftin% your %a.e mo0es the ima%e of the star from the fo0ea/ where it +oes not
248927081.doc 18
pro0i+e enou%h li%ht to stimulate the cones/ to the periphery/ where it can affect the more
sensiti0e ro+s-
Ro+s an+ cones synapse with rou%hly # million neurons calle+ bipolar cells ;see 4i%ure 17<
#a:=/ which in turn synapse within the layer of neurons calle+ ganglion cells a+?acent to the
posterior ca0ity- ! networ, of hori9ontal cells e5ten+s across the outer portion of the retina
at the le0el of the synapses 'etween photoreceptors an+ 'ipolar cells- ! compara'le layer of
amacrine ;!@>a>,rin= cells occurs where 'ipolar cells synapse with %an%lion cells-
(ori.ontal an+ amacrine cells can facilitate or inhi'it communication 'etween
photoreceptors an+ %an%lion cells/ there'y alterin% the sensiti0ity of the retina- The effect is
compara'le to a+?ustin% the contrast on a tele0ision set- These cells play an important role in
the eyeFs a+?ustment to +im or 'ri%htly lit en0ironments-
The Optic Disc !5ons from an estimate+ 1 million %an%lion cells con0er%e on the optic disc/
a circular re%ion ?ust me+ial to the fo0ea- The optic +isc is the ori%in of the optic ner0e ;77=-
4rom this point/ the a5ons turn/ penetrate the wall of the eye/ an+ procee+ towar+ the
+iencephalon ;4i%ure 17<#':=- The central retinal artery an+ central retinal vein/ which
supply the retina/ pass throu%h the center of the optic ner0e an+ emer%e on the surface of the
optic +isc ;4i%ure 17<#'/c:=- The optic +isc has no photoreceptors or other structures typical
of the rest of the retina- Because li%ht stri,in% this area %oes unnotice+/ the optic +isc is
commonly calle+ the blind spot- 6ou +o not notice a 'lan, spot in your fiel+ of 0ision/
primarily 'ecause in0oluntary eye mo0ements ,eep the 0isual ima%e mo0in% an+ allow your
'rain to fill in the missin% information- (owe0er/ a simple acti0ity usin% 4i%ure 17<7: will
pro0e that a 'lin+ spot really e5ists in your fiel+ of 0ision-
Clinical #ote
Photoreceptors are entirely +epen+ent on the +iffusion of o5y%en an+ nutrients from 'loo+
0essels in the choroi+- 7n a detached retina/ the neural part of the retina 'ecomes separate+
from the pi%mente+ part- This con+ition can result from a 0ariety of factors/ inclu+in% a
su++en 'low to the eye- Hnless the two parts of the neural tunic are reattache+/ the
photoreceptors will +e%enerate an+ 0ision will 'e lost- The reattachment is %enerally
performe+ 'y Dwel+in%E the two layers to%ether usin% laser 'eams focuse+ throu%h the
248927081.doc 19
cornea- These 'eams heat the layers/ there'y fusin% them to%ether at se0eral points aroun+
the retina- (owe0er/ the proce+ure +estroys the photoreceptors an+
other cells at the Dwel+s/E pro+ucin% permanent 'lin+ spots-
The 1ham'ers of the "ye
!s note+ earlier/ the ciliary 'o+y an+ lens +i0i+e the interior of the eye into a lar%e posterior
ca0ity/ or 0itreous cham'er/ an+ a smaller anterior ca0ity ;see 4i%ure 17<c:=- The anterior
ca0ity is su'+i0i+e+ into the anterior chamber/ which e5ten+s from the cornea to the iris/
an+ a posterior chamber/ 'etween the iris an+ the ciliary 'o+y an+ lens- The anterior an+
posterior cham'ers are fille+ with the flui+ a#ueous humor- The posterior ca0ity also
contains a&ueous humor/ 'ut most of its 0olume is ta,en up 'y a %elatinous su'stance
,nown as the vitreous body/ or vitreous humor-
!&ueous (umor Aqueous humor is a flui+ that circulates within the anterior ca0ity/ passin%
from the posterior to the anterior cham'er throu%h the pupil ;4i%ure 17<*:=- 7t also freely
+iffuses throu%h the 0itreous 'o+y an+ across the surface of the retina- !&ueous humor
forms throu%h acti0e secretion 'y epithelial cells of the ciliary 'o+yFs ciliary processes- The
epithelial cells re%ulate its composition/ which resem'les that of cere'rospinal flui+- Because
a&ueous humor circulates/ it pro0i+es an important route for nutrient an+ waste transport/ in
a++ition to formin% a flui+ cushion-
The eye is fille+ with flui+/ an+ flui+ pressure in the a&ueous humor helps retain the eyeFs
shape- 4lui+ pressure also sta'ili.es the position of the retina/ pressin% the neural part
a%ainst the pi%mente+ part- 7n effect/ the a&ueous humor acts li,e the air insi+e a 'alloon-
The eyeFs intraocular pressure can 'e measure+ in the anterior cham'er/ where the flui+
pushes a%ainst the inner surface of the cornea- 7ntraocular pressure is most often chec,e+ 'y
'ouncin% a tiny 'last of air off the surface of the eye an+ measurin% the +eflection pro+uce+-
Aormal intraocular pressure ran%es from 12 to 21 mm (%-
!&ueous humor is secrete+ into the posterior cham'er at a rate of 1<2 ml per minute- 7t
lea0es the anterior cham'er at the same rate- !fter filterin% throu%h a networ, of connecti0e
tissues locate+ near the 'ase of the iris/ a&ueous humor enters the canal of Schlemm/ or
scleral venous sinus/ a passa%eway that e5ten+s completely aroun+ the eye at the le0el of
the lim'us- 1ollectin% channels +eli0er the a&ueous humor from this canal to 0eins in the
248927081.doc 20
sclera- The rate of remo0al normally ,eeps pace with the rate of %eneration at the ciliary
processes/ an+ a&ueous humor is remo0e+ an+ recycle+ within a few hours of its formation-
The $itreous Bo+y The posterior ca0ity of the eye contains the !itreous body/ a %elatinous
mass- The 0itreous 'o+y helps sta'ili.e the shape of the eye/ which mi%ht otherwise +istort
as the e5tra>ocular muscles chan%e its position within the or'it- Speciali.e+ cells em'e++e+
in the 0itreous 'o+y pro+uce the colla%en fi'ers an+ proteo%lycans that account for the
%elatinous consistency of this mass- Hnli,e the a&ueous humor/ the 0itreous 'o+y is forme+
+urin% +e0elopment of the eye an+ is not replace+-
The 8ens
The lens lies posterior to the cornea/ hel+ in place 'y the suspensory li%aments that ori%inate
on the ciliary 'o+y of the choroi+ ;see 4i%ures 17<'/ p- 55#/ an+ 17<*:=- The primary
function of the lens is to focus the 0isual ima%e on the photoreceptors- The lens +oes so 'y
chan%in% its shape-
The lens consists of concentric layers of cells that are precisely or%ani.e+- ! +ense fi'rous
capsule co0ers the entire lens- @any of the capsular fi'ers are elastic- Hnless an outsi+e
force is applie+/ they will contract an+ ma,e the lens spherical- !roun+ the e+%es of the lens/
the capsular fi'ers intermin%le with those of the suspensory li%aments- The cells in the
interior of the lens are calle+ lens fibers- These hi%hly speciali.e+ cells ha0e lost their nuclei
an+ other or%anelles- They are slen+er an+ elon%ate an+ are fille+ with transparent proteins
calle+ crystallins/ which are responsi'le for 'oth the clarity an+ the focusin% power of the
lens- 1rystallins are e5tremely sta'le proteins that remain intact an+ functional for a lifetime
without the nee+ for replacement-
The transparency of the lens +epen+s on a precise com'ination of structural an+ 'iochemical
characteristics- 3hen that 'alance 'ecomes +istur'e+/ the lens loses its transparency2 this
a'normality is ,nown as a cataract- 1ataracts can result from in?uries/ ra+iation/ or reaction
to +ru%s/ 'ut senile cataracts/ a natural conse&uence of a%in%/ are the most common form-
O0er time/ the lens turns yellowish an+ e0entually 'e%ins to lose its transparency- !s the lens
'ecomes Dclou+y/E the in+i0i+ual nee+s 'ri%hter an+ 'ri%hter li%ht for rea+in%/ an+ 0isual
clarity 'e%ins to fa+e- 7f the lens 'ecomes completely opa&ue/ the person will 'e functionally
'lin+/ e0en thou%h the photoreceptors are normal- Sur%ical proce+ures in0ol0e remo0al of
248927081.doc 21
the lens/ either intact or after it has 'een shattere+ with hi%h>fre&uency soun+- The missin%
lens is replace+ 'y an artificial su'stitute/ an+ 0ision is then fine>tune+ with %lasses or
contact lenses-
Refraction The retina has a'out 130 million photoreceptors/ each monitorin% li%ht stri,in% a
specific site on the retina- ! 0isual ima%e results from the processin% of information from all
the receptors- The eye is often compare+ to a camera- To pro0i+e useful information/ the
lens of the eye/ li,e a camera lens/ must focus the arri0in% ima%e- To say that an ima%e is Din
focusE means that the rays of li%ht arri0in% from an o'?ect stri,e the sensiti0e surface of the
retina ;or photo%raphic film= in precise or+er so as to form a miniature ima%e of the o'?ect-
7f the rays are not perfectly focuse+/ the ima%e is 'lurry- 4ocusin% normally occurs in two
steps/ as li%ht passes throu%h first the cornea an+ then the lens-
8i%ht is refracted/ or 'ent/ when it passes from one me+ium to another me+ium with a
+ifferent +ensity- 6ou can +emonstrate this effect 'y stic,in% a pencil into a %lass of water-
Because refraction occurs as the li%ht passes into the air from the much +enser water/ the
shaft of the pencil appears to 'en+ sharply at the air<water interface-
7n the human eye/ the %reatest amount of refraction occurs when li%ht passes from the air
into the corneal tissues/ which ha0e a +ensity close to that of water- 3hen you open your
eyes un+er water/ you cannot see clearly 'ecause refraction at the air<water interface has
'een eliminate+2 li%ht passes un'ent from one watery me+ium to another-
!++itional refraction ta,es place when the li%ht passes from the a&ueous humor into the
relati0ely +ense lens- The lens pro0i+es the e5tra refraction nee+e+ to focus the li%ht rays
from an o'?ect towar+ a focal pointKa specific point of intersection on the retina- The
+istance 'etween the center of the lens an+ its focal point is the focal distance of the lens-
3hether in the eye or in a camera/ the focal +istance is +etermine+ 'y two factors9
&$ $he %istance from the Object to the &ens' The closer an o'?ect is to the lens/ the %reater
the focal +istance ;4i%ure 17<)a/':=-
%$ $he (hape of the &ens' The roun+er the lens/ the more refraction occurs/ so a 0ery roun+
lens has a shorter focal +istance than a flatter one ;4i%ure 17'/c:=-
!ccommo+ation ! camera focuses an ima%e 'y mo0in% the lens towar+ or away from the
film- This metho+ of focusin% cannot wor, in our eyes/ 'ecause the +istance from the lens to
248927081.doc 22
the macula lutea cannot chan%e- 3e focus ima%es on the retina 'y chan%in% the shape of the
lens to ,eep the focal len%th constant/ a process calle+ accommodation ;4i%ure 17<10:=-
Durin% accommo+ation/ the lens 'ecomes roun+er to focus the ima%e of a near'y o'?ect on
the retina2 the lens flattens when we focus on a +istant o'?ect-
The lens is hel+ in place 'y the suspensory li%aments that ori%inate at the ciliary 'o+y-
Smooth muscle fi'ers in the ciliary 'o+y act li,e sphincter muscles- 3hen the ciliary muscle
contracts/ the ciliary 'o+y mo0es towar+ the lens/ there'y re+ucin% the tension in the
suspensory li%aments- The elastic capsule then pulls the lens into a more spherical shape that
increases the refracti0e power of the lens/ ena'lin% it to 'rin% li%ht from near'y o'?ects into
focus on the retina ;4i%ure 17<10a:=- 3hen the ciliary muscle rela5es/ the suspensory
li%aments pull at the circumference of the lens/ ma,in% the lens flatter ;4i%ure 17<10':=-
The %reatest amount of refraction is re&uire+ to 0iew o'?ects that are 0ery close to the lens-
The inner limit of clear 0ision/ ,nown as the near point of vision/ is +etermine+ 'y the
+e%ree of elasticity in the lens- 1hil+ren can usually focus on somethin% 7<) cm from the
eye/ 'ut o0er time the lens ten+s to 'ecome stiffer an+ less responsi0e- ! youn% a+ult can
usually focus on o'?ects 15<20 cm away- !s a%in% procee+s/ this +istance %ra+ually
increases2 the near point at a%e #0 is typically a'out *3 cm- ;4or more information on
con%enital an+ a%e>relate+ chan%es in eye structure an+ function/ see the 1linical Aote
D!ccommo+ation Pro'lems-E=
7f li%ht passin% throu%h the cornea an+ lens is not refracte+ properly/ the 0isual ima%e will 'e
+istorte+- 7n the con+ition calle+ astigmatism/ the +e%ree of cur0ature in the cornea or lens
0aries from one a5is to another- @inor asti%matism is 0ery common2 the ima%e +istortion
may 'e so minimal that people are unaware of the con+ition-
7ma%e Re0ersal Thus far/ we ha0e consi+ere+ li%ht that ori%inates at a sin%le point/ either
near or far from the 0iewer- !n o'?ect in 0iew/ howe0er/ is a comple5 li%ht source that must
'e treate+ as a lar%e num'er of in+i0i+ual points- 8i%ht from each point is focuse+ on the
retina as in+icate+ in 4i%ure 17<12a/':- The result is the creation of a miniature ima%e of the
ori%inal/ 'ut the ima%e arri0es upsi+e +own an+ 'ac,war+-
To un+erstan+ why the ima%e is re0erse+ in this fashion/ consi+er 4i%ure 17<12c:/ a sa%ittal
section throu%h an eye that is loo,in% at a telephone pole- The ima%e of the top of the pole
lan+s at the 'ottom of the retina/ an+ the ima%e of the 'ottom hits the top of the retina- Aow
248927081.doc 23
consi+er 4i%ure 17<12+:/ a hori.ontal section throu%h an eye that is loo,in% at a pic,et
fence- The ima%e of the left e+%e of the fence falls on the ri%ht si+e of the retina/ an+ the
ima%e of the ri%ht e+%e falls on the left si+e of the retina- The 'rain compensates for this
ima%e re0ersal/ an+ we are not aware of any +ifference 'etween the orientation of the ima%e
on the retina an+ that of the o'?ect-
$isual !cuity 1larity of 0ision/ or !isual acuity/ is rate+ a%ainst a DnormalE stan+ar+- The
stan+ar+ 0ision ratin% of 20P 20 is +efine+ as the le0el of +etail seen at a +istance of 20 feet
'y an in+i0i+ual with normal 0ision- $ision rate+ as 20P 15 is 'etter than a0era%e/ 'ecause at
20 feet the person is a'le to see +etails that woul+ 'e clear to a normal eye only at a +istance
of 15 feet- 1on0ersely/ a person with 20P 30 0ision must 'e 20 feet from an o'?ect to +iscern
+etails that a person with normal 0ision coul+ ma,e
out at a +istance of 30 feet-
3hen 0isual acuity falls 'elow 20P 200/ e0en with the help of %lasses or contact lenses/ the
in+i0i+ual is consi+ere+ to 'e le%ally 'lin+- There are pro'a'ly fewer than 00/000 le%ally
'lin+ people in the Hnite+ States2 more than half are o0er #5 years ol+- The term blindness
implies a total a'sence of 0ision +ue to +ama%e to the eyes or to the optic pathways-
1ommon causes of 'lin+ness inclu+e +ia'etes mellitus/ cataracts/ %laucoma/ corneal
scarrin%/ +etachment of the retina/ acci+ental in?uries/ an+ here+itary factors that are as yet
poorly un+erstoo+-
!'normal 'lin+ spots/ or scotomas ;s,
Go
>T
GO
>muh.=/ may appear in the fiel+ of 0ision at positions other than at the optic
+isc- Scotomas are permanent a'normalities that are fi5e+ in position- They may result from
a compression of the optic ner0e/ +ama%e to photoreceptors/ or central +ama%e alon% the
0isual pathway- )loaters/ small spots that +rift across the fiel+ of 0ision/ are %enerally
temporary phenomena that result from 'loo+ cells or cellular +e'ris in the 0itreous 'o+y-
They can 'e +etecte+ 'y starin% at a 'lan, wall or a white sheet of paper-
248927081.doc 24
&11 Keys 2 8i%ht passes throu%h the con?uncti0a an+ cornea/ crosses the anterior ca0ity to
reach the lens/ transits the lens/ crosses the posterior cham'er/ an+ then penetrates the neural
tissue of the retina 'efore reachin% an+ stimulatin% the photoreceptors- 1ones are most
a'un+ant at the fo0ea an+ macula lutea/ an+ they pro0i+e hi%h>resolution color 0ision in
'ri%htly lit en0ironments- Ro+s +ominate the peripheral areas of the retina/ an+ they pro0i+e
relati0ely low>resolution 'lac,>an+>white 0ision in +imly lit en0ironments-
Concept Chec3
3hich layer of the eye woul+ 'e affecte+ first 'y the ina+e&uate pro+uction of tearsM
3hen the lens of your eye is 0ery roun+/ are you loo,in% at an o'?ect that is close to you
or far from youM
!s Renee enters a +ar, room/ most of the a0aila'le li%ht 'ecomes focuse+ on the fo0ea
of her eye- 3ill she 'e a'le to see 0ery clearlyM (ow woul+ a 'loc,a%e of the canal of
Schlemm affect your 0isionM
Ans"ers begin on p$ A4&
Visual )hysiology
The ro+s an+ cones of the retina are calle+ photoreceptors 'ecause they +etect photons/
'asic units of 0isi'le li%ht- 8i%ht ener%y is a form of radiant energy that tra0els in wa0es with
a characteristic wavelength ;+istance 'etween wa0e pea,s=-
Our eyes are sensiti0e to wa0elen%ths of 700<00 nm/ the spectrum of 0isi'le li%ht- This
spectrum/ seen in a rain'ow/ can 'e remem'ere+ 'y the acronym RO6 G- B7$ ;*e+/
Oran%e/ +ellow/ ,reen/ Blue/ -n+i%o/ .iolet=- Photons of re+ li%ht carry the least ener%y an+
ha0e the lon%est wa0elen%th/ an+ those from the 0iolet portion of the spectrum contain the
most ener%y an+ ha0e the shortest wa0elen%th- Ro+s pro0i+e the central ner0ous system
with information a'out the presence or a'sence of photons/ without re%ar+ to their
wa0elen%th- 1ones pro0i+e information a'out the wa0elen%th of arri0in% photons/ %i0in% us
a perception of color-
!natomy of Ro+s an+ 1ones
248927081.doc 25
4i%ure 17<13a: compares the structures of ro+s an+ cones- The elon%ate+ outer segment of
a photoreceptor contains hun+re+s to thousan+s of flattene+ mem'ranous plates/ or discs-
The names rod an+ cone refer to the outer se%mentFs shape- 7n a ro+/ each +isc is an
in+epen+ent entity/ an+ the outer se%ment forms an elon%ate+ cylin+er- 7n a cone/ the +iscs
are infol+in%s of the cell mem'rane/ an+ the outer se%ment tapers to a 'lunt point-
! narrow connectin% stal, attaches the outer se%ment to the inner segment/ a re%ion that
contains all the usual cellular or%anelles- The inner se%ment ma,es synaptic contact with
other cells/ an+ it is here that neurotransmitters are release+-
$isual Pi%ments The +iscs of the outer se%ment in 'oth ro+s an+ cones contain special
or%anic compoun+s calle+ !isual pigments- The a'sorption of photons 'y 0isual pi%ments is
the first ,ey step in the process of photoreceptionKthe +etection of li%ht- $isual pi%ments
are +eri0ati0es of the compoun+ rhodopsin ;roG>DOP>sin=/ or visual purple/ the 0isual
pi%ment foun+ in ro+s ;4i%ure 17<13':=- Rho+opsin consists of a protein/ opsin/ 'oun+ to
the pi%ment retinal ;R"T>i>nal=/ or retinene/ which is synthesi.e+ from !itamin A- One
form of opsin is characteristic of all ro+s-
1ones contain the same retinal pi%ment that ro+s +o/ 'ut in cones retinal is attache+ to other
forms of opsin- The type of opsin present +etermines the wa0elen%th of li%ht that can 'e
a'sor'e+ 'y retinal- Differential stimulation of these cone populations is the 'asis of color
0ision-
Aew +iscs containin% 0isual pi%ment are continuously assem'le+ at the 'ase of the outer
se%ment- ! complete+ +isc then mo0es towar+ the tip of the se%ment- !fter a'out 10 +ays/
the +isc will 'e she+ in a small +roplet of cytoplasm- Droplets with she+ +iscs are a'sor'e+
'y the pi%ment cells/ which 'rea, +own the mem'raneFs components an+ recon0ert the
retinal to 0itamin !- The 0itamin ! is then store+ within the pi%ment cells for su'se&uent
transfer to the photoreceptors-
The term retinitis pigmentosa ;RP= refers to a collection of inherite+ retinopathies-
To%ether/ they are the most common inherite+ 0isual a'normality/ affectin% appro5imately 1
in+i0i+ual in 3000- The 0isual receptors %ra+ually +eteriorate/ an+ 'lin+ness e0entually
results- The mutations that are responsi'le chan%e the structure of the photoreceptorsK
specifically/ the 0isual pi%ments of the mem'rane +iscs- 7t is not ,nown how the altere+
pi%ments lea+ to the +estruction of photoreceptors-
248927081.doc 26
Photoreception
The cell mem'rane in the outer se%ment of the photoreceptor contains chemically re%ulate+
so+ium ion channels- ;Refer to the +ia%ram of the restin% state in 4i%ure 17<1:-= 7n
+ar,ness/ these %ate+ channels are ,ept open in the presence of cyclic-,M/ ;cyclic
guanosine monophosphate/ or c,M/=/ a +eri0ati0e of the hi%h>ener%y compoun+ guanosine
triphosphate ;GTP=- Because the channels are open/ the transmem'rane potential is
appro5imately
>0 m$/ rather than the >70 m$ typical of restin% neurons- !t the >0>m$ transmem'rane
potential/ the photoreceptor is continuously releasin% neurotransmitters ;in this case/
%lutamate= across synapses at the inner se%ment- The inner se%ment also continuously pumps
so+ium ions out of the cytoplasm- The mo0ement of so+ium ions into the outer se%ment/ on
to the inner se%ment/ an+ out of the cell is ,nown as the dar current-
The process of rho+opsin>'ase+ photoreception 'e%ins when a photon stri,es the retinal
portion of a rho+opsin molecule em'e++e+ in the mem'rane of the +isc ;see 4i%ure 17<1:=9
(tep 0 Opsin 7s !cti0ate+- The 'oun+ retinal molecule has two possi'le confi%urations9 the
&&-cis form an+ the &&-trans form- Aormally/ the molecule is in the 11>cis form2 on
a'sor'in% li%ht/ it chan%es to the more linear 11>trans form- This chan%e acti0ates the opsin
molecule-
(tep 1 Opsin !cti0ates Trans+ucin/ 3hich in Turn !cti0ates Phospho+iesterase- ,ransducin
is a G proteinKa mem'rane>'oun+ en.yme comple5- lp$ (&& 7n this case/ trans+ucin is
acti0ate+ 'y opsin/ an+ trans+ucin in turn acti0ates phosphodiesterase :)+E;-
(tep 2 1yclic>G@P ;cG@P= 8e0els Decline/ an+ Gate+ So+ium 1hannels 1lose-
Phospho+iesterase is an en.yme that 'rea,s +own cG@P- The remo0al of cG@P from the
%ate+ so+ium channels results in their inacti0ation- The rate of Aa
I
entry into the cytoplasm
then +ecreases-
(tep 3 The Dar, 1urrent 7s Re+uce+ an+ the Rate of Aeurotransmitter Release Declines-
The re+uction in the rate of Aa
I
entry re+uces the +ar, current- Because acti0e transport
continues to remo0e Aa
I
from the cytoplasm/ when the so+ium channels close/ the
transmem'rane potential +rops towar+ >70 m$- !s the mem'rane hyperpolari.es/ the rate of
248927081.doc 27
neurotransmitter release +ecreases/ in+icatin% to the a+?acent 'ipolar cell that the
photoreceptor has a'sor'e+ a photon-
Reco0ery !fter Stimulation !fter a'sor'in% a photon/ retinal +oes not spontaneously re0ert
to the 11>cis form- 7nstea+/ the entire rho+opsin molecule must 'e 'ro,en +own an+
reassem'le+- Shortly after the chan%e in shape occurs/ the rho+opsin molecule 'e%ins to
'rea, +own into retinal an+ opsin/ a process ,nown as bleaching ;4i%ure 17<15:=- Before it
can recom'ine with opsin/ the retinal must 'e en.ymatically con0erte+ to the 11>cis form-
This con0ersion re&uires ener%y in the form of !TP ;a+enosine triphosphate=/ an+ it ta,es
time-
Bleachin% contri'utes to the lin%erin% 0isual impression you ha0e after you see a flash'ul'
%o off- 4ollowin% intense e5posure to li%ht/ a photoreceptor cannot respon+ to further
stimulation until its rho+opsin molecules ha0e 'een re%enerate+- !s a result/ a D%hostE ima%e
remains on the retina- Bleachin% is sel+om noticea'le un+er or+inary circumstances/ 'ecause
the eyes are constantly ma,in% small/ in0oluntary chan%es in position that mo0e the ima%e
across the retinaFs surface-
3hile the rho+opsin molecule is 'ein% reassem'le+/ mem'rane permea'ility is returnin% to
normal- Opsin is inacti0ate+ when 'leachin% occurs/ an+ the 'rea,+own of cG@P halts as a
result- !s other en.ymes %enerate cG@P in the cytoplasm/ the chemically %ate+ so+ium
channels reopen-
!s pre0iously note+/ the 0isual pi%ments of the photoreceptors are synthesi.e+ from 0itamin
!- The 'o+y contains 0itamin ! reser0es sufficient for se0eral months/ an+ a si%nificant
amount is store+ in the cells of the pi%mente+ part of the retina- 7f +ietary sources are
ina+e&uate/ these reser0es are %ra+ually e5hauste+ an+ the amount of 0isual pi%ment in the
photoreceptors 'e%ins to +ecline- Dayli%ht 0ision is affecte+/ 'ut in +aytime the li%ht is
usually 'ri%ht enou%h to stimulate any 0isual pi%ments that remain within the +ensely pac,e+
cone population of the fo0ea- !s a result/ the pro'lem first 'ecomes apparent at ni%ht/ when
the +im li%ht pro0es insufficient to acti0ate the ro+s- This con+ition/ ,nown as night
blindness/ can 'e treate+ 'y the a+ministration of 0itamin !- The 'o+y can con0ert the
carotene pi%ments in many 0e%eta'les to 0itamin !- 1arrots are a particularly %oo+ source of
caroteneKhence the ol+ a+a%e that carrots are %oo+ for your eyes-
248927081.doc 28
1olor $ision
!n or+inary li%ht'ul' or the sun emits photons of all wa0elen%ths- These photons stimulate
'oth ro+s an+ cones- 3hen all three types of cones are stimulate+/ or when ro+s alone are
stimulate+/ you see a DwhiteE li%ht- 6our eyes also +etect photons that reach your retina
after they 'ounce off o'?ects aroun+ you- 7f photons of all colors 'ounce off an o'?ect/ the
o'?ect will appear white to you2 if all the photons are a'sor'e+ 'y the o'?ect ;so that none
reach the retina=/ the o'?ect will appear 'lac,- !n o'?ect will appear to ha0e a particular
color if it reflects ;or transmits= photons from one portion of the 0isi'le spectrum an+
a'sor's the rest-
The three types of cones are blue cones/ green cones/ an+ red cones- "ach type has a
+ifferent form of opsin an+ a sensiti0ity to a +ifferent ran%e of wa0elen%ths- Their stimulation
in 0arious com'inations is the 'asis for color 0ision- 7n an in+i0i+ual with normal 0ision/ the
cone population consists of 1# percent 'lue cones/ 10 percent %reen cones/ an+ 7 percent
re+ cones- !lthou%h their sensiti0ities o0erlap/ each type is most sensiti0e to a specific
portion of the 0isual spectrum ;4i%ure 17<1#:=-
1olor +iscrimination occurs throu%h the inte%ration of information arri0in% from all three
types of cones- 4or e5ample/ the perception of yellow results from a com'ination of inputs
from hi%hly stimulate+ %reen cones/ less stron%ly stimulate+ re+ cones/ an+ relati0ely
unaffecte+ 'lue cones ;see 4i%ure 17<1#:=- 7f all three cone populations are stimulate+/ we
percei0e the color as white- Because we also percei0e white if ro+s/ rather than cones/ are
stimulate+/ e0erythin% appears 'lac,>an+>white when we enter +imly lit surroun+in%s or
wal, 'y starli%ht-
Persons who are una'le to +istin%uish certain colors ha0e a form of color blindness- The
stan+ar+ tests for color 0ision in0ol0e pic,in% num'ers or letters out of a comple5 colore+
picture ;4i%ure 17<17:=- 1olor 'lin+ness occurs when one or more classes of cones are
nonfunctional- The cones may 'e a'sent/ or they may 'e present 'ut una'le to manufacture
the necessary 0isual pi%ments- 7n the most common type of color 'lin+ness ;re+<%reen color
'lin+ness=/ the re+ cones are missin%/ so the in+i0i+ual cannot +istin%uish re+ li%ht from
%reen li%ht- 7nherite+ color 'lin+ness in0ol0in% one or two cone pi%ments is not unusual- Ten
percent of all men show some color 'lin+ness/ whereas the inci+ence amon% women is only
a'out 0-#7 percent- Total color 'lin+ness is e5tremely rare2 only 1 person in 300/000 fails to
248927081.doc 29
manufacture any cone pi%ments- 3e will consi+er the inheritance of color 'lin+ness in
1hapter 2)-
8i%ht an+ Dar, !+aptation
The sensiti0ity of your 0isual system 0aries with the intensity of illumination- !fter 30
minutes or more in the +ar,/ almost all 0isual pi%ments will 'e fully recepti0e to stimulation-
This is the dar3-adapted state- 3hen +ar,>a+apte+/ the 0isual system is e5tremely
sensiti0e- 4or e5ample/ a sin%le ro+ will hyperpolari.e in response to a sin%le photon of li%ht-
"0en more remar,a'le/ if as few as se0en ro+s a'sor' photons at one time/ you will see a
flash of li%ht-
3hen the li%hts come on/ at first they seem almost un'eara'ly 'ri%ht/ 'ut o0er the ne5t few
minutes your sensiti0ity +ecreases as 'leachin% occurs- "0entually/ the rate of 'leachin% is
'alance+ 'y the rate at which they re>form- This con+ition is the light-adapted state- 7f you
mo0e+ from the +epths of a ca0e to the full sunli%ht of mi++ay/ your receptor sensiti0ity
woul+ +ecrease 'y a factor of 25/000-
! 0ariety of central responses further a+?ust li%ht sensiti0ity- 1onstriction of the pupil/ 0ia the
pupillary constrictor reflex/ re+uces the amount of li%ht enterin% your eye to one>thirtieth
the ma5imum +ar,>a+apte+ le0el- Dilatin% the pupil fully can pro+uce a thirtyfol+ increase in
the amount of li%ht enterin% the eye/ an+ facilitatin% some of the synapses alon% the 0isual
pathway can perhaps triple its sensiti0ity- (ence/ the efficiency of the entire system may
increase 'y a factor of more than 1 million-
,he Visual )ath"ay
The 0isual pathway 'e%ins at the photoreceptors an+ en+s at the 0isual corte5 of the cere'ral
hemispheres- 7n other sensory pathways we ha0e e5amine+/ at most one synapse lies
'etween a receptor an+ a sensory neuron that +eli0ers information to the 1AS- 7n the 0isual
pathway/ the messa%e must cross two synapses ;photoreceptor to 'ipolar cell/ an+ 'ipolar
cell to %an%lion cell= 'efore it hea+s towar+ the 'rain- The e5tra synapse increases the
synaptic +elay/ 'ut it pro0i+es an opportunity for the processin% an+ inte%ration of 0isual
information 'efore it lea0es the retina-
248927081.doc 30
Processin% 'y the Retina
"ach photoreceptor in the retina monitors a specific recepti0e fiel+- The retina contains
a'out 130 million photoreceptors/ # million 'ipolar cells/ an+ 1 million %an%lion cells- Thus/
a consi+era'le amount of con0er%ence occurs at the start of the 0isual pathway- The +e%ree
of con0er%ence +iffers 'etween ro+s an+ cones- Re%ar+less of the amount of con0er%ence/
each %an%lion cell monitors a specific portion of the fiel+ of 0ision-
!s many as a thousan+ ro+s may pass information 0ia their 'ipolar cells to a sin%le %an%lion
cell- The %an%lion cells that monitor ro+s/ calle+ < cells ;magnocells2 magnus/ %reat=/ are
relati0ely lar%e- They pro0i+e information a'out the %eneral form of an o'?ect/ motion/ an+
sha+ows in +im li%htin%- Because so much con0er%ence occurs/ the acti0ation of an @ cell
in+icates that li%ht has arri0e+ in a %eneral area rather than at a specific location-
The loss of specificity +ue to con0er%ence is partially o0ercome 'y the fact that the acti0ity
of %an%lion cells 0aries accor+in% to the pattern of acti0ity in their recepti0e fiel+/ which is
%enerally circular- Typically/ a %an%lion cell respon+s +ifferently to stimuli that arri0e in the
center of its recepti0e fiel+ than to stimuli that arri0e at the e+%es ;4i%ure 17<1*:=- Some
%an%lion cells ;oncenter neurons= are e5cite+ 'y li%ht arri0in% in the center of their sensory
fiel+ an+ are inhi'ite+ when li%ht stri,es the e+%es of their recepti0e fiel+- Others ;off-center
neurons= are inhi'ite+ 'y li%ht in the central .one/ 'ut are stimulate+ 'y illumination at the
e+%es- On>center an+ off>center neurons pro0i+e information a'out which portion of their
recepti0e fiel+ is illuminate+-
1ones typically show 0ery little con0er%ence2 in the fo0ea/ the ratio of cones to %an%lion
cells is 191- The %an%lion cells that monitor cones/ calle+ ) cells ;parvo cells2 parvus/ small=/
are smaller an+ more numerous than @ cells- P cells are acti0e in 'ri%ht li%ht/ an+ they
pro0i+e information a'out e+%es/ fine +etail/ an+ color- Because little con0er%ence occurs/
the acti0ation of a P cell means that li%ht has arri0e+ at one specific location- !s a result/
cones pro0i+e more precise information a'out a 0isual ima%e than +o ro+s- 7n 0i+eo%raphic
terms/ ima%es forme+ 'y ro+s ha0e a coarse/ %rainy/ pi5elate+ appearance that 'lurs +etails2
'y contrast/ ima%es pro+uce+ 'y cones are fine>%raine+/ of hi%h +ensity/ sharp/ an+ clear-
1entral Processin% of $isual 7nformation
248927081.doc 31
!5ons from the entire population of %an%lion cells con0er%e on the optic +isc/ penetrate the
wall of the eye/ an+ procee+ towar+ the +iencephalon as the optic ner0e ;77=- The two optic
ner0es/ one from each eye/ reach the +iencephalon at the optic chiasm ;4i%ure 17<1):=-
4rom that point/ appro5imately half the fi'ers procee+ towar+ the lateral %eniculate nucleus
of the same si+e of
the 'rain/ whereas the other half cross o0er to reach the lateral %eniculate nucleus of the
opposite si+e- lp$ ('' 4rom each lateral %eniculate nucleus/ 0isual information tra0els to the
occipital corte5 of the cere'ral hemisphere on that si+e- The 'un+le of pro?ection fi'ers
lin,in% the lateral %eniculates with the 0isual corte5 is ,nown as the optic radiation-
1ollaterals from the fi'ers synapsin% in the lateral %eniculate continue to su'conscious
processin% centers in the +iencephalon an+ 'rain stem- 4or e5ample/ the pupillary refle5es
an+ refle5es that control eye mo0ement are tri%%ere+ 'y collaterals carryin% information to
the superior colliculi-
The 4iel+ of $ision The perception of a 0isual ima%e reflects the inte%ration of information
that arri0es at the 0isual corte5 of the occipital lo'es- "ach eye recei0es a sli%htly +ifferent
0isual ima%e/ 'ecause ;1= the fo0eas are 5<7-5 cm apart/ an+ ;2= the nose an+ eye soc,et
'loc, the 0iew of the opposite si+e- +epth perception/ an interpretation of the three>
+imensional relationships amon% o'?ects in 0iew/ is o'taine+ 'y comparin% the relati0e
positions of o'?ects within the ima%es recei0e+ 'y the two eyes-
3hen you loo, strai%ht ahea+/ the 0isual ima%es from your left an+ ri%ht eyes o0erlap ;see
4i%ure 17<1):=- The ima%e recei0e+ 'y the fo0ea of each eye lies in the center of the re%ion
of o0erlap- ! 0ertical line +rawn throu%h the center of this re%ion mar,s the +i0ision of
0isual information at the optic chiasm- $isual information from the left half of the com'ine+
fiel+ of 0ision reaches the 0isual corte5 of your ri%ht occipital lo'e2 0isual information from
the ri%ht half of the com'ine+ fiel+ of 0ision arri0es at the 0isual corte5 of your left occipital
lo'e-
The cere'ral hemispheres thus contain a map of the entire fiel+ of 0ision- !s in the case of
the primary sensory corte5/ the map +oes not faithfully +uplicate the relati0e areas within the
sensory fiel+- 4or e5ample/ the area assi%ne+ to the macula lutea an+ fo0ea co0ers a'out 35
times the surface it woul+ co0er if the map were proportionally accurate- The map is also
upsi+e +own an+ 'ac,war+/ +uplicatin% the orientation of the 0isual ima%e at the retina-
248927081.doc 32
The Brain Stem an+ $isual Processin% @any centers in the 'rain stem recei0e 0isual
information/ either from the lateral %eniculate nuclei or throu%h collaterals from the optic
tracts- 1ollaterals that 'ypass the lateral %eniculates synapse in the superior colliculi or in the
hypothalamus- The superior colliculi of the mesencephalon issue motor comman+s that
control unconscious eye/ hea+/ or nec, mo0ements in response to 0isual stimuli- $isual
inputs to the suprachiasmatic nucleus of the hypothalamus affect
the function of other 'rain stem nuclei- lp$ ('= The suprachiasmatic nucleus an+ the pineal
gland of the epithalamus recei0e 0isual information an+ use it to esta'lish a +aily pattern of
0isceral acti0ity that is tie+ to the +ay<ni%ht cycle- This circadian rhythm ;circa/ a'out I
dies/ +ay= affects your meta'olic rate/ en+ocrine function/ 'loo+ pressure/ +i%esti0e
acti0ities/ awa,e<asleep cycle/ an+ other physiolo%ical an+ 'eha0ioral processes-
Anatomy 5'1 2 Re0iew the anatomy of the eye on the Anatomy 5'1 C+-O<> #er!ous
System?Special Senses?Eye$
Concept Chec3
7f you ha+ 'een 'orn without cones in your eyes/ woul+ you still 'e a'le to seeM "5plain-
(ow coul+ a +iet +eficient in 0itamin ! affect 0isionM
3hat effect woul+ a +ecrease in phospho+iesterase acti0ity in photoreceptor cells ha0e
on 0isionM
Ans"ers begin on p$ A4&
Equilibrium and Hearing
Objectives
: Descri'e the structures of the e5ternal an+ mi++le ears an+ e5plain how they function-
: Descri'e the parts of the inner ear an+ their roles in e&uili'rium an+ hearin%-
: Trace the pathways for the sensations of e&uili'rium an+ hearin% to their respecti0e
+estinations in the 'rain-
248927081.doc 33
The special senses of e&uili'rium an+ hearin% are pro0i+e+ 'y the inner ear/ a receptor
comple5 locate+ in the petrous part of the temporal 'one of the s,ull- 4#uilibrium
sensations inform us of the position of the hea+ in space 'y monitorin% %ra0ity/ linear
acceleration/ an+ rotation- 5earing ena'les us to +etect an+ interpret soun+ wa0es- The
'asic receptor mechanism for 'oth senses is the same- The receptors/ calle+ hair cells/ are
simple mechanoreceptors- The comple5 structure of the inner ear an+ the +ifferent
arran%ement of accessory structures ena'le hair cells to respon+ to +ifferent stimuli an+ thus
to pro0i+e the input for 'oth senses-
Anatomy of the Ear
The ear is +i0i+e+ into three anatomical re%ions9 the e5ternal ear/ the mi++le ear/ an+ the
inner ear ;4i%ure 17<20:=- The external earKthe 0isi'le portion of the earKcollects an+
+irects soun+ wa0es towar+ the middle ear/ a cham'er locate+ within the petrous portion of
the temporal 'one- Structures of the mi++le ear collect soun+ wa0es an+ transmit them to an
appropriate portion of the inner ear/ which contains the sensory or%ans for hearin% an+
e&uili'rium-
The "5ternal "ar
The e8ternal ear inclu+es the fleshy an+ cartila%inous auricle/ or pinna/ which surroun+s
the e8ternal acoustic canal/ or ear canal- The auricle protects the openin% of the canal an+
pro0i+es +irectional sensiti0ity2 soun+s comin% from 'ehin+ the hea+ are 'loc,e+ 'y the
auricle/ whereas soun+s comin% from the si+e or front are collecte+ an+ channele+ into the
e5ternal acoustic canal- ;3hen you DcupE your ear with your han+ to hear a faint soun+
more clearly/ you are e5a%%eratin% this effect-= The e5ternal acoustic canal is a passa%eway
that en+s at the tympanic membrane/ also calle+ the tympanum or eardrum- The tympanic
mem'rane is a thin/ semitransparent sheet that separates the e5ternal ear from the mi++le ear-
The tympanic mem'rane is 0ery +elicate- The auricle an+ the narrow e5ternal acoustic canal
pro0i+e some protection from acci+ental in?ury- 7n a++ition/ ceruminous glandsK
inte%umentary %lan+s alon% the e5ternal acoustic canalKsecrete a wa5y material that helps
+eny access to forei%n o'?ects or small insects/ as +o many small/ outwar+ly pro?ectin% hairs-
These hairs also pro0i+e increase+ tactile sensiti0ity throu%h their root hair ple5uses- The
248927081.doc 34
sli%htly wa5y secretion of the ceruminous %lan+s/ calle+ cerumen/ also slows the %rowth of
microor%anisms in the e5ternal acoustic canal an+ re+uces the chances of infection-
The @i++le "ar
The middle ear/ or tympanic ca!ity/ is an air>fille+ cham'er separate+ from the e5ternal
acoustic canal 'y the tympanic mem'rane- The mi++le ear communicates 'oth with the
nasopharynx ;the superior portion of the pharyn5=/ throu%h the auditory tube/ an+ with the
mastoi+ air cells/ throu%h a num'er of small connections ;4i%ures 17<20 an+ 17<21:=- The
au+itory tu'e is also calle+ the pharyngotympanic tube or the 4ustachian tube- !'out cm
;1-# in-= lon%/ it consists of two portions- The portion near the connection to the mi++le ear
is relati0ely narrow an+ is supporte+ 'y elastic cartila%e- The portion near the openin% into
the nasopharyn5 is relati0ely 'roa+ an+ funnel shape+- The au+itory tu'e permits the
e&uali.ation of pressures on either si+e of the tympanic mem'rane- Hnfortunately/ the
au+itory tu'e can also allow microor%anisms to tra0el from the nasopharyn5 into the mi++le
ear- 7n0asion 'y microor%anisms can lea+ to an unpleasant mi++le ear infection ,nown as
otitis media- !@9 Otitis @e+ia an+ @astoi+itis
The !u+itory Ossicles The mi++le ear contains three tiny ear 'ones/ collecti0ely calle+
auditory ossicles- The ear 'ones connect the tympanic mem'rane with one of the receptor
comple5es of the inner ear ;see 4i%ures 17<20 an+ 17<21:=- The three au+itory ossicles are
the malleus/ the incus/ an+ the stapes- The malleus ;malleus/ hammer= attaches at three
points to the interior surface of the tympanic mem'rane- The incus ;incus/ an0il= the mi++le
ossicle/ attaches the malleus to the stapes ;stapes/ stirrup=/ the inner ossicle- The e+%es of
the 'ase of the stapes are 'oun+ to the e+%es of the oval window/ an openin% in the 'one
that surroun+s the inner ear- The articulations 'etween the au+itory ossicles are the smallest
syno0ial ?oints in the 'o+y- "ach has a tiny capsule an+ supportin% e5tracapsular li%aments-
$i'ration of the tympanic mem'rane con0erts arri0in% soun+ wa0es into mechanical
mo0ements- The au+itory ossicles act as le0ers that con+uct those 0i'rations to the inner ear-
The ossicles are connecte+ in such a way that an in<out mo0ement of the tympanic
mem'rane pro+uces a roc,in% motion of the stapes- The ossicles thus function as a le0er
system that collects the force applie+ to the tympanic mem'rane an+ focuses it on the o0al
win+ow- Because the tympanic mem'rane is 22 times lar%er an+ hea0ier than the o0al
248927081.doc 35
win+ow/ consi+era'le amplification occurs/ so we can hear 0ery faint soun+s- But that
+e%ree of amplification can 'e a pro'lem when we are e5pose+ to 0ery lou+ noises- 7n the
mi++le ear/ two small muscles protect the tympanic mem'rane an+ ossicles from 0iolent
mo0ements un+er 0ery noisy con+itions9
&$ The tensor tympani ;T"A>sor tim>P!A>eG= muscle is a short ri''on of muscle whose
ori%in is the petrous portion of the temporal 'one an+ the au+itory tu'e/ an+ whose insertion
is on the Dhan+leE of the malleus- 3hen the tensor tympani contracts/ the malleus is pulle+
me+ially/ stiffenin% the tympanic mem'rane- This increase+ stiffness re+uces the amount of
mo0ement
possi'le- The tensor tympani muscle is inner0ate+ 'y motor fi'ers of the man+i'ular 'ranch
of the tri%eminal ner0e ;$=-
%$ The stapedius ;sta>P
G"
>+
Ge
>us= muscle/ inner0ate+ 'y the facial ner0e ;$77=/ ori%inates from the posterior wall of the
mi++le
ear an+ inserts on the stapes- 1ontraction of the stape+ius pulls the stapes/ re+ucin%
mo0ement of the stapes at the o0al win+ow-
The 7nner "ar
The senses of e&uili'rium an+ hearin% are pro0i+e+ 'y receptors in the inner ear ;4i%ures
17<20 an+ 17<22a:=-
The superficial contours of the inner ear are esta'lishe+ 'y a layer of +ense 'one ,nown as
the bony labyrinth ;labyrinthos/ networ, of canals=- The walls of the 'ony la'yrinth are
continuous with the surroun+in% temporal 'one- The inner contours of the 'ony la'yrinth
closely follow the contours of the membranous labyrinth/ a +elicate/ interconnecte+
networ, of flui+>fille+ tu'es- The receptors of the inner ear are foun+ within those tu'es-
Between the 'ony an+ mem'ranous la'yrinths flows the perilymph ;P"R>i>limf=/ a li&ui+
248927081.doc 36
whose properties closely resem'le those of cere'rospinal flui+- The mem'ranous la'yrinth
contains
endolymph ;"A>+
Go
>limf=/ a flui+ with electrolyte concentrations +ifferent from those of typical 'o+y flui+s- The
physical rela
tionships are in+icate+ in 4i%ure 17<22':- ;See !ppen+i5 7$ for a chemical analysis of
perilymph/ en+olymph/ an+ other 'o+y flui+s-=
The 'ony la'yrinth can 'e su'+i0i+e+ into the vestibule/ three semicircular canals/ an+ the
cochlea ;see 4i%ure 17<22a:=- The
!estibule ;$"S>ti>'
Gu
l= consists of a pair of mem'ranous sacs9 the saccule ;S!N>
Gu
l= an+ the utricle ;
GH
>tri>,ul=/ or sacculus an+
utriculus- Receptors in the saccule an+ utricle pro0i+e sensations of %ra0ity an+ linear
acceleration-
The semicircular canals enclose slen+er semicircular ducts- Receptors in the semicircular
+ucts are stimulate+ 'y rotation of the hea+- The com'ination of 0esti'ule an+ semicircular
canals is calle+ the !estibular comple8- The flui+>fille+ cham'ers within the 0esti'ule are
'roa+ly continuous with those of the semicircular canals-
The cochlea ;NON>l
Ge
>uh2 cochlea/ a snail shell= is a spiral>shape+/ 'ony cham'er that contains the cochlear duct
of the mem
'ranous la'yrinth- Receptors within the cochlear +uct pro0i+e the sense of hearin%- The +uct
is san+wiche+ 'etween a pair of perilymph>fille+ cham'ers- The entire comple5 ma,es turns
aroun+ a central 'ony hu'/ much li,e a snail shell-
248927081.doc 37
The walls of the 'ony la'yrinth consist of +ense 'one e0erywhere e5cept at two small areas
near the 'ase of the cochlear spiral ;see 4i%ure 17<20:=- The round "indo" is a thin/
mem'ranous partition that separates the perilymph of the cochlear cham'ers from the air>
fille+ mi++le ear- 1olla%en fi'ers connect the 'ony mar%ins of the openin% ,nown as the o!al
"indo" to the 'ase of the stapes-
Equilibrium
!s ?ust note+/ e&uili'rium sensations are pro0i+e+ 'y receptors of the 0esti'ular comple5-
The semicircular +ucts con0ey information a'out rotational mo0ements of the hea+- 4or
e5ample/ when you turn your hea+ to the left/ receptors stimulate+ in the semicircular +ucts
tell you how rapi+ the mo0ement is/ an+ in which +irection- The saccule an+ the utricle
con0ey information a'out your position with respect to %ra0ity- 7f you stan+ with your hea+
tilte+ to one si+e/ these receptors report the an%le in0ol0e+ an+ whether your hea+ tilts
forwar+ or 'ac,war+- The saccule an+ the utricle are also stimulate+ 'y su++en acceleration-
3hen your car accelerates from a stop/ the saccular an+ utricular receptors %i0e you the
impression of increasin% spee+-
The Semicircular Ducts
Sensory receptors in the semicircular +ucts respon+ to rotational mo0ements of the hea+-
These hair cells are acti0e +urin% a mo0ement/ 'ut are &uiet when the 'o+y is motionless-
The anterior/ posterior/ an+ lateral semicircular ducts are continuous with the utricle
;4i%ure 17<23a:=- "ach semicircular +uct contains an ampulla/ an e5pan+e+ re%ion that
contains the receptors- The re%ion in the wall of the ampulla that contains the receptors is
,nown as a crista ;4i%ure 17<23':=- "ach crista is 'oun+ to a cupula
;N
GH
>p
Gu
248927081.doc 38
>luh=/ a %elatinous structure that e5ten+s the full wi+th of the ampulla- The receptors in the
cristae are calle+ hair cells
;4i%ure 17<23'/+:=-
(air cells are the receptors foun+ in other portions of the mem'ranous la'yrinth as well-
Re%ar+less of location/ they are always surroun+e+ 'y supportin% cells an+ monitore+ 'y the
+en+rites of sensory neurons- The free surface of each hair cell supports *0<100 lon%
stereocilia which resem'le 0ery lon% micro0illi ;see 4i%ure 17<23+:=- "ach hair cell in the
0esti'ule also con
G
7 when an e5ternal force pushes a%ainst these processes/ the +istortion of the cell mem'rane
alters the rate at which the hair cell releases chemical transmitters-
(air cells pro0i+e information a'out the +irection an+ stren%th of mechanical stimuli- The
stimuli in0ol0e+/ howe0er/ +epen+ on the hair cellFs location9 %ra0ity or acceleration in the
0esti'ule/ rotation in the semicircular canals/ an+ soun+ in the cochlea- The sensiti0ities of
the hair cells +iffer/ 'ecause each of these re%ions has +ifferent accessory structures that
+etermine which stimulus will pro0i+e the force to +eflect the ,inocilia an+ stereocilia-
!t a crista/ the ,inocilia an+ stereocilia of the hair cells are em'e++e+ in the cupula ;see
4i%ure 17<23':=- Because the cupula has a +ensity 0ery close to that of the surroun+in%
en+olymph/ it essentially floats a'o0e the receptor surface- 3hen your hea+ rotates in the
plane of the +uct/ the mo0ement of en+olymph alon% the len%th of the semicircular +uct
pushes the cupula to the si+e an+ +istorts the receptor processes ;4i%ure 17<23c:=-
@o0ement of flui+ in one +irection stimulates the hair cells/ an+ mo0ement in the opposite
+irection inhi'its them- 3hen the en+olymph stops mo0in%/ the elastic nature of the cupula
ma,es it return to its normal position-
"0en the most comple5 mo0ement can 'e analy.e+ in terms of motion in three rotational
planes- "ach semicircular +uct re
tains a 3inocilium ;N
Go
>S78>
Ge
248927081.doc 39
>um=/ a sin%le lar%e cilium- (air cells +o not acti0ely mo0e their ,inocilia or stereocilia-
(owe0er/
>n
spon+s to one of these rotational mo0ements- ! hori.ontal rotation/ as in sha,in% your hea+
Dno/E stimulates the hair cells of the lateral semicircular +uct- Ao++in% DyesE e5cites the
anterior +uct/ an+ tiltin% your hea+ from si+e to si+e acti0ates receptors in the posterior
+uct-
The Htricle an+ Saccule
The utricle an+ saccule pro0i+e e&uili'rium sensations/ whether the 'o+y is mo0in% or is
stationary- The two cham'ers are connecte+ 'y a slen+er passa%eway that is continuous with
the narrow en+olymphatic +uct- The endolymphatic duct en+s in a 'lin+ pouch calle+ the
endolymphatic sac ;see 4i%ure 17<23a:=- This sac pro?ects throu%h the +ura mater that
lines the temporal 'one an+ into the su'arachnoi+ space/ where it is surroun+e+ 'y a
capillary networ,- Portions of the cochlear +uct secrete en+olymph continuously/ an+ at the
en+olymphatic sac e5cess flui+ returns to the %eneral circulation as the capillaries a'sor'
en+olymph remo0e+ 'y a com'ination of acti0e transport an+ 0esicular transport-
The hair cells of the utricle an+ saccule are clustere+ in o0al structures calle+ maculae
;@!N>
Gu
>l
Ge
2 macula/ spot=
;4i%ure 17<2a:=- !s in the ampullae/ the hair cell processes are em'e++e+ in a %elatinous
mass- (owe0er/ the surface of this %elatinous material contains +ensely pac,e+ calcium
car'onate crystals ,nown as statoconia ;statos/ stan+in% I conia/ +ust=- The comple5 as a
whole ;%elatinous matri5 an+ statoconia= is calle+ an otolith ;Dear stoneE=-
The macula of the saccule is +ia%ramme+ in 4i%ure 17<2':/ an+ its function is shown in
4i%ure 17<2c:- 3hen your hea+ is in the normal/ upri%ht position/ the statoconia sit atop
the macula ;ST"P 1=- Their wei%ht presses on the macular surface/ pushin% the hair cell
processes +own rather than to one si+e or another- 3hen your hea+ is tilte+/ the pull of
248927081.doc 40
%ra0ity on the statoconia shifts them to the si+e/ there'y +istortin% the hair cell processes
;ST"P 2=- The chan%e in receptor acti0ity tells the 1AS that your hea+ is no lon%er le0el-
! similar mechanism accounts for your perception of linear acceleration when you are in a
car that spee+s up su++enly- The statoconia la% 'ehin+/ an+ the effect on the hair cells is
compara'le to tiltin% your hea+ 'ac,- Hn+er normal circumstances/ your ner0ous system
+istin%uishes 'etween the sensations of tiltin% an+ linear acceleration 'y inte%ratin%
0esti'ular sensations with 0isual information- @any amusement par, ri+es confuse your
sense of e&uili'rium 'y com'inin% rapi+ rotation with chan%es in position an+ acceleration
while pro0i+in% restricte+ or mislea+in% 0isual information-
Pathways for "&uili'rium Sensations
(air cells of the 0esti'ule an+ semicircular +ucts are monitore+ 'y sensory neurons locate+
in a+?acent !estibular ganglia- Sensory fi'ers from these %an%lia form the !estibular
branch of the 0esti'ulocochlear ner0e ;$777=- lp$ (6@ These fi'ers inner0ate neurons within
the pair of !estibular nuclei at the 'oun+ary 'etween the pons an+ the me+ulla o'lon%ata-
The two 0esti'ular nuclei ha0e four functions9
&$ 7nte%ratin% sensory information a'out 'alance an+ e&uili'rium that arri0es from 'oth si+es
of the hea+-
%$ Relayin% information from the 0esti'ular comple5 to the cere'ellum-
5$ Relayin% information from the 0esti'ular comple5 to the cere'ral corte5/ pro0i+in% a
conscious sense of hea+ position an+ mo0ement-
($ Sen+in% comman+s to motor nuclei in the 'rain stem an+ spinal cor+-
The refle5i0e motor comman+s issue+ 'y the 0esti'ular nuclei are +istri'ute+ to the motor
nuclei for cranial ner0es in0ol0e+
with eye/ hea+/ an+ nec, mo0ements ;777/ 7$/ $7/ an+ J7=- 7nstructions +escen+in% in the
vestibulospinal tracts of the spinal cor+ a+?ust peripheral muscle tone an+ complement the
refle5i0e mo0ements of the hea+ or nec,- lp$ @&& These pathways are in+icate+ in 4i%ure
17<25:-
The automatic mo0ements of the eyes that occur in response to sensations of motion are
+irecte+ 'y the superior colliculi of the mesencephalon- lp$ ('( These mo0ements attempt
248927081.doc 41
to ,eep your %a.e focuse+ on a specific point in space/ +espite chan%es in 'o+y position an+
orientation- 7f your 'o+y is turnin% or spinnin% rapi+ly/ your eyes will fi5 on one point for a
moment an+ then ?ump ahea+ to another in a series of short/ ?er,y mo0ements- This type of
eye mo0ement can occur e0en when the 'o+y is stationary if either the 'rain stem or the
inner ear is +ama%e+- 7n+i0i+uals with this con+ition/ which is calle+ nystagmus ;nis>T!G>
mus=/ ha0e trou'le controllin% their eye mo0ements- Physicians commonly chec, for
nysta%mus 'y as,in% patients to watch a small penli%ht as it is mo0e+ across the fiel+ of
0ision- !@9 $erti%o/ @otion Sic,ness/ an+ @QniRreFs Disease
Hearing
The receptors of the cochlear +uct pro0i+e a sense of hearin% that ena'les us to +etect the
&uietest whisper/ yet remain functional in a noisy room- The receptors responsi'le for
au+itory sensations are hair cells similar to those of the 0esti'ular comple5- (owe0er/ their
placement within the cochlear +uct an+ the or%ani.ation of the surroun+in% accessory
structures shiel+ them from stimuli other than soun+-
7n con0eyin% 0i'rations from the tympanic mem'rane to the o0al win+ow/ the au+itory
ossicles con0ert pressure fluctuations in air into much %reater pressure fluctuations in the
perilymph of the cochlea- These fluctuations stimulate hair cells alon% the cochlear spiral-
The fre#uency of the percei0e+ soun+ is +etermine+ 'y which part of the cochlear +uct is
stimulate+- The intensity ;0olume= of the percei0e+ soun+ is +etermine+ 'y how many of the
hair cells at that location are stimulate+- 3e will now consi+er the mechanics of this
remar,a'ly ele%ant process-
The 1ochlear Duct
7n sectional 0iew ;4i%ure 17<2#a/' an+ 17<27a/':=/ the cochlear +uct/ or scala media/ lies
'etween a pair of perilymphatic cham'ers9 the !estibular duct ;scala vestibuli= an+ the
tympanic duct ;scala tympani=- The outer surfaces of these +ucts are encase+ 'y the 'ony
la'yrinth e0erywhere e5cept at the o0al win+ow ;the 'ase of the 0esti'ular +uct= an+ the
roun+ win+ow ;the 'ase of the tympanic +uct=- Because the 0esti'ular an+ tympanic +ucts
are interconnecte+ at the tip of the cochlear spiral/ they really form one lon% an+ continuous
perilymphatic cham'er- This cham'er 'e%ins at the o0al win+ow2 e5ten+s throu%h the
248927081.doc 42
0esti'ular +uct/ aroun+ the top of the cochlea/ an+ alon% the tympanic +uct2 an+ en+s at the
roun+ win+ow-
The cochlear +uct is an elon%ate+ tu'eli,e structure suspen+e+ 'etween the 0esti'ular +uct
an+ the tympanic +uct- The hair cells of the cochlear +uct are locate+ in a structure calle+
the organ of Corti ;see 4i%ures 17<2#' an+ 17<27a/':=- This sensory structure sits on the
basilar membrane/ a mem'rane that separates the cochlear +uct from the tympanic +uct-
The hair cells are arran%e+ in a series of lon%itu+inal rows- They lac, ,inocilia/ an+ their
stereocilia are in contact with the o0erlyin% tectorial ;te,>TOR>eG>al= membrane ;tectum/
roof=- This mem'rane is firmly attache+ to the inner wall of the cochlear +uct- 3hen a
portion of the 'asilar mem'rane 'ounces up an+ +own/ the stereocilia of the hair cells are
presse+ a%ainst the tectorial mem'rane an+ 'ecome +istorte+- The 'asilar mem'rane mo0es
in response to pressure fluctuations within the perilymph- These pressure chan%es are
tri%%ere+ 'y soun+ wa0es arri0in% at the tympanic mem'rane- To un+erstan+ this process/
we must consi+er the 'asic properties of soun+-
!n 7ntro+uction to Soun+
(earin% is the perception of soun+/ which consists of wa0es of pressure con+ucte+ throu%h
a me+ium such as air or water- 7n air/ each pressure wave consists of a re%ion where the air
molecules are crow+e+ to%ether an+ an a+?acent .one where they are farther apart ;4i%ure
17<2*a:=- These wa0es are sine wa0esKthat is/ S>shape+ cur0es that repeat in a re%ular
patternKan+ tra0el throu%h
the air at a'out 1235 ,m P h ;7#* mph=-
The wavelength of soun+ is the +istance 'etween two a+?acent wa0e crests ;pea,s= or/
e&ui0alently/ the +istance 'etween two a+?acent wa0e trou%hs ;4i%ure 17<2*':=- 3a0elen%th
is in0ersely relate+ to frequencyKthe num'er of wa0es that pass a fi5e+ reference point in a
%i0en time- Physicists use the term cycles rather than waves- (ence/ the fre&uency of a
soun+ is measure+ in terms of the num'er of cycles per secon+ ;cps=/ a unit calle+ hert9
:H9;- 3hat we percei0e as the pitch of a soun+ is our sensory response to its fre&uency- !
high-fre#uency soun+ ;hi%h pitch/ short wa0elen%th= mi%ht ha0e a fre&uency of 15/000 (.
or more2 a 0ery low-fre#uency soun+ ;low pitch/ lon% wa0elen%th= coul+ ha0e a fre&uency of
100 (. or less-
248927081.doc 43
7t ta,es ener%y to pro+uce soun+ wa0es- 3hen you stri,e a tunin% for,/ it 0i'rates an+
pushes a%ainst the surroun+in% air/ pro+ucin% soun+ wa0es whose fre&uency +epen+s on the
instrumentFs fre&uency of 0i'ration- The har+er you stri,e the tunin% for,/ the more ener%y
you pro0i+e2 the ener%y increases the amplitude of the soun+ wa0e ;see 4i%ure 17<2*':=-
The amplitu+e/ or intensity/ of a soun+ +etermines how lou+ it seems2 the %reater the ener%y
content/ the lar%er the amplitu+e/ an+ the lou+er the soun+- Soun+ ener%y is reporte+ in
decibels ;D"S>i>'el.=- Ta'le 17<1 in+icates the +eci'el le0els of familiar soun+s-
3hen soun+ wa0es stri,e an o'?ect/ their ener%y is a physical pressure- 6ou may ha0e seen
win+ows mo0e in a room in which a stereo is 'lastin%- The more fle5i'le the o'?ect/ the
more easily it will respon+ to soun+ pressure- "0en soft stereo music will 0i'rate a sheet of
paper hel+ in front of the spea,er- Gi0en the ri%ht com'ination of fre&uencies an+
amplitu+es/ an o'?ect will 'e%in to 0i'rate at the same fre&uency as the soun+/ a
phenomenon calle+ resonance- The hi%her the +eci'el le0el/ the %reater the amount of
0i'ration- 4or you to 'e a'le to hear any soun+/ your thin/ fle5i'le tympanic mem'rane must
0i'rate in resonance with the soun+ wa0es-
Pro'a'ly more than # million people in the Hnite+ States alone ha0e at least a partial hearin%
+eficit/ +ue to pro'lems with either the transfer of 0i'rations 'y the au+itory ossicles or
+ama%e to the receptors or the au+itory pathways- !@9 (earin% Deficits
The (earin% Process
The process of hearin% can 'e +i0i+e+ into si5 'asic steps ;4i%ure 17<2):=9
(tep 0 Soun+ 3a0es !rri0e at the Tympanic @em'rane- Soun+ wa0es enter the e5ternal
acoustic canal an+ tra0el towar+ the tympanic mem'rane- The orientation of the canal
pro0i+es some +irectional sensiti0ity- Soun+ wa0es approachin% a particular si+e of the hea+
ha0e +irect access to the tympanic mem'rane on that si+e/ whereas soun+s arri0in% from
another +irection must 'en+ aroun+ corners or pass throu%h the auricle or other 'o+y
tissues-
(tep 1 @o0ement of the Tympanic @em'rane 1auses Displacement of the !u+itory
Ossicles- The tympanic mem'rane pro0i+es a surface for the collection of soun+/ an+ it
0i'rates in resonance to soun+ wa0es with fre&uencies 'etween appro5imately 20 an+
248927081.doc 44
20/000 (.- 3hen the tympanic mem'rane 0i'rates/ so +o the malleus an+/ throu%h their
articulations/ the incus an+ stapes- 7n this way/ the soun+ is amplifie+-
(tep 2 @o0ement of the Stapes at the O0al 3in+ow "sta'lishes Pressure 3a0es in the
Perilymph of the $esti'ular Duct- 8i&ui+s are incompressi'le9 7f you push +own on one part
of a water 'e+/ the 'e+ 'ul%es somewhere else- Because the rest of the cochlea is sheathe+
in 'one/ pressure applie+ at the o0al win+ow can 'e relie0e+ only at the roun+ win+ow-
!lthou%h the stapes actually has a roc,in% mo0ement/ the in<out component is easiest to
0isuali.e an+ +escri'e- Basically/ when the stapes mo0es inwar+/ the roun+ win+ow 'ul%es
outwar+/ into the mi++le ear ca0ity- !s the stapes mo0es in an+ out/ 0i'ratin% at the
fre&uency of the soun+ arri0in% at the tympanic mem'rane/ it creates pressure wa0es within
the perilymph-
(tep 3 The Pressure 3a0es Distort the Basilar @em'rane on Their 3ay to the Roun+
3in+ow of the Tympanic Duct- The pressure wa0es esta'lishe+ 'y the mo0ement of the
stapes tra0el throu%h the perilymph of the 0esti'ular an+ tympanic +ucts to reach the roun+
win+ow- 7n +oin% so/ the wa0es +istort the 'asilar mem'rane- The location of ma5imum
+istortion 0aries with the fre&uency of the soun+/ owin% to re%ional +ifferences in the wi+th
an+ fle5i'ility of the 'asilar mem'rane alon% its len%th- (i%h>fre&uency soun+s/ which ha0e a
0ery short wa0elen%th/ 0i'rate the 'asilar mem'rane near the o0al win+ow- The lower the
fre&uency of the soun+/ the lon%er the wa0elen%th/ an+ the farther from the o0al win+ow the
area of ma5imum +istortion will 'e ;4i%ure 17<30a<c:=- Thus/ information a'out fre&uency
is translate+ into information a'out position alon% the 'asilar mem'rane-
The amount of mo0ement at a %i0en location +epen+s on the amount of force applie+ 'y the
stapes/ which in turn is a function of ener%y content of the soun+- The lou+er the soun+/ the
more the 'asilar mem'rane mo0es-
(tep 6 $i'ration of the Basilar @em'rane 1auses $i'ration of (air 1ells a%ainst the
Tectorial @em'rane- $i'ration of the affecte+ re%ion of the 'asilar mem'rane mo0es hair
cells a%ainst the tectorial mem'rane- This mo0ement lea+s to the +isplacement of the
stereocilia/ which in turn opens ion channels in the hair cell mem'ranes- The resultin% inrush
of ions +epolari.es the hair cells/ lea+in% to the release of neurotransmitters an+ thus to the
stimulation of sensory neurons-
248927081.doc 45
The hair cells of the or%an of 1orti are arran%e+ in se0eral rows- ! 0ery soft soun+ may
stimulate only a few hair cells in a portion of one row- !s the intensity of a soun+ increases/
not only +o these hair cells 'ecome more acti0e/ 'ut a++itional hair cellsK at first in the
same row an+ then in a+?acent rowsKare stimulate+ as well- The num'er of hair cells
respon+in% in a %i0en re%ion of the or%an of 1orti thus pro0i+es information on the intensity
of the soun+-
(tep 7 7nformation a'out the Re%ion an+ 7ntensity of Stimulation 7s Relaye+ to the 1AS
o0er the 1ochlear Branch of the $esti'ulocochlear Aer0e ;$777=- The cell 'o+ies of the
'ipolar sensory neurons that monitor the cochlear hair cells are locate+ at the center of the
'ony cochlea/ in the spiral ganglion ;see 4i%ure 17<27a:=- 4rom there/ the information is
carrie+ 'y the cochlear 'ranch of cranial ner0e $777 to the cochlear nuclei of the me+ulla
o'lon%ata for su'se&uent +istri'ution to other centers in the 'rain-
!u+itory Pathways
Stimulation of hair cells acti0ates sensory neurons whose cell 'o+ies are in the a+?acent
spiral %an%lion- The afferent fi'ers of those neurons form the cochlear branch of the
0esti'ulocochlear ner0e ;$777= ;4i%ure 17<31:=- These a5ons enter the me+ulla o'lon%ata/
where they synapse at the dorsal an+ !entral cochlear nuclei- 4rom there/ the information
crosses to the opposite si+e of the 'rain an+ ascen+s to the inferior colliculus of the
mesencephalon- This processin% center coor+inates a num'er of responses to acoustic
stimuli/ inclu+in% au+itory refle5es that in0ol0e s,eletal muscles of the hea+/ face/ an+ trun,-
These refle5es automatically chan%e the position of your hea+ in response to a su++en lou+
noise2 you usually turn your hea+ an+ your eyes towar+ the source of the soun+-
Before reachin% the cere'ral corte5 an+ your awareness/ ascen+in% au+itory sensations
synapse in the me+ial %eniculate nucleus of the thalamus- Pro?ection fi'ers then +eli0er the
information to the au+itory corte5 of the temporal lo'e- 7nformation tra0els to the corte5
o0er la'ele+ lines9 (i%h>fre&uency soun+s acti0ate one portion of the corte5/ low>fre&uency
soun+s another- 7n effect/ the au+itory corte5 contains a map of the or%an of 1orti- Thus/
information a'out fre#uency/ translate+ into information a'out position on the 'asilar
mem'rane/ is pro?ecte+ in that form onto the au+itory corte5/ where it is interprete+ to
pro+uce your su'?ecti0e sensation of pitch-
248927081.doc 46
!n in+i0i+ual whose au+itory corte5 is +ama%e+ will respon+ to soun+s an+ ha0e normal
acoustic refle5es/ 'ut will fin+ it +ifficult or impossi'le to interpret the soun+s an+ reco%ni.e
a pattern in them- Dama%e to the a+?acent association area lea0es the a'ility to +etect the
tones an+ patterns/ 'ut pro+uces an ina'ility to comprehen+ their meanin%-
!u+itory Sensiti0ity
Our hearin% a'ilities are remar,a'le/ 'ut it is +ifficult to assess the a'solute sensiti0ity of the
system- The ran%e from the softest au+i'le soun+ to the lou+est tolera'le 'last represents a
trillionfol+ increase in power- The receptor mechanism is so sensiti0e that/ if we were to
remo0e the stapes/ we coul+/ in theory/ hear air molecules 'ouncin% off the o0al win+ow- 3e
ne0er use the full potential of this system/ 'ecause 'o+y mo0ements an+ our internal or%ans
pro+uce s&uea,s/ %roans/ thumps/ an+ other soun+s that are tune+ out 'y central an+
peripheral a+aptation- 3hen other en0ironmental noises fa+e away/ the le0el of a+aptation
+rops an+ the system 'ecomes increasin%ly sensiti0e- 4or e5ample/ when you rela5 in a &uiet
room/ your heart'eat seems to %et lou+er an+ lou+er as the au+itory system a+?usts to the
le0el of 'ac,%roun+ noise-
6oun% chil+ren ha0e the %reatest hearin% ran%e9 They can +etect soun+s ran%in% from a 20>
(. 'u.. to a 20/000>(. whine- 3ith a%e/ +ama%e +ue to lou+ noises or other in?uries
accumulates- The tympanic mem'rane %ets less fle5i'le/ the articulations 'etween the
ossicles stiffen/ an+ the roun+ win+ow may 'e%in to ossify- !s a result/ ol+er in+i0i+uals
show some +e%ree of hearin% loss-
&11 Keys 2 Balance an+ hearin% rely on the same 'asic types of sensory receptors ;hair
cells=- The anatomical structure of the associate+ sense or%an +etermines what stimuli affect
the hair cells- 7n the semicircular +ucts/ the stimulus is flui+ mo0ement cause+ 'y hea+
rotation in the hori.ontal/ sa%ittal/ or frontal planes- 7n the utricle an+ saccule/ the stimuli are
%ra0ity>in+uce+ shifts in the position of attache+ otoliths- 7n the cochlea/ the stimulus is
mo0ement of the tectorial mem'rane as pressure wa0es +istort the 'asilar mem'rane-
Concept Chec3
7f the roun+ win+ow were not a'le to 'ul%e out with increase+ pressure in the perilymph/
how woul+ the perception of soun+ 'e affecte+M
(ow woul+ the loss of stereocilia from hair cells of the or%an of 1orti affect hearin%M
248927081.doc 47
3hy +oes a 'loc,a%e of the au+itory tu'e pro+uce an earacheM
Ans"ers begin on p$ A4&
Anatomy 5'1 2 Re0iew the anatomy of the ear on the Anatomy 5'1 C+-O<> #er!ous
System?Special Senses?Ear$
Chapter e!ie"
Selected Clinical ,erminology
astigmatism> Re+uction in 0isual acuity +ue to a cur0ature irre%ularity in the cornea or lens-
;p- 5#5=
cataract> !n a'normal con+ition in which the lens has lost its transparency- ;p- 5#2=
color blindness> ! con+ition in which a person is una'le to +istin%uish certain colors- ;p-
570=
conducti!e deafness> Deafness resultin% from con+itions in the outer or mi++le ear that
'loc, the transfer of 0i'rations from the tympanic mem'rane to the o0al win+ow- S!@T
detached retina> Delamination of a portion of the neural retina/ which separates the
photoreceptor layer from the pi%ment layer- 7f untreate+/ 'lin+ness can result in the affecte+
area- ;p- 5#1=
diabetic retinopathy> Deterioration of the retinal photoreceptor layer +ue to 0ascular
+ama%e an+ the o0er%rowth an+ rupture of 'loo+ 0essels on the retinal surface- ;p- 55)=
glaucoma> ! con+ition characteri.e+ 'y increase+ intraocular pressure +ue to the impaire+
rea'sorption of a&ueous humor2 can result in 'lin+ness- ;p- 5#2=
hyperopiaA or farsightedness: ! con+ition in which near'y o'?ects are 'lurry/ 'ut +istant
o'?ects are clear- ;p- 5#=
motion sic3ness> ! con+ition resultin% from conflictin% 0isual an+ e&uili'rium sensory
stimuli- Si%ns an+ symptoms can inclu+e hea+ache/ sweatin%/ nausea/ 0omitin%/ an+ chan%es
in mental state- S!@T
myopiaA or nearsightedness: ! con+ition in which 0ision at close ran%e is normal/ 'ut
+istant o'?ects appear 'lurry- ;p- 5#=
248927081.doc 48
ner!e deafness> Deafness resultin% from pro'lems within the cochlea or alon% the au+itory
pathway- S!@T
night blindness> 8oss of 0isual acuity un+er +im li%ht con+itions +ue to ina+e&uate 0isual
pi%ment pro+uction/ usually as a result of 0itamin ! +eficiency- ;p- 570=
nystagmus> !'normal eye mo0ements that may appear after 'rain stem or inner ear
+ama%e- ;p- 57)=
otitis media> 7nfection an+ tissue inflammation within the mi++le ear ca0ity- ;p- 57 an+
S!@T=
presbyopia> ! type of hyperopia that +e0elops with a%e as lenses 'ecome less elastic- ;p-
5#=
retinitis pigmentosa> ! %roup of inherite+ retinopathies characteri.e+ 'y the pro%ressi0e
+eterioration of photoreceptors/ e0entually resultin% in 'lin+ness- ;p- 5#7=
scotomas> !'normal 'lin+ spots that are fi5e+ in position- ;p- 5##=
Study Outline
Olfaction p- 550 Olfactory Receptors p- 551
&$ The olfactory organs contain the olfactory epithelium with olfactory receptors/
supportin% cells/ an+ basal ;stem= cells- The surfaces of the olfactory or%ans are coate+ with
the secretions of the olfactory glands- !)igure 0890"
%$ The olfactory receptors are hi%hly mo+ifie+ neurons-
5$ Olfactory reception in0ol0es +etectin% +issol0e+ chemicals as they interact with
o+orant>'in+in% proteins-
Olfactory Pathways p- 551
($ 7n olfaction/ the arri0in% information reaches the information centers without first
synapsin% in the thalamus- !)igure 0890"
Olfactory Discrimination p- 551
@$ The olfactory system can +istin%uish thousan+s of chemical stimuli- The 1AS
interprets smells 'y the pattern of receptor acti0ity-
'$ The olfactory receptor population shows consi+era'le turno0er- The num'er of
olfactory receptors +eclines with a%e-
248927081.doc 49
Gustation p- 552
&$ ,aste ;%ustatory= receptors are clustere+ in taste buds-
%$ Taste 'u+s are associate+ with epithelial pro?ections ;lingual papillae= on the +orsal
surface of the ton%ue- !)igure 0891"
Taste Receptors p- 553
5$ "ach taste 'u+ contains basal cells/ which appear to 'e stem cells/ an+ gustatory
cells/ which e5ten+ taste hairs throu%h a narrow taste pore- !)igure 0891"
Gustatory Pathways p- 553
($ The taste 'u+s are monitore+ 'y cranial ner0es that synapse within the solitary
nucleus of the me+ulla o'lon%ata an+ then on to the thalamus an+ the primary sensory
corte5-
Gustatory Discrimination p- 553
@$ The primary taste sensations are sweet/ salt/ sour/ an+ 'itter- Receptors also e5ist
for umami an+ "ater-
'$ Taste sensiti0ity e5hi'its si%nificant in+i0i+ual +ifferences/ some of which are
inherite+-
*$ The num'er of taste 'u+s +eclines with a%e-
&11 Keys 2 p- 55
Vision p- 55 !ccessory Structures of the "ye p- 55
&$ The accessory structures of the eye inclu+e the eyeli+s :palpebrae;/ separate+ 'y
the palpebral fissure/ the eyelashes/ an+ the tarsal glands- !)igures 0892: 0893"
%$ !n epithelium calle+ the con7uncti!a co0ers most of the e5pose+ surface of the eye-
The cornea is transparent- !)igures 0892: 0893"
5$ The secretions of the lacrimal gland contain lyso9yme- Tears collect in the lacrimal
lae an+ reach the inferior meatus of the nose after they pass throu%h the lacrimal puncta/
the lacrimal canaliculi/ the lacrimal sac/ an+ the nasolacrimal duct- !)igure 0892"
The "ye p- 557
248927081.doc 50
($ The eye has three layers9 an outer fibrous tunic/ a mi++le !ascular tunic/ an+ an
inner neural tunic- !)igure 0893"
@$ The fi'rous tunic consists of the sclera/ the cornea/ an+ the limbus- !)igure 0893"
'$ The 0ascular tunic/ or u!ea/ inclu+es the iris/ the ciliary body/ an+ the choroid- The
iris contains muscle fi'ers that chan%e the +iameter of the pupil- The ciliary 'o+y contains
the ciliary muscle an+ the ciliary processes/ which attach to the suspensory ligaments of
the lens- !)igures 0893: 0896"
*$ The neural tunic/ or retina/ consists of an outer pigmented part an+ an inner neural
part2 the latter contains 0isual receptors an+ associate+ neurons- !)igures 0893: 0897"
6$ The retina contains two types of photoreceptors> rods an+ cones-
=$ 1ones are +ensely clustere+ in the fo!ea/ at the center of the macula lutea- !)igure
0897"
&1$ The +irect line to the 1AS procee+s from the photoreceptors to bipolar cells/ then to
ganglion cells/ an+/ finally/ to the 'rain 0ia the optic ner0e- The a5ons of %an%lion cells
con0er%e at the optic disc/ or blind spot- Hori9ontal cells an+ amacrine cells mo+ify the
si%nals passe+ amon% other components of the retina- !)igures 0897: 0898"
&&$ The ciliary 'o+y an+ lens +i0i+e the interior of the eye into a lar%e posterior ca!ity/ or
vitreous chamber/ an+ a smaller anterior ca!ity- The anterior ca0ity is su'+i0i+e+ into the
anterior chamber/ which e5ten+s from the cornea to the iris/ an+ a posterior chamber/
'etween the iris an+ the ciliary 'o+y an+ lens- !)igure 089;"
&%$ The flui+ aqueous humor circulates within the eye an+ reenters the circulation after
+iffusin% throu%h the walls of the anterior cham'er an+ into the canal of Schlemm- !)igure
089;"
&5$ The lens lies posterior to the cornea an+ forms the anterior 'oun+ary of the posterior
ca0ity- This ca0ity contains the !itreous body/ a %elatinous mass that helps sta'ili.e the
shape of the eye an+ support the retina- !)igure 089;"
&($ The lens focuses a 0isual ima%e on the photoreceptors- The con+ition in which a lens has
lost its transparency is a cataract-
&@$ 8i%ht is refracted ;'ent= when it passes throu%h the cornea an+ lens- Durin%
accommodation/ the shape of the lens chan%es to focus an ima%e on the retina- DAormalE
!isual acuity is rate+ 20 P 20- !)igures 089< to 08901"
248927081.doc 51
&11 Keys 2 p- 5##
$isual Physiolo%y p- 5##
&'$ The two types of photoreceptors are ro+s/ which respon+ to almost any photon/
re%ar+less of its ener%y content/ an+ cones/ which ha0e characteristic ran%es of sensiti0ity-
!)igure 08902"
&*$ "ach photoreceptor contains an outer segment with mem'ranous discs- ! narrow stal,
connects the outer se%ment to the inner segment- 8i%ht a'sorption occurs in the !isual
pigments/ which are +eri0ati0es of rhodopsin ;opsin plus the pi%ment retinal/ which is
synthesi.e+ from 0itamin !=- !)igures 08902 to 08906"
&6$ 1olor sensiti0ity +epen+s on the inte%ration of information from red/ green/ an+ blue
cones- Color blindness is the ina'ility to +etect certain colors- !)igures 08907: 08908"
&=$ 7n the dar3-adapted state/ most 0isual pi%ments are fully recepti0e to stimulation- 7n
the light-adapted state/ the pupil constricts an+ bleaching of the 0isual pi%ments occurs-
The $isual Pathway p- 571
%1$ The %an%lion cells that monitor ro+s/ calle+ < cells ;magnocells=/ are relati0ely lar%e-
The %an%lion cells that monitor cones/ calle+ ) cells ;parvo cells=/ are smaller an+ more
numerous- !)igure 0890;"
%&$ $isual +ata from the left half of the com'ine+ fiel+ of 0ision arri0e at the 0isual corte5 of
the ri%ht occipital lo'e2 +ata from the ri%ht half of the com'ine+ fiel+ of 0ision arri0e at the
0isual corte5 of the left occipital lo'e- !)igure 0890<"
%%$ +epth perception is o'taine+ 'y comparin% relati0e positions of o'?ects 'etween the
left> an+ ri%ht>eye ima%es !)igure 0890<"
%5$ $isual inputs to the suprachiasmatic nucleus of the hypothalamus affect the function of
other 'rain stem nuclei- This nucleus esta'lishes a 0isceral circadian rhythm/ which is tie+
to the +ay<ni%ht cycle an+ affects other meta'olic processes-
Anatomy 5'1 2 Aer0ous SystemUSpecial SensesU"ye
248927081.doc 52
Equilibrium and Hearing p- 573
&$ The senses of e&uili'rium an+ hearin% are pro0i+e+ 'y the receptors of the inner ear-
!natomy of the "ar p- 573
%$ The ear is +i0i+e+ into the e8ternal ear/ the middle ear/ an+ the inner ear- !)igure
0891="
5$ The e5ternal ear inclu+es the auricle/ or pinna/ which surroun+s the entrance to the
e8ternal acoustic canal/ which en+s at the tympanic membrane ;eardrum=- !)igure 089
1="
($ The mi++le ear communicates with the nasopharyn5 0ia the auditory
;pharyngotympanic= tube- The mi++le ear encloses an+ protects the auditory ossicles-
!)igures 0891=: 08910"
@$ The membranous labyrinth ;the cham'ers an+ tu'es= of the inner ear contains the
flui+ endolymph- The bony labyrinth surroun+s an+ protects the mem'ranous la'yrinth
an+ can 'e su'+i0i+e+ into the !estibule/ the semicircular canals/ an+ the cochlea-
!)igures 0891=: 08911"
'$ The 0esti'ule of the inner ear encloses the saccule an+ utricle- The semicircular
canals contain the semicircular ducts- The cochlea contains the cochlear duct/ an elon%ate+
portion of the mem'ranous la'yrinth- !)igure 08911"
*$ The round "indo" separates the perilymph from the air spaces of the mi++le ear-
The o!al "indo" is connecte+ to the 'ase of the stapes- !)igure 0891="
"&uili'rium p- 57#
6$ The 'asic receptors of the inner ear are hair cells/ which pro0i+e information a'out
the +irection an+ stren%th of mechanical stimuli- !)igure 08912"
=$ The anterior/ posterior/ an+ lateral semicircular ducts are continuous with the
utricle- "ach +uct contains an ampulla with a %elatinous cupula an+ associate+ sensory
receptors- !)igures 08911: 08912"
&1$ The saccule an+ utricle are connecte+ 'y a passa%eway that is continuous with the
endolymphatic duct/ which terminates in the endolymphatic sac- ln the saccule an+
248927081.doc 53
utricle/ hair cells cluster within maculae/ where their cilia contact the otolith ;+ensely
pac,e+ mineral crystals/ calle+ statoconia/ in a matri5=- !)igures 08912: 08913"
&&$ The 0esti'ular receptors acti0ate sensory neurons of the !estibular ganglia- The a5ons
form the !estibular branch of the 0esti'ulocochlear ner0e ;$777=/ synapsin% within the
!estibular nuclei- !)igure 08916"
(earin% p- 57)
&%$ The cochlear +uct lies 'etween the !estibular duct an+ the tympanic duct- The hair
cells of the cochlear +uct lie within the organ of Corti- !)igures 08917: 08918"
&5$ The ener%y content of a soun+ +etermines its intensity/ measure+ in decibels- Soun+
wa0es tra0el towar+ the tympanic mem'rane/ which 0i'rates2 the au+itory ossicles con+uct
these 0i'rations to the inner ear- @o0ement at the o0al win+ow applies pressure to the
perilymph of the 0esti'ular +uct- !)igures 0891;: 0891<> $able 0890"
&($ Pressure wa0es +istort the basilar membrane an+ push the hair cells of the or%an of
1orti a%ainst the tectorial membrane- The tensor tympani an+ stapedius muscles
contract to re+uce the amount of motion when 0ery lou+ soun+s arri0e- !)igures 0891<:
-0892="
&@$ The sensory neurons are locate+ in the spiral ganglion of the cochlea- The afferent
fi'ers of these neurons form the cochlear branch of the 0esti'ulocochlear ner0e ;$777=/
synapsin% at the cochlear nuclei- !)igure 08920"
&11 Keys 2 p- 5*#
Anatomy 5'1 2 Aer0ous SystemUSpecial SensesU"ar
e!ie" Buestions
<yAC) 2 !ccess more re0iew material online at <yAC)- There youFll fin+ learnin%
acti0ities/ case stu+ies/ &ui..es/ 7nteracti0e Physiolo%y e5ercises/ an+ more to help you
succee+- To access the site/ %o to www-myaan+p-com-
?nswers to the *eview @uestions begin on page ?90'
8"$"8 1 Re0iewin% 4acts an+ Terms
248927081.doc 54
&$ ! re+uction in sensiti0ity in the presence of a constant stimulus is
:a; trans+uction :b; sensory co+in%
:c; line la'elin% :d; a+aptation
%$ ! 'lin+ spot in the retina occurs where
:a; the fo0ea is locate+
:b; %an%lion cells synapse with 'ipolar cells
:c; the optic ner0e attaches to the retina
:d; ro+ cells are clustere+ to form the macula
:e; amacrine cells are locate+
5$ Soun+ wa0es are con0erte+ into mechanical mo0ements 'y the
:a; au+itory ossicles :b; cochlea
:c; o0al win+ow :d; roun+ win+ow
:e; tympanic mem'rane
($ The 'asic receptors in the inner ear are the
:a; utricles :b; saccules
:c; hair cells :d; supportin% cells
:e; ampullae
@$ The retina is the
:a; 0ascular tunic :b; fi'rous tunic
:c; neural tunic :d; a/'/ an+ c are correct
'$ !t sunset/ your 0isual system a+apts to
:a; fo0ea 0ision :b; ro+>'ase+ 0ision
:c; macular 0ision :d; cone>'ase+ 0ision
*$ ! 'etter>than>a0era%e 0isual acuity ratin% is
:a; 20 P 20 :b; 20 P 30 :c; 15 P 20 :d; 20 P 15
6$ The malleus/ incus/ an+ stapes are the tiny 'ones locate+ in the
:a; outer ear :b; mi++le ear
:c; inner ear :d; mem'ranous la'yrinth
=$ Receptors in the saccule an+ utricle pro0i+e sensations of
:a; an%ular acceleration
:b; hearin%
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:c; 0i'ration
:d; %ra0ity an+ linear acceleration
&1$ The or%an of 1orti is locate+ in the VVVVV of the inner ear-
:a; utricle :b; 'ony la'yrinth
:c; 0esti'ule :d; cochlea
&&$ !u+itory information a'out the fre&uency an+ intensity of stimulation is relaye+ to the
1AS o0er the cochlear 'ranch of cranial ner0e
:a; 7$ :b; $7 :c; $777 :d; J
&%$ 3hat are the three types of papillae on the human ton%ueM
&5$ :a; 3hat structures ma,e up the fi'rous tunic of the eyeM
:b; 3hat are the functions of the fi'rous tunicM
&($ 3hat structures ma,e up the 0ascular tunic of the eyeM
&@$ 3hat are the three au+itory ossicles in the mi++le ear/ an+ what are their functionsM
8"$"8 2 Re0iewin% 1oncepts
&'$ Trace the olfactory pathway from the time an o+or reaches the olfactory epithelium until
it reaches its final +estination in the 'rain-
&*$ 3hy are olfactory sensations lon%>lastin% an+ an important part of our memories an+
emotionsM
&6$ 3hat is the usual result if a se'aceous %lan+ of an eyelash or a tarsal %lan+ 'ecomes
infecte+M
&=$ Displacement of sterocilia towar+ the ,inocilium of a hair cell
:a; pro+uces a +epolari.ation of the mem'rane
:b; pro+uces a hyperpolari.ation of the mem'rane
:c; +ecreases the mem'rane permea'ility to so+ium ions
:d; increases the mem'rane permea'ility to potassium ions
:e; +oes not affect the transmem'rane potential of the cell
%1$ Dama%e to the cupula of the lateral semicircular +uct woul+ interfere with the perception
of
:a; the +irection of %ra0itational pull
:b; linear acceleration
248927081.doc 56
:c; hori.ontal rotation of the hea+
:d; 0ertical rotation of the hea+
:e; an%ular rotation of the hea+
%&$ 3hen 0iewin% an o'?ect close to you/ your lens shoul+ 'e more VVVVV-
:a; roun+e+ :b; flattene+
:c; con0e5 :d; lateral
:e; me+ial
8"$"8 3 1ritical Thin,in% an+ 1linical !pplications
%%$ 6ou are at a par, watchin% some +eer 35 feet away from you- ! frien+ taps you on the
shoul+er to as, a &uestion- !s you turn to loo, at your frien+/ who is stan+in% ?ust 2 feet
away/ what chan%es woul+ your eyes un+er%oM
%5$ 6our frien+ Shelly suffers from myopia ;nearsi%hte+ness=- 6ou remem'er from your
physics class thay conca0e lenses cause li%ht wa0es to con0er%e an+ con0e5 lenses sprea+
li%ht wa0es- 3hat type of correcti0e lenses woul+ you su%%est to your frien+M
:a; conca0e lenses
:b; con0e5 lenses
%($ Tom has sur%ery to remo0e polyps ;%rowths= from his sinuses- !fter he heals from the
sur%ery/ he notices that his sense of smell is not as ,een as it was 'efore the sur%ery- 1an you
su%%est a reason for thisM
%@$ 4or a few secon+s after you ri+e the e5press ele0ator from the 20th floor to the %roun+
floor/ you still feel as if you are +escen+in%/ e0en thou%h you ha0e come to a stop- 3hyM
%'$ Wuan has a +isor+er in0ol0in% the saccule an+ the utricle- (e is as,e+ to stan+ with his
feet to%ether an+ arms e5ten+e+ forwar+- !s lon% as he ,eeps his eyes open/ he e5hi'its 0ery
little mo0ement- But when he closes his eyes/ his 'o+y 'e%ins to sway a %reat +eal/
an+ his arms ten+ to +rift in the +irection of the impaire+ 0esti'ular receptors- 3hy +oes this
occurM
,A.DE &*4& The Power 1ontent of Representati0e Soun+s
,ypical +angerous
+ecibel ,ime
248927081.doc 57
De!el E8ample E8posure
0 8owest au+i'le soun+
30 Xuiet li'rary2 soft whisper
0 Xuiet office2 li0in% room2
'e+room away from traffic
50 8i%ht traffic at a +istance2
refri%erator2 %entle 'ree.e
#0 !ir con+itioner at 20 feet2
con0ersation2 sewin%
machine in operation
70 Busy traffic2 noisy restaurant Some +ama%e
if continuous
*0 Su'way2 hea0y city traffic2 @ore than
alarm cloc, at 2 feet2 * hours
factory noise
)0 Truc, traffic2 noisy home 8ess than
appliances2 shop tools2 %as * hours
lawn mower
100 1hain saw2 'oiler shop2 2 hours
pneumatic +rill
120 D(ea0y metalE roc, concert2 7mme+iate
san+'lastin%2 thun+erclap +an%er
near'y
10 Gunshot2 ?et plane 7mme+iate
+an%er
1#0 Roc,et launchin% pa+ (earin% loss
ine0ita'le
: EIG/E &*4& The Olfactory Or%ans- :a; The olfactory or%an on the left si+e of the nasal
septum- :b; !n olfactory receptor is a mo+ifie+ neuron with multiple cilia e5ten+in% from its
free surface-
248927081.doc 58
: EIG/E &*4% Gustatory Receptors- :a; 8an+mar,s an+ receptors on the ton%ue- :b; The
structure an+ representati0e locations of the three types of lin%ual papillae- Taste receptors
are locate+ in taste 'u+s/ which form poc,ets in the epithelium of fun%iform or circum0allate
papillae-
:c; Taste 'u+s in a circum0allate papilla- ! +ia%rammatic 0iew of a taste 'u+/ showin%
receptor ;%ustatory= cells an+ supportin% cells-
: EIG/E &*45 "5ternal 4eatures an+ !ccessory Structures of the "ye- :a; Gross an+
superficial anatomies of the accessory structures-
:b; The or%ani.ation of the lacrimal apparatus- !T8!S9 3c2 12a2 1#a/'
: EIG/E &*4( The Sectional !natomy of the "ye- :a; ! sa%ittal section throu%h the left
eye/ showin% the position of the forni5- :b; @a?or lan+mar,s in the eye- This hori.ontal
section shows the anterior an+ posterior ca0ities an+ the three layers/ or tunics/ in the wall of
the ri%ht eye- :c; ! +etaile+ hori.ontal section of the ri%ht eye- !T8!S9 Plates 12a2 1#a/'
: EIG/E &*4@ The Pupillary @uscles
: EIG/E &*4' The Or%ani.ation of the Retina- :a; The cellular or%ani.ation of the retina-
The photoreceptors are closest to the choroi+/ rather than near the posterior ca0ity ;0itreous
cham'er=- :b; The optic +isc in +ia%rammatic sa%ittal section- :c; ! photo%raph of the retina
as seen throu%h the pupil-
: EIG/E &*4* ! Demonstration of the Presence of a Blin+ Spot- 1lose your left eye an+
stare at the cross with your ri%ht eye/ ,eepin% the cross in the center of your fiel+ of 0ision-
Be%in with the pa%e a few inches away from your eye/ an+ %ra+ually increase the +istance-
The +ot will +isappear when its ima%e falls on the 'lin+ spot/ at your optic +isc- To chec, the
'lin+ spot in your left eye/ close your ri%ht eye an+ repeat the se&uence while you stare at
the +ot-
: EIG/E &*46 The 1irculation of !&ueous (umor- !&ueous humor/ which is secrete+ at
the ciliary 'o+y/ circulates throu%h the posterior an+ anterior cham'ers 'efore it is
rea'sor'e+ throu%h the canal of Schlemm-
: EIG/E &*4= 4actors !ffectin% 4ocal Distance- 8i%ht rays from a source are refracte+
when they reach the lens of the eye- The rays are then focuse+ onto a sin%le focal point-
248927081.doc 59
: EIG/E &*4&1 !ccommo+ation- 4or the eye to form a sharp ima%e/ the focal +istance
must e&ual the +istance 'etween the center of the lens an+ the retina-
: EIG/E &*4&% 7ma%e 4ormation- :aAb; 8i%ht from each portion of an o'?ect is focuse+
on a +ifferent part of the retina- The resultin% ima%e arri0es upsi+e +own :c; an+ 'ac,war+
:d;-
: EIG/E &*4&5 Structure of Ro+s an+ 1ones- :a; The structures of ro+s an+ cones-
Aotice the shapes of their outer se%ments- :b; The structure of a rho+opsin molecule within
the mem'rane of a +isc-
: EIG/E &*4&(
Photoreception
: EIG/E &*4&@ Bleachin% an+ Re%eneration of $isual Pi%ments
: EIG/E &*4&' 1one Types an+ Sensiti0ity to 1olor- ! %raph comparin% the a'sorpti0e
characteristics of 'lue/ %reen/ an+ re+ cones with those of typical ro+s- Aotice that the
sensiti0ities of the ro+s o0erlap those of the cones/ an+ that the three types of cones ha0e
o0erlappin% sensiti0ity cur0es-
: EIG/E &*4&* ! Stan+ar+ Test for 1olor $ision- 7n+i0i+uals lac,in% one or more
populations of cones are una'le to +istin%uish the patterne+ ima%e ;the num'er 12=-
: EIG/E &*4&6 1on0er%ence an+ Gan%lion 1ell 4unction- Photoreceptors are or%ani.e+
in %roups within a recepti0e fiel+2 each %an%lion cell monitors a well>+efine+ portion of that
fiel+- Some %an%lion cells ;on>center neurons/ la'ele+ != respon+ stron%ly to li%ht arri0in% at
the center of their recepti0e fiel+- Others ;off>center neurons/ la'ele+ B= respon+ most
stron%ly to illumination of the e+%es of their recepti0e fiel+-
: EIG/E &*4&= The $isual Pathways- The crosso0er of some ner0e fi'ers occurs at the
optic chiasm- !s a result/ each hemisphere recei0es 0isual information from the me+ial half
of the fiel+ of 0ision of the eye on that si+e/ an+ from the lateral half of the fiel+ of 0ision of
the eye on the opposite si+e- $isual association areas inte%rate this information to +e0elop a
composite picture of the entire fiel+ of 0ision-
: EIG/E &*4%1 The !natomy of the "ar- The 'oun+aries separatin% the three anatomical
re%ions of the ear ;e5ternal/ mi++le/ an+ inner= are in+icate+ 'y the +ashe+ lines-
248927081.doc 60
: EIG/E &*4%& The @i++le "ar- :a; The structures of the mi++le ear- :b; The tympanic
mem'rane an+ au+itory ossicles-
: EIG/E &*4%% The 7nner "ar- :a; The 'ony an+ mem'ranous la'yrinths- !reas of the
mem'ranous la'yrinth containin% sensory receptors ;cristae/ maculae/ an+ the or%an of
1orti= are shown in purple- :b; ! section throu%h one of the semicircular canals/ showin%
the relationship 'etween the 'ony an+ mem'ranous la'yrinths/ an+ the locations of
perilymph an+ en+olymph-
: EIG/E &*4%5 The Semicircular Ducts- :a; !n anterior 0iew of the ri%ht semicircular
+ucts/ the utricle/ an+ the saccule/ showin% the locations of sensory receptors- :b; ! cross
section throu%h the ampulla of a semicircular +uct- :c; "n+olymph mo0ement alon% the
len%th of the +uct mo0es the cupula an+ stimulates the hair cells- :d; ! representati0e hair
cell ;receptor= from the 0esti'ular comple5- Ben+in% the stereocilia towar+ the ,inocilium
+epolari.es the cell an+ stimulates the sensory neuron- Displacement in the opposite
+irection inhi'its the sensory neuron-
: EIG/E &*4%( The Saccule an+ Htricle- :a; The location of the maculae- :b; The
structure of an in+i0i+ual macula- :c; ! +ia%rammatic 0iew of macular function when the
hea+ is hel+ hori.ontally ;S,E) &= an+ then tilte+ 'ac, ;S,E) %=-
of 1orti- :b; Dia%rammatic an+ sectional 0iews of the receptor hair cell comple5 of the
or%an of 1orti- ;8@ Y 1233=
: EIG/E &*4%6 The Aature of Soun+- :a; Soun+ wa0es ;here/ %enerate+ 'y a tunin% for,=
tra0el throu%h the air as pressure wa0es- :b; ! %raph showin% the soun+ ener%y arri0in% at
the tympanic mem'rane- The +istance 'etween wa0e pea,s is the wa0elen%th- The num'er
of wa0es arri0in% each secon+ is the fre&uency/ which we percei0e as pitch- 4re&uencies are
reporte+ in cycles per secon+ ;cps=/ or hert. ;(.=- The amount of ener%y in each wa0e
+etermines the wa0eFs amplitu+e/ or intensity/ which we percei0e as the lou+ness of the
soun+-
: EIG/E &*4%= Soun+ an+ (earin%- Steps in the reception an+ trans+uction of soun+
ener%y-
: EIG/E &*451 4re&uency Discrimination- :a; The fle5i'ility of the 'asilar mem'rane
0aries alon% its len%th/ so pressure wa0es of +ifferent fre&uencies affect +ifferent parts of the
mem'rane- :bA c; The effects of a 0i'ration of the stapes at a fre&uency of #000 (.- 3hen
248927081.doc 61
the stapes mo0es inwar+/ as in part ;'=/ the 'asilar mem'rane +istorts towar+ the roun+
win+ow/ which 'ul%es into the mi++le>ear ca0ity- 3hen the stapes mo0es outwar+/ as in part
;c=/ the 'asilar mem'rane re'oun+s an+ +istorts towar+ the o0al win+ow-
: EIG/E &*45& Pathways for !u+itory Sensations- !u+itory sensations are carrie+ 'y the
cochlear 'ranch of cranial ner0e $777 to the cochlear nuclei of the me+ulla o'lon%ata- 4rom
there/ the information is relaye+ to the inferior colliculus/ a center that +irects a 0ariety of
unconscious motor responses to soun+s- !scen+in% acoustic information %oes to the me+ial
%eniculate nucleus 'efore 'ein% forwar+e+ to the au+itory corte5 of the temporal lo'e-
: EIG/E &*4%@ Pathways for "&uili'rium Sensations
: EIG/E &*4%' The 1ochlea- :a; The structure of the cochlea- :b; Dia%rammatic an+
sectional 0iews of the cochlear spiral-
: EIG/E &*4%* The Or%an of 1orti- :a; ! three>+imensional section of the cochlea/
showin% the compartments/ tectorial mem'rane/ an+ or%an
248927081.doc 62