Biol

ogicalMembr anes
Thephy si
calorganizat
ionandf unct
ioningofal l
biol
ogi calmembr anesdependont heir
consti
tuent
s:li
pids,protei
ns,
andcar bohy drat
es.Youar eal
readyfami l
iarwi tht
hese
molecules
andwithmembr anesthatenclosecellsandor ganel
les.Lipidsestabli
sht hephy sical
i
ntegri
tyofthemembr aneandcr eateabar ri
ertother apidpassageofhy dr ophi
lic
materi
alssuchaswat erandions.Inaddition,t
hephosphol ipi
dbi l
ayerser v
esasal ipi
d
“l
ake”inwhichav ariet
yofproteins“fl
oat”Thisgener aldesignisknownast hefluid
mosaicmodel .I smosai
ti cbecausei tismadeupofmanydi scret
ecomponent s, and
fl
uidbecausethecomponent scanmov efreel
y .

Flui
dMosai cModelAmolecul
armodelfort
hestruct
ureofbi
ologicalmembr
anes
consisti
ng
ofafluidphosphol
i
pidbi
l
ayeri
nwhichsuspendedprotei
nsar
ef r
eet omovei
nthepl
ane
of
thebilay
er.

Thefluidmosaicmodel depi
ctspr
otei
nsasnoncovalentl
yembeddedinthe
phospholi
pidbi
layer
bytheirhy
drophobicregi
ons(ordomains)ort
ether
edt ol
ipi
dsi
nsert
edintothe
membr ane.

Protei
nsmayspanthemembr aneormaybeboundonthesurf
ace.Thei
rhy
drophil
i
c
regi
onsare
exposedtothewat
erycondi
ti
onsoneit
hersi
deofthebi
lay
er.Membraneprot
einshav
e
several
functi
ons,includi
ngmovingmater
ial
sthroughthemembr aneandr
ecei
vingchemical
signals
from thecell
’sexter
nal
envir
onment.Eachmembr anehasasetofprot
einssui
tabl
efor
the
speciali
zedfuncti
onsofthecel
lororganel
lei
tsurrounds.

Thecar bohy dratesassociatedwi thmembr anesar eattachedei thertot helipidsort o

protein
mol ecules.I
ncel lmembr anes, carbohydr
at esar elocatedont heout sideoft hecel l
,
wher ethey
mayi nteractwi t
hsubstancesi ntheexternal envir
onment .Likesomeoft hemembr ane
proteins,
carbohy drat
esar ecruciali
nr ecognizingspeci fi
cmol ecules, suchast hoseont he
surfacesof
adjacentcells.
Althought hefluidmosai cmodel i
slargel
yv ali
dformembr anest r
uct ure,i
tdoesnotsay
much
aboutmembr anecomposi tion.Asy oureadaboutt hevariousmol ecul esinmembr anes
i
nt he
nextsect i
ons, keepinmi ndt hatsomemembr aneshav emor eproteint hanlipids,other
s
are
l
ipid-r
ich,othershav esignif
icantamount sofchol ester
ol orot hersterols,andst il
lothers
arer i
ch
i
ncar bohy drates.

Lipidsf ormt hehy drophobi ccor eoft hemembr ane

Thel ipidsinbi ol
ogical membr anesar eusually*phospholipids.thatsomecompounds
arehy drophil
ic(“wat er-
lovi
ng”)andot hersarehy dr
ophobic( “waterhat
ing”),
Aphosphol i
pidmol ecul ehasr egionsofbot hkinds:
1.Hy drophil
icregions:Thephosphat e-contai
ning“head”oft hephosphol i
pidis
electricall
y
char gedandt hereforeassoci ateswi thpolarwatermol ecules.
2.Hy drophobicregions:Thel ong, nonpolarfatt
yacid“ t
ail
s”oft hephosphol i
pid
associ atewi th
othernonpol armat er i
als;
theydonotdi ssol
veinwat erorassoci atewithhydrophi
l
ic
subst ances.
Thechemi calproper ti
esofphosphol ipidsaresuchthatwhenphosphol i
pidscoexi
st
withwat er,
theyf orm abi l
ayer, withthefat t
yaci d“tail
s”ofthetwol ayersinteract
ingwitheach
otherandt hepol ar“heads”f acingt heout si
deaqueousenv ironment

Thet hicknessofabiol
ogical
membr aneisabout8nanomet er
s(0.008μm),whi
chis
twi
cet helengthofatypi
calphosphol
ipi
d—anotherindi
cat
ionthatthemembrane
consi
st sofalipi
dbil
ayer
.
Allbi
ologicalmembraneshaveasimilarst
ruct
ure,butt
heydiff
erintheki
ndsofprot
eins
and
l
ipidstheycont ai
n.Membr anesf r
om di
fferentcel
lsororganel
lesmaydi ffergreat
lyi
n
theirl
i
pid
composi ti
on.Phospholipi
dscandi ff
eri
nt ermsoff at
tyaci
dchainl engt
h( numberof
carbon
atoms), degreeofunsatur
ation(numberofdoubl ebonds)inthefattyaci
ds, andthe
polargroups
present.
Thesat uratedchainsal
lowcl osepacki
ngoff att
yacidsinthebil
ayer,wher easthe
“ki
nks”inunsat ur
at edf
att
yaci dsmakeforal essdense,moref l
uidpacking.

Upt o25per centofthel i

pidcont entofanani mal cell
’scell
membr anemaybethe
steroi
d
cholester ol
.Cholesterolpref
erent i
allyassociateswi thsaturat
edf att
yaci
ds.When
present, cholest
erolisimpor t
antf ormembr aneint egri
ty;t
hechol est
erol
inyour
membr anesi snothazar doust oy ourhealth.
Thef at
tyaci dsoft hephosphol ipidsmaket hehy drophobicinteri
orofthemembrane
somewhatf l
uid—aboutasf l
uidasl i
ghtweightoli
v eoil.Thi
sf l
uidi
typermit
ssome
molecul est o
mov elat eral
lywithi
nt heplaneoft hemembr ane.

Agivenphospholi
pidmolecul
eint hecellmembr anecant ravelfr
om oneendoft hecell
tothe
otherinali
ttl
emorethanasecond!Howev er,aphosphol i
pidmoleculeinonehalfofthe
bil
ayer
i
sunl i
kelyt
ospontaneousl
yfli
pov ert
ot heotherside.Forthattohappen, t
hepolarpart
ofthe
moleculewouldhavetomov ethroughthehy drophobi
ci nt
eriorofthemembr ane.Since
spontaneousphopchol
ipi
dfli
p-f
lopsarer ar
e,theinnerandout erhalvesofthebil
ayer
maybe
quit
edi f
fer
entint
hekindsofphospholipidstheycontain.

Membr anef l
uidit
yisaf f
ect edbysev er
al f
actors,t
woofwhi char eparti
cularl
yimport
ant:
1.Lipidcomposi t
ion:Chol esterolandlong-chain,saturatedf att
yaci dspackt i
ght
ly
besideone
another,wi t
hlitt
leroom f ormov ement .Thi
sclosepacki ngresultsinless-fl
uid
membr anes.A
membr anewi t
hshor t
er-
chai nf att
yacids,unsat ur
atedf attyacids,orlesscholest
eroli
s
mor e
fl
uid.
2.Temper atur
e:Becausemol eculesmov emor eslowlyandf lui
ditydecreasesat
reduced
temper atures,cel
lularprocessest hattakeplacewi t
hint hemembr anemaysl owdown
orstop
undercol dcondi t
ionsinor gani smst hatcannotkeept hei rbodieswar m.Toaddr essthi
s
problem, insomeor ganismst helipi
dcomposi t
ionofthei rmembr aneschangeswhen
theyget
cold,r
eplaci
ngsat
urat
edwithunsat
uratedfat
tyaci
dsandusingf
att
yacidswithshor
ter
tai
ls.
Thesechangesplayarol
einthesur
vivalofpl
ants,
bact
eri
a,andhi
bernat
inganimal
s
during
thewinter.
Allbiologi
calmembranescontainprotei
ns.Typicall
y,cell
membr aneshav e1protei
n
mol ecule
forev ery25l
ipi
dmol ecul
es.Thisrat
iovari
esdependi ngonmembr anefuncti
on.I
nt he
i
nner
membr aneofthemitochondri
on,whichisspecial
izedforenergyprocessing,t
her
ei s1
protein
forev ery15l
ipi
ds.Howev er
,my el
i
n—amodi fi
edcellmembr anethatenclosesporti
ons
ofsome
neur ons(ner
vecell
s)andactsasanel ectr
icali
nsulator—hasonly1proteinforev
ery70
l
ipids.

Membr anepr otei nsar ev er ydi ver se.I nf act ,

aboutone- four t
hoft heprotein-coding
genesi n
theeukar yot i
cgenomeencodemembr anepr ot eins.Ther ear et wogener al typesof
membr ane
prot eins:i
nt egral pr oteinsandper ipher al protei ns.
Integr almembr anepr ot einsar eatl eastpar tlyembeddedi nt hephosphol i
pidbi l
ayer.
Likephosphol i
pi ds, t
hesepr otei nshav ebot hhy drophi l
i
candhy drophobicr egions
(domai ns).
i
nt egr almembr anepr ot einsPr oteinst hatar eatl eastpar tiall
yembeddedi ntheplasma
membr ane.( Cont rastwi thper ipher almembr anepr otei
ns. )
Hy dr ophil
icdomai ns:St ret chesofami noaci dswi thhy drophi licsidechai ns(Rgr oups)
givecer t
ainr egionsoft hepr ot ei napol archar act er.Thesehy drophili
cdomai nsinteract
withwat erandst ickouti nt ot heaqueousenv i
r onmenti nsi deorout si
det hecel l
.
2.Hy drophobi cdomai ns:St ret chesofami noaci dswi thhy drophobi csidechai nsgive
other
regionsoft hepr ot einanonpol archar act er.Thesedomai nsi nteractwitht hef att
yacids
i
nt he
i
nt erioroft hephosphol ipidbi lay er ,awayf r
om wat er.Somepr oteinscont aincov al
ently
l
inked
l
ipidssuchasf at tyaci dchai nst hatdot hej obofi nser t
ingi ntot hemembr ane( i
nstead
oft he
prot einhav inghy dr ophobi cami noaci dr egions) .
Per ipheralmembr anepr oteinsl ackexposedhy drophobi cgr oupsandar enot
embeddedi n
thebi l
ayer.Instead, t
heyhav epol arorchar gedr egi onst hatint eractwithexposedpar t
s
of
i
nt egr almembr anepr otei ns, orwi tht hepol arheadsofphosphol ipi
dmol ecul es;
Aspeci alpr eparat ionmet hodf orel ect ronmi cr oscopy ,calledf reeze-f
racturing,reveals
prot einsthatar eembeddedi nt hephosphol i
pi dbi layersofcel l
ularmembr anes.
Whent hetwol i
pi dl eaflets( orl ay er s)t hatmakeupt hebi layerar esepar ated, t
he
prot
einscan
beseenasbumpst hatprotr
udefr
om t
heint
eri
orofeachmembr ane.Thebumpsar
e
not
observedwhenar
ti
fici
albil
ayer
sofpur
eli
pidar
efreeze-
fr
act
ured.

Membr anepr oteinsandl ipidsgener al lyint eractonl ynoncov al

ent ly.Thepol arendsof
prot einscani nteractwi tht hepol arendsofl i
pids,andt henonpol arregionsofbot h
mol ecul es
cani nter acthy drophobi call
y .Asment ionedabov e,howev er,somemembr anepr ot eins
hav e
hy drophobi cl i
pidcomponent scov al ent lyat tachedt ot hem thatal l
owt hesepr otei nst o
tether
themsel vest ot hephosphol ipidbi lay er .
Pr oteinsar easy mmet ri
cal l
ydi str i
but edont hei nnerandout ersur f
acesofmembr anes.
An
i
nt egr al pr oteint hatext endsal l thewayt hr ought hephosphol i
pidbi layerandpr ot rudes
onbot h
sidesi sknownasat r ansmembr anepr otein.I naddi t
iontooneormor etransmembr ane
domai nst hatext endt hrought hebi lay er,suchapr oteinmayhav edomai nswi t
hot her
speci fic
funct ionsont hei nnerandout ersi desoft hemembr ane.Per i
pher al membr anepr ot eins,
by
cont rast , arel ocat edononesi deoft hemembr aneort heot her.Thi sasy mmet rical
arrangement
ofmembr anepr oteinsgi vest het wosur facesoft hemembr anedi f
ferentpr oper ties.As
youwi l
l
soonsee, thesedi fferenceshav egr eatf unct ionalsignifi
cance.
transmembr anepr ot einAni ntegr al membr anepr oteinthatspanst hephosphol ipi d
bilayer .
transmembr anedomai nApr oteinr egi onr ichi nhy drophobi cami noaci dst hatspans
the
phosphol i
pi dbilay er.
Likel ipids, somemembr anepr oteinsmov ear oundr elativel
yf reelywi t
hint he
phosphol i
pi d
bilayer .Exper i
ment st hati nv olvet het echni queofcel lfusioni l
lust r
atethismi grat i
on
dramat i
cal ly.
Whent wocel l
sar ef used, asi ngl econt i
nuousmembr anef ormsandsur r
oundsbot h
cel l
s, and
somepr ot einsf rom eachcel ldi stribut ethemsel v
esuni f
orml yar oundt hismembr ane

Althoughsomepr oteinsarefr
eetomi gr
atei
nthemembrane,other
sarenot
,butrat
her
appeart obe“ anchored”toaspecif
icregi
onofthemembrane.Thesemembrane
regionsar el
ike
anencl osureofanimal satthezoo:t
heanimalsar
efr
eetomov earoundwit
hint
he
fencedar ea
butnotout sidei t.Anexampl eist hepr oteinint hecel lmembr aneofamuscl ecel lthat
recogni zesachemi cal signal fr
om aneur on.Thi sprot einisnor mal l
yf oundonl yatt he
speci fi
c
regi onwher et heneur onmeet sthemuscl ecel l
.Pr oteinsi nsi
det hecel lcanr est ri
ctt he
mov ementofpr oteinswi thinamembr ane.Thecy t
oskel etonhascomponent sjust
bel owt he
i
nnerf aceoft hemembr anet hatar eattachedt omembr anepr ot einspr otrudingi ntot he
cy toplasm.Thest abi l
it
yoft hecy toskel etalcomponent smayt husr estrictmov ementof
attached
membr anepr oteins.
Membr anesar econst ant l
ychangi ng
Membr anesi neukar yot i
ccel lsareconst antlyformi ng, transformi ngf rom onet ypet o
anot her,
fusi ngwi t
honeanot her ,andbr eaki ngdown.f r
agment sofmembr anemov e,int hef orm
ofv esicles,from t heendopl asmi cr eti
culum ( ER)t otheGol giappar atus, andf rom t he
Gol gi appar atust ot hecel lmembr ane.Secondar ylysosomesf orm whenpr i
mar y
l
y sosomesf r
om t heGol gi apparatusf usewi t
hphagosomesf rom t hecel lmembr ane.
Becauseal l membr anesappearsi mi l
arundert heel ectronmi croscope, andbecause
they
i
nt er conv ertreadi l
y, wemi ghtexpectal lsubcellularmembr anest obechemi cal l
y
i
dent i
cal.
Howev er,thati snott hecase: therear emaj orchemi cal diff
erencesamongt he
membr anesof
ev enasi ngleeukar y oti
ccel l
.Membr anesar echangedchemi callywhent heyf orm par ts
of
cer tainor ganel l
es.I nt heGol giappar atus, f
orexampl e,themembr anesoft heci sf ace
closel y
resembl ethoseoft heERi nchemi calcomposi ti
on, butt hoseoft hetransf acear emor e
simi lar
tot hecel lmembr ane.

Cellmembr anecar bohy dr atesar erecognitionsi t

es
Inaddi ti
ont ol i
pidsandpr oteins,
thecellmembr anecont ainscarbohydrates.
Thecar bohy dratesar elocat edont heout ersur f
aceoft hecellmembr aneandser v
eas
recogni ti
onsi tesforot hercel lsandmol ecul es.
Membr ane- associatedcar bohy dr
atesmaybecov al
ent l
ybondedt oli
pidsort oprotei
ns:
Agl ycolipidconsi st
sofacar bohydratecov alentl
ybondedt oal i
pid.Extendingoutf r
om
the
cellsurface, thecarbohy dratemayser veasar ecognitionsi gnalf
orinteract
ions
betweencel ls.
Forexampl e, thecarbohy dratesonsomegl ycoli
pidschangewhencel lsbecome
cancer ous.
Thischangemayal lowwhi t
ebl oodcellstot argetcancercel l
sfordestructi
on.
Agl ycopr oteinconsi stsofoneormor eshor tcarbohy dratechainscov alent
lybondedt o
a
protein.Theboundcar bohy dratesareoligosacchar ides, usuall
ynotexceedi ng15
monosacchar ideunitsinlength.
Apr ot eoglycani samor eheav i
lygl
ycosy l
atedpr otei
n: i
thasmor ecarbohy drate
mol ecul esat tachedt oit,andthecarbohy dratechainsar eoftenlongerthantheyar ein
gly
copr otei
ns.Thecar bohy dr
atesofglycopr ot
einsandpr ot
eoglycansoftenf uncti
oni n
cel
l recogni t
ionandadhesi on.
glycol i
pi dAl i
pi dtowhi chsugar sar
eattached.
glycopr oteinApr ot
eintowhi chsugarsar eat t
ached.
The“ alphabet ”ofmonosacchar idesont heout ersurfacesofmembr anescangener ate
alar ge
div
er sit yofmessages.Recal lt
hatmonosacchar idesar esimplecarbohydrates, of
ten
cont ainingf iveorsi xcarbonsinar i
ngst ructure,whichcanbondwi thoneanot herin
variousConf i
gur ati
ons.Theymayf ormlinearorbr anchedol i
gosaccharideswi thmany
dif
fer entt hree-dimensional shapes.Anol i
gosacchar ideofaspeci fi
cshapeononecel l
canbi nd
toacompl ement aryshapeonanadj acentcel l
.Thisbi ndi
ngist hebasisofcel l
–cell
adhesi on.