GuideQuestionsforpp.

164172

1. Listanddescribethefunctionsofmembranes.
(1)Membranesdefinetheboundariesofacellandactaspermeabilitybarrierssothatthe
appropriatesubstancesarekeptinsideandtheunneeded/harmfulonesarekeptout.Additionally,
intracellularmembranescompartmentalizesubstanceswithinthecellforperformingspecific
functionslikeinthemitochondria.(2)Membraneshavespecificfunctionsbasedonthe
molecules(proteins)embeddedinthem.Enzymescanserveasmarkersforspecific
membranesliketheERorGolgi.(3)Membranesregulatethetransportofsolutesintoandoutof
thecell.Theyareabletodothisthroughpassive(alongtheconcentrationgradient)andactive
(againstthegradient)transport.(4)Membranesdetectandtransmitelectricalandchemical
signals(throughsignaltransductionpathways)tocausecertaincellularbehavior(likethe
productionofcertainproteins)ortocausetheacceptanceofcertainmolecules.(5)Membranes
mediatecelltocellcommunicationthroughchannelslikeplasmodesmata(forplants)andgap
junctions(foranimals).
2. Identifythescientiststhatmadecontributionstothestudyofmembranestructure.For
each,describethesignificanceoftheirstudiesindeterminingthestructureofbiological
membranes.
Overton&Langmuir(1890s):Overtonobservedacorrelationbetweenlipophiliccells
andtheireaseinenteringamembrane.Healsoknewabouttheprominenceofsterolsand
phospholipidsinmembranestructure.Langmuirdissolvedphopholipidsinbenzeneandlayered
themonawatersurface,discoveringthelipidmonolayer.Hehypothesizedthatthehydrophilic
endsofthelayerfacedthewaterandthehydrophobicendsprotrudedawayfromthewater.
GorterandGrendel(1925):experimentedwithredbloodcellsandextractedthelipids
fromthemtodeterminethenumberoflipidlayersinthecellscoat.Theyfoundthatthelipids
spreadoutasamonolayeronwaterhadabouttwicethesurfaceareaofthecell,sothey
concludedthatthecoatwasaphospholipidbilayer.Theyfounditwasthermodynamically
favorableforthehydrocarbontailsofthephopholipidstobothfaceinwardandforthephophates
tofaceoutward.
DavsonandDanielli(1935):Asimplelipidbilayerdidnotexplainmanyofthespecial
propertiesofthemembraneslikehowhydrophilicsolutesmovedmorereadilyintoandoutofthe
cellthancanbeexplainedbybilayerpermeability.DavsonandDanielliproposedthatproteinsin
themembranesbroughtabouttheseproperties,describingitasasandwichmodelwhere
proteinlayerssandwichedthelipidbilayer.Thistheorywasmodifiedbyaddingpolarpores
wherehydrophilicproteinscanpenetratethroughanotherwisehydrophobicbilayer.
Robertson(1950s):Withtheadventofelectronmicroscopy,cellbiologistsnoticedthat
everycellhadamembraneandfoundthatadjacentcellshadatrilaminarstructure(membrane

ofonecellfollowedbyintercellularspacefollowedbythemembraneofthenextcell).Robertson
proposedtheideaofaunitmembranethatallcellmembranessharethesameunderlying
structure.
SingerandNicolson(1972):proposedthefluidmosaicmodelwhereproteinswere
embeddedintothemembrane.Themosaicpartofthemodeldescribeshowvariousproteinsare
embeddedatdifferentdepthsinthemembranetocarryouttheirspecificfunction(tofacilitate
diffusionoflargemoleculesforexample).Thefluidpartofthemodeldescribeshowthelipid
componentsareinconstantmotionandtheproteinsareabletomovelaterallywithinthe
membrane.
3. Whatarethemajorproblemswiththeearlymembranemodels?
Electronmicroscopyfoundthatthemembranesweretoothintoaccommodateforthe
DavsonandDaniellimodel.Themodelalsodidnotaccountforthevariabilityinchemical
compositionandproteintolipidratioindifferenttypesofmembranes.Italsodidnottakeinto
accountphospholipiddegradationbyphospholipases,whichwasfoundtobeaverycommon
occurrence,asthebilayerwouldbeprotectedbytheproteinlayers.Manyproteinswerefoundto
behydrophobicaswell,sotheyhadtobeembeddedinthehydrophobicinteriorofthe
membrane.TheModelalsodidnotaccountforthefluidityofthemembrane,asstackedlipids
wouldformaveryrigidstructure.

GuideQuestionsforpp.171180

1. Listthemajorclassesofmembranelipidsandgiveexamplesforeachclass.
a. Phospholipids
i. Prominentcomponentofmembranes
ii. Examples:phosphoglyceridesandsphingolipids
b. Glycolipids
i. Formedbytheadditionofcarbohydrategroupstolipids.Someare
glycerolbased,butmostarederivativesofsphingosineandaretherefore
calledglycosphingolipids
ii. Examples:cerebrosides,gangliosides
c. Sterols
i. Membranescontainsignificantamountsofsterol
ii. Examples:cholesterol,phytosterols
2. Whyisitsaidthatmembranesareasymmetricwithrespecttomembranelipids?
a. Therearedifferentproportionsoflipidsineachofthetwomonolayers.
Asymmetryismaintainedbecauseofthemovementofthehydrophilicheads
throughthehydrophobicinteriorofthemembrane.Thisiscalledtransverse
diffusion.
3. Howaresterolsorientedinthecellmembrane?

4.

5.

6.
7.

8.

a. Itorientsitselfinthelayerwithitssinglehydroxylgroupclosetothepolarhead
groupofaneighboringphospholipidmolecule,whereitcanformahydrogenbond
withtheoxygenoftheesterbondbetweentheglycerolbackboneandthefatty
acidofthesterol.
Howdoescholesterolregulatemembranefluidity?
a. Iteffectivelypreventsthehydrocarbonchainsofphospholipidsfromfitting
togetherasthetemperaturedecreases,thusincreasingfluidity.
Whatcharacteristicsoffattyacidsdeterminethefluidityofmembranes?
a. Themorecompactandclosertogethertheyare,themorerigidandlessfluida
membraneis.Thefurthertheyareapartandtheirinabilitytobindtootherlipids
wouldgiverisetoafluidmembrane.
Howdoestemperatureaffectthefluidityofmembranes?
a. Decreaseinthemembranefluidityatgreatertemperatures.
Howdopoikilothermscompensatefortemperaturechanges?
a. Astemperaturesincrease,thelipidbilayersbecomesofluidthattheynolonger
serveasapermeabilitybarrier.However,mostpoikilothermscancompensatefor
thetemperaturechangesbyalteringthelipidcompositionoftheirmembranes.
Describehomeoviscousadaptationincoldtemperatures.
a. Itallowsorganismstochangetheproportionsandconcentrationsofdifferent
lipidsinthebilayersothatthechangeintemperaturedoesnotaffectfluidityor
permeability.

GuideQuestionsforTheMosaicPartoftheModel(p.181191)

1.Brieflydescribethefreezefracturetechniqueusedtostudymembranes.
a.Thefreezefracturetechniqueisperformedbyfirstquicklyfreezingalipidbilayeror
membrane.Thenthelayerissubjecttosharpblowsfromamicrotomeknife.Thisthensplitsthe
specimenintothetheinnerandoutermonolayerrevealingtheinnersurfaceofeach.Thisallow
researchertoseetheproteinssuspendedinthemembranes.
2.Whatarethetwoclassesofintegralmembraneproteins?Howaretheydifferent?
a.IntegralMonotopicproteinsAclassofintegralmembraneproteinsthatprotrudeononly
onesideofthebilayer
b.TransmembraneproteinsThesemakeupmostoftheintegralmembraneprotiensand
theyspanthemembranehavinghydrophilicregionsonbothprotrudingfrombothsidesofthe
membrane
3.Describephysicalpropertiesofsinglepassandmultipassmembraneproteins.
a.Singlepassmembraneproteinshavejustonetransmembranesegmentwitha
hydrophiliccarboxylterminusextendingoutofthemembraneononesideandahydrophilic
aminoterminusprotrudingontheotherside
b. Multipassmembraneproteinshaveseveraltransmembranesegmentsrangingfrom3
to20ormoresuchsegments

4.Whatdoesahydrophobicity(hydropathy)plotshow?
a.Ahydrophobicityplotshowsthenumberandpositionsoftransmembranesegmentsin
aprotein
5.Whatarephysicalpropertiesofperipheralmembraneproteins?
a.Peripheralmembraneproteinsassociatewiththemembranesurfacesthroughweak
electrostaticforcesandhydrogenbondingwiththehydrophilicprotonsofintegralproteinsand
perhapswithpolarheadgroupsofmembranelipidsthatextendoutofthemembrane.Peripheral
proteinsaremuchmorereadilyremovedfromthemembranethatintegralproteinsandcan
usuallybeextractedbychangingPH.

GuideQuestionspp.192193ManyMembraneProteinsareGlycosylated

1.DifferentiatebetweenNlinkedglycosylationandOlinkedglycosylation.
a.Glycosylationistheadditionofacarbohydratesidechaintoaprotein.Nlinked
glycosylationisthelinkageofthatcarbohydrategrouptoaaminogroupandOlinked
glycosylationisthelinkageofthatcarbohydrategrouptotheoxygenofahydroxylgroup.
is
2.Whatisthenatureofthecarbohydratechainsonthemembranesurface?
a.InbothNlinkedandOlinkedthecarbohydrategroupterminatesinnegativelycharged
sialicacid.Thiscausestheerythrocytestorepeleachothertoreducebloodviscosity.
3.Whatisthefunctionofcarbohydrategroupsonthecellsurface?
a.Glycoproteinsarefoundmostlyintheplasmamembranewheretheyplananimportant
roleincellcellrecognition.Thisisbecausetheglycoproteinsarepositionedinsuchawaythat
thecarbohydrategroupsstickoutoftheexternalmembrane.