10/8/2016

GlucagonWikipedia,thefreeencyclopedia

Glucagon
FromWikipedia,thefreeencyclopedia

Glucagonisapeptidehormone,producedbyalphacellsofthepancreas.Itworkstoraisethe
concentrationofglucoseinthebloodstream.Itseffectisoppositethatofinsulin,whichlowersthe
glucose.[2]
Thepancreasreleasesglucagonwhentheconcentrationofglucoseinthebloodstreamfallstoolow.
Glucagoncausesthelivertoconvertstoredglycogenintoglucose,whichisreleasedintothe
bloodstream.[3]Highbloodglucoselevelsstimulatethereleaseofinsulin.Insulinallowsglucosetobe
takenupandusedbyinsulindependenttissues.Thus,glucagonandinsulinarepartofafeedback
systemthatkeepsbloodglucoselevelsatastablelevel.Itincreasesenergyexpenditureandiselevated
underconditionsofstress.[4]Glucagonbelongstoafamilyofseveralotherrelatedhormones.
ItisontheWHOModelListofEssentialMedicines,themostimportantmedicationsneededinabasic
healthsystem.[5]

Contents
1 Function
2 Medicaluses
2.1 Hypoglycemia
2.2 Betablockeroverdose
2.3 Anaphylaxis
2.4 Impactedfoodbolus
2.5 Endoscopicretrogradecholangiopancreatography
3 Adverseeffects
3.1 Contraindications
4 Mechanismofaction
5 Physiology
5.1 Production
5.2 Regulation
6 Structure
7 Pathology
8 History
8.1 Etymology
https://en.wikipedia.org/wiki/Glucagon

GCG

Availablestructures
PDB

HumanUniProtsearch:PDBe(http://www.ebi.ac.u
k/pdbe/searchResults.html?display=both&term=P01
275)RCSB(http://www.rcsb.org/pdb/search/smartS
ubquery.do?smartSearchSubtype=UpAccessionIdQu
ery&accessionIdList=P01275)
ListofPDBidcodes
1BH0(http://www.rcsb.org/pdb/explore/explore.d
o?pdbId=1BH0),1D0R(http://www.rcsb.org/pdb/
explore/explore.do?pdbId=1D0R),1NAU(http://w
ww.rcsb.org/pdb/explore/explore.do?pdbId=1NA
U),2G49(http://www.rcsb.org/pdb/explore/explor
e.do?pdbId=2G49),2L63(http://www.rcsb.org/pd
b/explore/explore.do?pdbId=2L63),2L64(http://
www.rcsb.org/pdb/explore/explore.do?pdbId=2L6
4),2M5P(http://www.rcsb.org/pdb/explore/explor
e.do?pdbId=2M5P),2M5Q(http://www.rcsb.org/p
db/explore/explore.do?pdbId=2M5Q),3IOL(htt
p://www.rcsb.org/pdb/explore/explore.do?pdbId=3
IOL),4APD(http://www.rcsb.org/pdb/explore/exp
lore.do?pdbId=4APD),4ZGM(http://www.rcsb.or
g/pdb/explore/explore.do?pdbId=4ZGM)
Identifiers
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8.1 Etymology
9 Seealso
10 References

Function
Glucagongenerallyelevatestheconcentrationofglucoseinthebloodbypromotinggluconeogenesis
andglycogenolysis.
Glucoseisstoredintheliverintheformofthepolysaccharideglycogen,whichisaglucan(apolymer
madeupofglucosemolecules).Livercells(hepatocytes)haveglucagonreceptors.Whenglucagon
bindstotheglucagonreceptors,thelivercellsconverttheglycogenintoindividualglucosemolecules
andreleasethemintothebloodstream,inaprocessknownasglycogenolysis.Asthesestoresbecome
depleted,glucagonthenencouragestheliverandkidneytosynthesizeadditionalglucoseby
gluconeogenesis.Glucagonturnsoffglycolysisintheliver,causingglycolyticintermediatestobe
shuttledtogluconeogenesis.
Glucagonalsoregulatestherateofglucoseproductionthroughlipolysis.Glucagoninduceslipolysisin
humansunderconditionsofinsulinsuppression(suchasdiabetesmellitustype1).[6]
Glucagonproductionappearstobedependentonthecentralnervoussystemthroughpathwaysyetto
bedefined.Ininvertebrateanimals,eyestalkremovalhasbeenreportedtoaffectglucagonproduction.
Excisingtheeyestalkinyoungcrayfishproducesglucagoninducedhyperglycemia.[7]

Medicaluses
Hypoglycemia
Aninjectableformofglucagonisvitalfirstaidincasesofseverehypoglycemiawhenthevictimis
unconsciousorforotherreasonscannottakeglucoseorally.Thedoseforanadultistypically1
milligram,andtheglucagonisgivenbyintramuscular,intravenousorsubcutaneousinjection,and
quicklyraisesbloodglucoselevels.Tousetheinjectableform,itmustbereconstitutedpriortouse,a
stepthatrequiresasterilediluenttobeinjectedintoavialcontainingpowderedglucagon,becausethe
hormoneishighlyunstablewhendissolvedinsolution.Whendissolvedinafluidstate,glucagoncan
formamyloidfibrils,ortightlywovenchainsofproteinsmadeupoftheindividualglucagonpeptides,
https://en.wikipedia.org/wiki/Glucagon

Aliases(ht GCG(http://www.genenames.org/cgibin/gene_sy
tps://en.wi mbol_report?hgnc_id=4191),GLP1,GLP2,
kipedia.or GRPP,glucagon,Glucagon
g/wiki/Hu
man_Gen
ome_Orga
nisation)
External
IDs

OMIM:138030(https://omim.org/entry/138030)
HomoloGene:136497(https://www.ncbi.nlm.ni
h.gov/entrez/query.fcgi?cmd=Retrieve&db=homo
logene&dopt=HomoloGene&list_uids=136497)
GeneCards:2641(http://www.genecards.org/cgi
bin/carddisp.pl?id_type=entrezgene&id=2641)
Geneontology

Molecular
hormoneactivity(http://amigo.geneontology.
function org/amigo/term/GO:0005179)
proteinbinding(http://amigo.geneontology.or
g/amigo/term/GO:0005515)
identicalproteinbinding(http://amigo.geneon
tology.org/amigo/term/GO:0042802)
receptorbinding(http://amigo.geneontology.o
rg/amigo/term/GO:0005102)
glucagonreceptorbinding(http://amigo.geneo
ntology.org/amigo/term/GO:0031769)
Cellular
cytoplasm(http://amigo.geneontology.org/ami
component go/term/GO:0005737)
endoplasmicreticulumlumen(http://amigo.ge
neontology.org/amigo/term/GO:0005788)
plasmamembrane(http://amigo.geneontology.
org/amigo/term/GO:0005886)
extracellularspace(http://amigo.geneontolog
y.org/amigo/term/GO:0005615)
extracellularregion(http://amigo.geneontolog
y.org/amigo/term/GO:0005576)
secretorygranulelumen(http://amigo.geneont
ology.org/amigo/term/GO:0034774)
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andonceglucagonbeginstofibrilize,itbecomesuselesswheninjected,astheglucagoncannotbe
absorbedandusedbythebody.Thereconstitutionprocessmakesusingglucagoncumbersome,
althoughthereareanumberofproductsnowindevelopmentfromanumberofcompaniesthataimto
maketheproducteasiertouse.

Betablockeroverdose
Anecdotalevidencesuggestsabenefitofhigherdosesofglucagoninthetreatmentofoverdosewith
betablockersthelikelymechanismofactionistheincreaseofcAMPinthemyocardium,ineffect
bypassingtheadrenergicsecondmessengersystem.[8]

Anaphylaxis
Somepeoplewhohaveanaphylaxisandareonbetablockersareresistanttoepinephrine.Inthis
situationglucagonintravenouslymaybeusefultotreattheirlowbloodpressure.[9]

Impactedfoodbolus
Glucagonrelaxestheloweresophagealsphincterandmaybeusedinthosewithanimpactedfood
bolusintheesophagus("steakhousesyndrome").[10]Thereislittleevidenceforglucagon's
effectivenessinthiscondition,[11][12][13]andglucagonmayinducenauseaandvomiting,[13]but
consideringthesafetyofglucagonthisisstillconsideredanacceptableoptionaslongitdoesnotlead
todelaysinarrangingothertreatments.[14][15]

Endoscopicretrogradecholangiopancreatography
Glucagon'seffectofdecreasingcAMPcausesrelaxationofsplanchnicsmoothmuscle,allowing
cannulationoftheduodenumduringtheendoscopicretrogradecholangiopancreatography(ERCP)
procedure.

Adverseeffects
Glucagonactsveryquicklycommonsideeffectsincludeheadacheandnausea.

https://en.wikipedia.org/wiki/Glucagon

Biological
negativeregulationofexecutionphaseof
process
apoptosis(http://amigo.geneontology.org/amig
o/term/GO:1900118)
positiveregulationofcAMPbiosynthetic
process(http://amigo.geneontology.org/amigo/t
erm/GO:0030819)
Gproteincoupledreceptorsignalingpathway
(http://amigo.geneontology.org/amigo/term/GO:
0007186)
regulationofinsulinsecretion(http://amigo.g
eneontology.org/amigo/term/GO:0050796)
negativeregulationofintrinsicapoptotic
signalingpathway(http://amigo.geneontology.o
rg/amigo/term/GO:2001243)
cellularresponsetoglucagonstimulus(http://
amigo.geneontology.org/amigo/term/GO:00713
77)
negativeregulationofapoptoticprocess(htt
p://amigo.geneontology.org/amigo/term/GO:00
43066)
positiveregulationofpeptidylserine
phosphorylation(http://amigo.geneontology.or
g/amigo/term/GO:0033138)
positiveregulationofcalciumionimport(htt
p://amigo.geneontology.org/amigo/term/GO:00
90280)
positiveregulationofhistoneH3K4
methylation(http://amigo.geneontology.org/ami
go/term/GO:0051571)
positiveregulationofpeptidylthreonine
phosphorylation(http://amigo.geneontology.or
g/amigo/term/GO:0010800)
feedingbehavior(http://amigo.geneontology.o
rg/amigo/term/GO:0007631)
positiveregulationofERK1andERK2
cascade(http://amigo.geneontology.org/amigo/t
erm/GO:0070374)
proteinkinaseAsignaling(http://amigo.geneo
ntology.org/amigo/term/GO:0010737)
responsetostarvation(http://amigo.geneontol
ogy.org/amigo/term/GO:0042594)
positiveregulationofinsulinsecretion
involvedincellularresponsetoglucose
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stimulus(http://amigo.geneontology.org/amigo/
term/GO:0035774)
positiveregulationofproteinbinding(http://a
migo.geneontology.org/amigo/term/GO:003209
2)
cellproliferation(http://amigo.geneontology.o
rg/amigo/term/GO:0008283)
positiveregulationofgluconeogenesisby
positiveregulationoftranscriptionfromRNA
polymeraseIIpromoter(http://amigo.geneontol
ogy.org/amigo/term/GO:0035948)
signaltransduction(http://amigo.geneontolog
y.org/amigo/term/GO:0007165)
positiveregulationofproteinkinaseactivity
(http://amigo.geneontology.org/amigo/term/GO:
0045860)
adenylatecyclasemodulatingGprotein
coupledreceptorsignalingpathway(http://amig
o.geneontology.org/amigo/term/GO:0007188)

Druginteractions:Glucagoninteractsonlywithoralanticoagulants,increasingthetendencyto
bleed.[16]

Contraindications
Whileglucagoncanbeusedclinicallytotreatvariousformsofhypoglycemia,itisseverely
contraindicatedinpatientswithpheochromocytoma,asthedruginteractionwithelevatedlevelsof
adrenalineproducedbythetumormayproduceanexponentialincreaseinbloodsugarlevels,leading
toahyperglycemicstate,whichmayincurafatalelevationinbloodpressure.[17]Likewise,glucagonis
contraindicatedinpatientswithaninsulinoma,asitsusemayleadtoreboundhypoglycemia.[17]

Mechanismofaction
Glucagonbindstotheglucagonreceptor,aGproteincoupledreceptor,locatedintheplasma
membrane.TheconformationchangeinthereceptoractivatesGproteins,aheterotrimericproteinwith
,,andsubunits.WhentheGproteininteractswiththereceptor,itundergoesaconformational
changethatresultsinthereplacementoftheGDPmoleculethatwasboundtothesubunitwitha
GTPmolecule.Thissubstitutionresultsinthereleasingofthesubunitfromtheandsubunits.The
alphasubunitspecificallyactivatesthenextenzymeinthecascade,adenylatecyclase.

Sources:Amigo(http://amigo.geneontology.org/)/QuickGO
(http://www.ebi.ac.uk/QuickGO/)
RNAexpressionpattern

Adenylatecyclasemanufacturescyclicadenosinemonophosphate(cyclicAMPorcAMP),which
activatesproteinkinaseA(cAMPdependentproteinkinase).Thisenzyme,inturn,activates
phosphorylasekinase,whichthenphosphorylatesglycogenphosphorylaseb,convertingitintothe
activeformcalledphosphorylasea.Phosphorylaseaistheenzymeresponsibleforthereleaseof
glucose1phosphatefromglycogenpolymers.
Additionally,thecoordinatedcontrolofglycolysisandgluconeogenesisintheliverisadjustedbythe
phosphorylationstateoftheenzymesthatcatalyzetheformationofapotentactivatorofglycolysis
calledfructose2,6bisphosphate.[18]TheenzymeproteinkinaseAthatwasstimulatedbythecascade
initiatedbyglucagonwillalsophosphorylateasingleserineresidueofthebifunctionalpolypeptide
chaincontainingboththeenzymesfructose2,6bisphosphataseandphosphofructokinase2.This
covalentphosphorylationinitiatedbyglucagonactivatestheformerandinhibitsthelatter.This
regulatesthereactioncatalyzingfructose2,6bisphosphate(apotentactivatorofphosphofructokinase
1,theenzymethatistheprimaryregulatorystepofglycolysis)[19]byslowingtherateofitsformation,
therebyinhibitingthefluxoftheglycolysispathwayandallowinggluconeogenesistopredominate.
Thisprocessisreversibleintheabsenceofglucagon(andthus,thepresenceofinsulin).
https://en.wikipedia.org/wiki/Glucagon

Morereferenceexpressiondata(http://biogps.org/gene/2641/)
Orthologs
Species

Human

Mouse

Entrez
2641(https://www.ncbi.nlm.nih.gov/ent n/a
rez/query.fcgi?db=gene&cmd=retrieve
&dopt=default&list_uids=2641&rn=1)

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GlucagonstimulationofPKAalsoinactivatestheglycolyticenzymepyruvatekinase.[20]

Ensembl

Physiology
Production
Thehormoneissynthesizedandsecretedfromalphacells(cells)oftheisletsofLangerhans,which
arelocatedintheendocrineportionofthepancreas.Production,whichisotherwisefreerunning,is
suppressed/regulatedbyinsulinfromtheadjacentbetacells.Whenbloodsugardrops,insulin
productiondropsandmoreglucagonisproduced.[21]Inrodents,thealphacellsarelocatedintheouter
rimoftheislet.Humanisletstructureismuchlesssegregated,andalphacellsaredistributed
throughouttheisletincloseproximitytobetacells.Glucagonisalsoproducedbyalphacellsinthe
stomach.[22]

Hypoglycemia
Epinephrine(via2,2,[23]and1[24]adrenergicreceptors)
Arginine
Alanine(oftenfrommusclederivedpyruvate/glutamatetransamination(seealanine
transaminasereaction).
Acetylcholine[25]
Cholecystokinin
Secretionofglucagonisinhibitedby:
Somatostatin
Insulin(viaGABA)[26]
PPAR/retinoidXreceptorheterodimer.[27]
Increasedfreefattyacidsandketoacidsintotheblood.[28]
Increasedureaproduction

n/a

P01275(http://www.uniprot.org/unipro
t/P01275)

n/a

UniProt

RefSeq
(mRNA)

RefSeq
(protein)

Regulation
Secretionofglucagonisstimulatedby:

ENSG00000115263(http://www.ensem
bl.org/Homo_sapiens/geneview?gene=E
NSG00000115263db=core)

NM_002054(https://www.ncbi.nlm.ni
n/a
h.gov/entrez/viewer.fcgi?val=NM_0020
54)

NP_002045.1(https://www.ncbi.nlm.ni n/a
h.gov/entrez/viewer.fcgi?val=NP_00204
5.1)

Location
(UCSC)

Chr2:162.14162.15Mb(http://geno n/a
me.ucsc.edu/cgibin/hgTracks?org=Hum
an&db=hg38&position=chr2:162142873
162152404)

PubMed
search

[1]

n/a
Wikidata
View/EditHuman

Glucagon

Structure
https://en.wikipedia.org/wiki/Glucagon

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Glucagonisa29aminoacidpolypeptide.Itsprimarystructureinhumansis:NH2HisSerGlnGlyThrPhe
ThrSerAspTyrSerLysTyrLeuAspSerArgArgAlaGlnAspPheValGlnTrpLeuMetAsnThrCOOH.
Thepolypeptidehasamolecularweightof3485daltons.Glucagonisapeptide(nonsteroid)hormone.
Glucagonisgeneratedfromthecleavageofproglucagonbyproproteinconvertase2inpancreaticisletcells.In
intestinalLcells,proglucagoniscleavedtothealternateproductsglicentin,GLP1(anincretin),IP2,andGLP
2(promotesintestinalgrowth).[29]

Pathology
Abnormallyelevatedlevelsofglucagonmaybecausedbypancreatictumors,suchasglucagonoma,symptoms
ofwhichincludenecrolyticmigratoryerythema,reducedaminoacids,andhyperglycemia.Itmayoccuraloneor
inthecontextofmultipleendocrineneoplasiatype1.

Glucagonballandstickmodel,withthecarboxyl
terminusaboveandtheaminoterminusbelow
Identifiers

History
Inthe1920s,KimballandMurlinstudiedpancreaticextracts,andfoundanadditionalsubstancewith
hyperglycemicproperties.Theydescribedglucagonin1923.[30]Theaminoacidsequenceofglucagonwas
describedinthelate1950s.[31]Amorecompleteunderstandingofitsroleinphysiologyanddiseasewasnot
establisheduntilthe1970s,whenaspecificradioimmunoassaywasdeveloped.

Etymology
Glucagonwasnamedin1923,probablyfromtheGreeksweet,andtolead.[32]

PubChem

IUPHAR/BPS 1136(http://www.guidetopharmacol
ogy.org/GRAC/LigandDisplayForwa
rd?ligandId=1136)
ChemSpider 10481928(http://www.chemspider.c
om/ChemicalStructure.10481928.ht
ml)
UNII

Seealso
Cortisol
Diabetesmellitus
Glucagonlikepeptide1
Glucagonlikepeptide2
Insulin
IsletsofLangerhans
Pancreas
https://en.wikipedia.org/wiki/Glucagon

CID16186314(https://pubchem.ncb
i.nlm.nih.gov/summary/summary.cg
i?cid=16186314)

76LA80IG2G(http://fdasis.nlm.nih.
gov/srs/srsdirect.jsp?regno=76LA80I
G2G)

ChEMBL

CHEMBL266481(https://www.ebi.a
c.uk/chembldb/index.php/compound/
inspect/CHEMBL266481)
Chemicaldata

Formula

C153H225N43O49S
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Proglucagon
Tyrosinekinase

References
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https://en.wikipedia.org/wiki/Glucagon

Molarmass

3482.747314g/mol

InChI

(whatisthis?)(verify)

Metabolicregulationofglycogenbyglucagon.

Amicroscopicimagestainedfor
glucagon

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17."InformationforthePhysician:GlucagonforInjection(rDNAorigin)"(PDF).EliLillyandCompany.
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SJ(1984)."Theroleoffructose2,6bisphosphateintheregulationofcarbohydratemetabolism".Current
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therapeuticmakeover".NCBI.JClinInvest.doi:10.1172/JCI60016.Retrieved10February2016.
22.UngerRH,CherringtonAD(2012)."Glucagonocentricrestructuringofdiabetes:apathophysiologicand
therapeuticmakeover".TheJournalofClinicalInvestigation.122(1):412.doi:10.1172/JCI60016.
PMC3248306 .PMID22214853.
23.LaydenBT,DuraiV,LoweWL(2010)."GProteinCoupledReceptors,PancreaticIslets,andDiabetes".
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24.SkoglundG,LundquistI,AhrnB(Nov1987)."Alpha1andalpha2adrenoceptoractivationincreases
plasmaglucagonlevelsinthemouse".EuropeanJournalofPharmacology.143(1):838.
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system".CellMetabolism.3(1):4758.doi:10.1016/j.cmet.2005.11.015.PMID16399504.
27.KrtznerR,FrhlichF,LeplerK,SchrderM,RherK,DickelC,TzvetkovMV,QuentinT,OetjenE,
KnepelW(Feb2008)."AperoxisomeproliferatoractivatedreceptorgammaretinoidXreceptorheterodimer
physicallyinteractswiththetranscriptionalactivatorPAX6toinhibitglucagongenetranscription".
MolecularPharmacology.73(2):509517.doi:10.1124/mol.107.035568.PMID17962386.
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proglucagoninman".Diabetologia.30(11):87481.doi:10.1007/BF00274797.PMID3446554.
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Biol.Chem.58(1):337348.
31.BromerW,WinnL,BehrensO(1957)."TheaminoacidsequenceofglucagonV.Locationofamidegroups,
aciddegradationstudiesandsummaryofsequentialevidence".J.Am.Chem.Soc.79(11):28072810.
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32.glucagon(http://dictionary.reference.com/browse/glucagon)ondictionary.com

https://en.wikipedia.org/wiki/Glucagon

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GlucagonWikipedia,thefreeencyclopedia

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https://en.wikipedia.org/wiki/Glucagon

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