BCH204

DigestionandAbsorptionofLipids

Digestionenzymeswhichregulateandactasacatalysttospeeduptherateofchemicalreactions.Lipids,o
ftenreferredtoasfatshydrophobicordonotmixwithwater.Lipiddigestionandabsorptionarecomplexp
rocessesandinvolvedifferentenzymes.

LIPIDDIGESTION:PHYSIOCHEMICALANDENZYMATICSTEPS.

Dietarylipidsconsumedbymonogastricsarepredominantlytriacylglycerols.Theutilizationoffatinani
malsandhumansrequiresthedigestionandabsorptionofdietaryfatsourcesduringthepassagethroughth
egastrointestinaltract(GIT)
(TheGItractconsistsoftheoralcavity,pharynx,esophagus,stomach,smallintestine,largeintestine,andanalc
anal.).Gastrointestinallipiddigestionconsistsofseveralsequentialstepsthatincludephysicochemicala
ndenzymaticevents.Sincefatisinsolubleinwater,emulsificationisrequiredforthedigestionoffatinana
queousmediumasitexistsintheGIT.Thisemulsificationleadstotheorganizationofdietarylipidsinthefo
rmofdropletsintheaqueousdigestivesystem

Digestionoflipidsbeginsinthemouththenmovestothestomachandsmallintestine.Althoughfatsaregen
erallythoughtofasharmful,thebodyneedsacertainamountoffatinordertofunction.Fatsplayseveralimp
ortantrolesinthebody.

 Animportantsourceofessentialfattyacids-thesearefattyacidsthebodycannotmakeonitsown
 VitaminssuchasA,D,andEarestoredinfattytissue
 Storeenergy,insulate,andprotectvitalorgans
 Actaschemicalmessengers
 Playaroleinchemicalreactionsinvolvinggrowth,immunefunction,andmetabolism
 Theyformtheplasmamembraneofcells

Triglycerides

StepsofFatDigestionandAbsorption
Whenwaterandoilaremixed,theoil(fat)separates.Thedigestivetractcanbedescribedasawateryenviro
nmentcausinglipidstohavesimilarbehaviorwhenmovingthroughthetract.Whenlipidsentertheenviro
nment,theyformlargedropletsoffat.Duringlipiddigestion,theselargedropletsmustbebrokendownand
digestedbyspecialenzymes.

Lipiddigestioncanbebrokenintothreemainsteps:

 Lipiddigestioninthemouth
 Lipiddigestioninthestomach
 Lipiddigestioninthesmallintestine

Boththemouthandstomachplayaroleinlipiddigestion,butthemajorityoflipiddigestionoccursinthesm
allintestine.Oncelipidsarebrokendownthroughthesesteps,theyarethencirculatedandtransportedarou
ndthebody.Sincebloodislargelycomposedofwater,lipidsrequirespecialtransportmechanismstomov
ethemtotheirdestinations
 1. Lipiddigestioninthemouth

Duringchewing,foodparticlesaremechanicallybrokendownintosmalleronesandmixedwithsaliva.B
yproducinglinguallipase,thetongue’scellsbegintodigesttriglycerides,breakingthemdownintoindivi
dualfattyacids.
 2. Lipiddigestioninthestomach

Mixingandchurninginthestomachdispersesfatmoleculesandfoodparticles.Aspartoftheenzymaticdi
gestionoftriglycerides,thestomachproducesanotherlipasecalledgastriclipase.Whenswallowedwithf
oodandsaliva,thelinguallipasestaysactiveinthestomach.However,bothoftheselipasestogetherdonot
playasignificantroleinfatdigestion,andthemajorityofenzymaticdigestiontakesplaceinthesmallintest
ine.

3. Lipiddigestioninthesmallintestine

Mostofthedietarylipidsremainundigestedinthesmallintestineafterthestomachcontentsareswallowed
.Aspartofthedigestiveprocess,thelivermakesandstoresbile,whichenterstheduodenum,thefirstsectio
nofthesmallintestine.Bileisbothhydrophobicandhydrophilic,sotheyaredrawntofatsandwater.There
sultisthattheyworkasgoodemulsifiers,i.e.theybreakdownfatglobulesintosmallerdroplets.Emulsific
ationincreasesthesurfaceareaonwhichdigestionenzymescanactonlipids.

GoingtotheBloodstream

Asstomachcontentsenterthesmallintestine,thedigestivesystemsetsouttomanageasmallhurdle,namel
y,tocombinetheseparatedfatswithitsownwateryfluids.Thesolutiontothishurdleisbile.Bilecontainsbi
lesalts,lecithin,andsubstancesderivedfromcholesterolsoitactsasanemulsifier.Itattractsandholdsont
ofatwhileitissimultaneouslyattractedtoandheldontobywater.Emulsificationincreasesthesurfaceare
aoflipidsoverathousand-fold,makingthemmoreaccessibletothedigestiveenzymes.

Oncethestomachcontentshavebeenemulsified,fat-
breakingenzymesworkonthetriglyceridesanddiglyceridestoseverfattyacidsfromtheirglycerolfound
ations.Aspancreaticlipaseentersthesmallintestine,itbreaksdownthefatsintofreefattyacidsandmonog
lycerides.Yetagain,anotherhurdlepresentsitself.Howwillthefatspassthroughthewaterylayerofmucu
sthatcoatstheabsorptiveliningofthedigestivetract?
Asbefore,theanswerisbile.Bilesaltsenvelopthefattyacidsandmonoglyceridestoformmicelles.Micell
eshaveafattyacidcorewithawater-
solubleexterior.Thisallowsefficienttransportationtotheintestinalmicrovillus.Here,thefatcomponent
sarereleasedanddisseminatedintothecellsofthedigestivetractlining.

MicelleFormation
Chylomicrons

Justaslipidsrequirespecialhandlinginthedigestivetracttomovewithinawater-
basedenvironment,theyrequiresimilarhandlingtotravelinthebloodstream.Insidetheintestinalcells,th
emonoglyceridesandfattyacidsreassemblethemselvesintotriglycerides.Triglycerides,cholesterol,a
ndphospholipidsformlipoproteinswhenjoinedwithaproteincarrier.Lipoproteinshaveaninnercoretha
tisprimarilymadeupoftriglyceridesandcholesterolesters(acholesterolesterisacholesterollinkedtoafa
ttyacid).Theouterenvelopeismadeofphospholipidsinterspersedwithproteinsandcholesterol.Togethe
rtheyformachylomicron,whichisalargelipoproteinthatnowentersthelymphaticsystemandwillsoonb
ereleasedintothebloodstreamviathejugularveinintheneck.Chylomicronstransportfoodfatsperfectlyt
hroughthebody’swater-
basedenvironmenttospecificdestinationssuchastheliverandotherbodytissues.

Cholesterolsarepoorlyabsorbedwhencomparedtophospholipidsandtriglycerides.Cholesterolabsorp
tionisaidedbyanincreaseindietaryfatcomponentsandishinderedbyhighfibercontent.Thisisthereason
thatahighintakeoffiberisrecommendedtodecreasebloodcholesterol.Foodshighinfibersuchasfreshfr
uits,vegetables,andoatscanbindbilesaltsandcholesterol,preventingtheirabsorptionandcarryingthem
outofthecolon.

Iffatsarenotabsorbedproperlyasisseeninsomemedicalconditions,aperson’sstoolwillcontainhigham
ountsoffat.Iffatmalabsorptionpersiststheconditionisknownassteatorrhea.Steatorrheacanresultfrom
diseasesthataffectabsorption,suchasCrohn’sdiseaseandcysticfibrosis.
BiochemistryofDigestionandAbsorptionofLipids

Assimilationoflipidsbeginswiththeactionofbileacids,whichpromoteemulsification.Bileacidsareder
ivedfromcholesterolandhavebothhydrophobicandhydrophilicdomains.Ifexposedtoalipidaggregate
containingalargefractionoftriglycerides,thehydrophobicpartofbileacidswillintercalateintothelipida
ndthehydrophilicpartwillremainonthesurface.Bileacids,whencoatedonaggregatesordroplets,aidinb
reakingthemdownintosmallerpieces.

Thehydrolysisoftriglyceridesformsmonoglyceridesandfreefattyacids.Inadditiontoclippingthefatty
acidsatpositions1and3ofthetriglyceride,thisenzymealsoleavesa2-monoglycerideafterdoingso.

Inthesmallintestine,lipaseisveryabundantshortlyafterameal,butitonlyhasthecapabilityofactingonth
esurfaceoftriglyceridedroplets.Asmallerdropletsizehasmoresurfaceareacomparedtolargerdropletsi
ze,enablingmorelipasemoleculestogettoworkonagivenvolumeoflipid.

Byactingonlipase,monoglyceridesandfattyacidsarereleasedandremainboundtobileacids.Thesefatty
acidsthenjoinwithotherlipidstoformmicelles.Amicelleisasmallaggregateofmixedbileacidsandlipid
s(4-
8nmdiameter)containedwithintheintake.Uponmixingtheingesta,micellesbumpintobrushborderson
smallintestinalenterocytes,allowinglipidsincludingmonoglyceridesandfattyacids,toenterepithelial
cells.

AbsorptionofLipids

Asaresultoffatdigestion,fat-
solublevitamins,monoglycerides,glycerol,cholesterol,andmonoglyceridesneedtoenterthecirculator
ysystem,wheretheycanbeusedbycellsaroundthebody.Thisprocessisalsoaidedbybile.

 Bilesaltsbondwiththeproductsoffatdigestiontoformmicelles,whichhelpabsorbedfatsreachth
eintestinalmicrovilli.Bydiffusingacrosstheintestinalcellmembrane,fatdigestionproductsare
transformedintobilesalts,whicharereusedtodissolvefatandformmicelles
  Thebloodstreamcandirectlyabsorbshort-andmedium-
chainfattyacidsandglyceroloncetheselipidshaveenteredtheintestinalcells,butlargerlipidssuc
hasmonoglycerides,cholesterol,andfat-
solublevitaminsrequiremoreassistancetoabsorbandtransferintothebloodstream

 Withcholesterolandfat-solublevitamins,long-
chainfattyacidsandmonoglyceridesareconvertedintotriglycerideswithintheintestinalcells,a
ndthesetriglyceridesarethenincorporatedintotransportcarrierscalledchylomicrons

 Thechylomicronsarelipoproteins—
largestructuresmadeupoftriglycerideandcholesterolinthecoreandphospholipidsintheouterm
embraneinterspersedwiththeproteins.Sincetheyarecoveredbythisoutermembrane,theycanm
oveinthebody’saqueousenvironmentofbloodandlymph

 Fromthesmallintestine,chylomicronstravelintothelymphaticsystembeforereachingtheblood
stream

 Dietaryfats,afterabsorption,appearinthecirculationasachylomicron

Bileandpancreaticjuiceprovidepancreaticlipase,bilesalts,andcolipase,whichfunctioncooperativelyt
oensuretheefficiencyoflipiddigestionandabsorption.Theimportanceofbiletotheefficiencyofthesepr
ocessesisindicatedbythedecreasedrateoflipidabsorptioninhumanswithbilefistulas.Thegreatlyreduc
edconcentrationofbileacidintheduodenumofsuchindividualssuggeststhatbilesalts,althoughpossibly
notabsolutelynecessaryfordigestion,areessentialforthecompleteabsorptionofdietaryfats.Elevatedc
oncentrationsofbilesaltshavebeenshowntoinhibitpancreaticlipaseactivityintheduodenum.Suchinhi
bitionisoffset,however,bycolipase,whichhasbeenshowninvitrotorestorepancreaticlipaseactivityun
dersuchcircumstances.Theimportanceofcolipaseinthedigestionoffatwasindicatedinaclinicalreport
ofsteatorrheaintwobrotherswithacongenitalabsenceofcolipase,whichindicatedthatthesteatorrheade
creasedwiththeadministrationofcolipase.Ithasalsobeendemonstratedincolipase-
deficientmice,which,whenplacedonahigh-
fatdiet,developedsteatorrheathatwassoseverethatundigestedlipidscouldbeseenintheirfeces.Whenth
esemicewereplacedonalow-
fatdiet,theirabilitytodigestfatreturnedtonormal.Thesefindingssuggestthatcolipaseplaysacritical,but
notessential,roleinthedigestionofdietarylipidsbypancreaticlipases.