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Samenvatting Metabolisme
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Dit document bevat alle modules van Sofia, gegeven door Prof. Schuit. De hoofdstukken volgen het handboek en extra modules nadien.
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InhoudHoofdstuk 1: biomoleculen ................................................................................................................... 10
Inleiding ............................................................................................................................................. 10
Koolhydraten (suikers) ...................................................................................................................... 10
Monosachariden en disachariden ................................................................................................. 11
Oligosachariden ............................................................................................................................. 12
Polysachariden .............................................................................................................................. 13
Glycosaminoglycanen .................................................................................................................... 14
Lipiden (vetten) ................................................................................................................................. 15
Vetzuren (VZ) ................................................................................................................................. 15
Triglyceriden = triacyl-glycerolen .................................................................................................. 17
Glycerolipiden................................................................................................................................ 17
Sfingolipiden .................................................................................................................................. 17
Glycolipiden ................................................................................................................................... 18
Overige lipiden .............................................................................................................................. 18
Membraanlipiden .......................................................................................................................... 19
Membranen: mozaïek van vloeibare en stijve domeinen ............................................................. 19
Aminozuren, peptiden en proteïnen (eiwitten) ................................................................................ 20
Polaire en apolaire zijketens ......................................................................................................... 21
Peptidebinding .............................................................................................................................. 22
Primaire eiwitstructuur ................................................................................................................. 22
Secundaire structuur ..................................................................................................................... 24
Tertiaire structuur ......................................................................................................................... 24
Quaternaire structuur ................................................................................................................... 25
Basen, nucleotiden en nucleïnezuren ............................................................................................... 25
Dubbele helixstructuur .................................................................................................................. 26
Enzymen ............................................................................................................................................ 26
Katalysemechanisme ..................................................................................................................... 27
Classificatie .................................................................................................................................... 27
Polymorfisme .................................................................................................................................... 29
Kenmerken van biologische evolutie ............................................................................................ 30
Hoofdstuk 2: wat is metabolisme? ........................................................................................................ 31
Inleiding ............................................................................................................................................. 31
1
, Metabolisme ..................................................................................................................................... 31
Metabolisme is een groot actieterrein.............................................................................................. 32
Metabole paden ................................................................................................................................ 33
Metabole flux .................................................................................................................................... 33
Anabole en katabole wegen .............................................................................................................. 33
Katabolisme ................................................................................................................................... 33
Anabolisme .................................................................................................................................... 35
Ribonucleotiden als metabole dragers.............................................................................................. 35
ATP................................................................................................................................................. 36
NADH, NADPH en FADH2 .............................................................................................................. 37
Co-enzym A.................................................................................................................................... 38
Andere dragers .............................................................................................................................. 38
Metabolisme wordt intensief geregeld ............................................................................................. 39
Regeling van de hoeveelheid parallel werkende enzymmoleculen .............................................. 39
Regeling van de enzymactiviteit .................................................................................................... 40
Regeling van de toegankelijkheid van het enzym voor het substraat........................................... 40
Beperken van diffusieafstanden en ongewenste reacties ............................................................ 40
Verdeling van metabole paden over celcompartimenten ............................................................ 41
Hormonen en neurotransmitters regelen de metabole flux ......................................................... 43
Fouten in het metabolisme ............................................................................................................... 43
Stofwisselingsziekten..................................................................................................................... 43
Metaboliet herstelmechanismen .................................................................................................. 48
Hoofdstuk 3: glycolyse........................................................................................................................... 49
Inleiding ............................................................................................................................................. 49
Een oude heirbaan met wetenschappelijke mijlpalen ...................................................................... 49
Overzicht van de glycolyse ................................................................................................................ 49
Glucoseopname in de cel .............................................................................................................. 50
Tien reactiestappen van de glycolyse (nog foto’s invoegen van de reacties) ............................... 51
Verdere metabole bestemming van pyruvaat .............................................................................. 54
Regeling van de glycolyse .................................................................................................................. 56
Regeling van bestaande enzymen ................................................................................................. 56
Regeling door aanmaak van nieuwe enzymen .............................................................................. 57
Metabolisme van fructose en galactose ....................................................................................... 57
Gevolgen van toegenomen sucrosegebruik .................................................................................. 58
2
, Lactose is de voedingssuiker van zoogdierenmelk........................................................................ 59
Lactosetolerantie ........................................................................................................................... 59
Fructosemie ................................................................................................................................... 59
Galactosemie ................................................................................................................................. 59
Hoofdstuk 4: mitochondriale koolstofverbranding ............................................................................... 60
Inleiding ............................................................................................................................................. 60
........................................................................................................................................................... 61
Mitochondriën................................................................................................................................... 61
Oxidatieve decarboxylering ............................................................................................................... 62
Pyruvaatdehydrogenasecomplex .................................................................................................. 62
Regeling van pyruvaatdehydrogenasecomplex ............................................................................ 63
Citroenzuurcyclus .............................................................................................................................. 63
Organisatie van een bijzondere metabole weg............................................................................. 63
Regeling van de flux van de citroenzuurcyclus.............................................................................. 63
Anabole functies van de citroenzuurcyclus: anaplerose ............................................................... 64
Bèta-oxidatie van vetzuren ............................................................................................................... 64
Principe van bèta-oxidatie............................................................................................................. 64
Eigenlijke bèta-oxidatie ................................................................................................................. 65
Peroxisomale vetzuuroxidatie ....................................................................................................... 66
Oxidatie van ketonlichamen (zie verder hfdst 6 en 12) .................................................................... 67
Aminozuuroxidatie (zie hfdst 9) ........................................................................................................ 68
Hoofdstuk 5: oxidatieve fosforylering ................................................................................................... 69
Inleiding ............................................................................................................................................. 69
Overzicht van de oxidatieve fosforylering......................................................................................... 69
Energetica van redoxreacties ............................................................................................................ 69
Overzicht van de elektronentransportketen ..................................................................................... 70
Eiwitcomplexen van de elektronentransportketen .......................................................................... 70
Ziekten tgv ROS ............................................................................................................................. 71
Mitochondriale ATP-synthase (complex V) ....................................................................................... 71
Maximaal rendement van de oxidatieve fosforylering ..................................................................... 71
Respiratoire ontkoppeling ................................................................................................................. 72
Shuttles voor cytoplasmatisch NADH ................................................................................................ 73
Mitochondriale ziekten ..................................................................................................................... 73
Hoofdstuk 6: glycogeenmetabolisme, gluconeogenese en ketogenese ............................................... 75
3
, Inleiding ............................................................................................................................................. 75
Glycogeenmetabolisme ..................................................................................................................... 75
Glucose-1-fosfaat in lever en spieren............................................................................................ 76
Glycogenolyse................................................................................................................................ 76
Glycogeensynthese........................................................................................................................ 76
Regeling van glycogeenmetabolisme ............................................................................................ 77
Cascade-effect → amplificatie ...................................................................................................... 78
Gluconeogenese ................................................................................................................................ 78
Verloop gluconeogenese (vergelijkbaar met glycolyse)................................................................ 79
Drie bijzondere reacties (anders dan glycolyse, want die waren onomkeerbaar)........................ 79
Reciproke regeling van gluconeogenese en glycolyse .................................................................. 80
Glycogenosen .................................................................................................................................... 81
Ketogenese ........................................................................................................................................ 82
Paradigmaverschuiving ................................................................................................................. 82
Verloop ketogenese ...................................................................................................................... 82
Ketonlichamen............................................................................................................................... 83
Adaptaties bij vasten ..................................................................................................................... 83
Ketogeen dieet .............................................................................................................................. 83
Hoofdstuk 7: vetzuursynthese, cholesterolsynthese en pentosefosfaatweg ....................................... 84
Vetzuursynthese ................................................................................................................................ 84
Aanmaak triglyceriden................................................................................................................... 86
Mevalonaatweg (de novo cholesterolsynthese) ............................................................................... 86
Lipidentransport ................................................................................................................................ 87
Lipoproteïnen verpakken wateronoplosbare cholesterolesters en triglyceriden......................... 87
Verloop lipidentransport ............................................................................................................... 88
3 mechanismen van cholesteroltransport .................................................................................... 88
Homeostase van cholesterol ......................................................................................................... 89
Regulatie van expressie HMG-CoA reductase ............................................................................... 89
Atherosclerose............................................................................................................................... 90
Paradigmaverschuiving ................................................................................................................. 91
Pentosefosfaatweg ............................................................................................................................ 91
4 verschillende modaliteiten van werking pentosefosfaatweg .................................................... 93
Glucose-6-fosfaat dehydrogenase-deficiëntie (G6PD-deficiëntie) ............................................... 94
Hoofdstuk 8: vetzuur- en cholesterolafgeleiden ................................................................................... 96
4
, Inleiding ............................................................................................................................................. 96
Van vetzuren afgeleide biomoleculen ............................................................................................... 96
Glycerolipiden................................................................................................................................ 96
Sfingolipiden .................................................................................................................................. 97
Lipid messengers ........................................................................................................................... 98
Van cholesterol afgeleide biomoleculen ......................................................................................... 102
Cytochroom P450 (CYP’s) (zie hfdst 16) ...................................................................................... 102
Galzouten .................................................................................................................................... 102
Steroïdhormonen ........................................................................................................................ 103
Retinoïnezuur (> vit A) ................................................................................................................. 106
Vitamine D ................................................................................................................................... 106
Hoofdstuk 9: aminozuurmetabolisme ................................................................................................. 107
Essentiële en niet-essentiële AZ ...................................................................................................... 107
Inleiding ....................................................................................................................................... 107
Synthese van niet-essentiële AZ .................................................................................................. 108
Eiwitbehoefte en voedingsproducten die eiwitrijk zijn ............................................................... 110
Aminozuurkatabolisme ................................................................................................................... 110
Detoxificatie van ammoniak: ureumcyclus ................................................................................. 111
Opbouw lever .............................................................................................................................. 112
Van aminozuren afgeleide stoffen .................................................................................................. 112
Inleiding ....................................................................................................................................... 112
Creatine ....................................................................................................................................... 113
Carnitine ...................................................................................................................................... 113
Carnosine ..................................................................................................................................... 113
Arginine ....................................................................................................................................... 113
Polyaminen .................................................................................................................................. 114
Heemsynthese ............................................................................................................................. 114
Afbraak van heem ....................................................................................................................... 116
Stikstofmonoxide (NO) ................................................................................................................ 118
Melanine...................................................................................................................................... 118
Metabole paden voor synthese/afbraak van NT......................................................................... 121
Catecholamines ........................................................................................................................... 121
Serotonine ................................................................................................................................... 122
Melatonine .................................................................................................................................. 123
5
, Glutamaat .................................................................................................................................... 123
GABA............................................................................................................................................ 124
Acetylcholine ............................................................................................................................... 124
Endocannabinoïden ..................................................................................................................... 125
Thyroxine ..................................................................................................................................... 126
Verschillen tussen neurotransmitters en neuropeptiden ............................................................... 127
Hoofdstuk 10: nucleotidenmetabolisme ............................................................................................. 128
Inleiding ........................................................................................................................................... 128
Purinesynthese ................................................................................................................................ 129
De novo purinesynthese.............................................................................................................. 129
Salvageweg van purinesynthese ................................................................................................. 130
Antimetabolieten van de purinesynthese ................................................................................... 130
Pyrimidinesynthese ......................................................................................................................... 131
De novo pyrimidinesynthese ....................................................................................................... 131
Salvageweg van pyrimidinesynthese........................................................................................... 131
Synthese van DNA-bouwstenen .................................................................................................. 132
Synthese van dTMP ..................................................................................................................... 132
Antimetabolieten van pyrimidinesynthese ................................................................................. 133
Jicht.................................................................................................................................................. 134
Oorzaken ..................................................................................................................................... 134
Normale metabole weg afbraak purinebasen............................................................................. 134
Metabole weg bij jicht ................................................................................................................. 135
Behandeling ................................................................................................................................. 136
Hoofdstuk 12: lichaamsmetabolisme .................................................................................................. 138
Glucosehomeostase ........................................................................................................................ 138
Postprandiaal............................................................................................................................... 138
Interprandiaal .............................................................................................................................. 138
Hormonen van eilandjes van Langerhans (1-2% van pancreas).................................................. 139
Energievoorraad .............................................................................................................................. 139
Postprandiaal............................................................................................................................... 140
Interprandiaal .............................................................................................................................. 140
Scenario’s in opbouwen van vetreserve ..................................................................................... 140
Diabetes........................................................................................................................................... 141
Diabetes type I (10%) .................................................................................................................. 142
6
, Diabetes type II (90%) ................................................................................................................. 142
Diabetes tussenvorm ................................................................................................................... 143
Problemen diabetes .................................................................................................................... 143
Genetische factoren .................................................................................................................... 144
Obesitas ........................................................................................................................................... 145
Problemen obesitas ..................................................................................................................... 145
Vetweefsel ................................................................................................................................... 147
Genetische factoren .................................................................................................................... 148
Bariatrische chirurgie .................................................................................................................. 149
Hoofdstuk 13: stofwisselingsziekten ................................................................................................... 150
Soorten ............................................................................................................................................ 150
Fenylketonurie (zie p111) ............................................................................................................ 150
Albinisme ..................................................................................................................................... 150
Tyrosinemie ................................................................................................................................. 151
Alkaptonurie ................................................................................................................................ 151
Glycogenose type II (ziekte van Pompe) ..................................................................................... 151
Fructosurie................................................................................................................................... 151
Fructosemie ................................................................................................................................. 151
ADA (adenosine deaminase deficiëntie) ..................................................................................... 151
Neonatale hypothyroïdie ............................................................................................................ 151
Adrenogenitaal syndroom = congenitale bijnierhyperplasie ...................................................... 152
Behandeling stofwisselingsziekte .................................................................................................... 152
Vormen van therapie................................................................................................................... 152
CRISPR -Cas9 ................................................................................................................................ 153
Screenen ...................................................................................................................................... 153
Metaboliet herstelmechanismen ................................................................................................ 153
Hoofdstuk 14: NT en neuropeptiden .................................................................................................. 154
Inleiding peptiden (algemeen) ........................................................................................................ 154
Post-translationele bewerkingen van preprohormoon .............................................................. 154
Afbraak van peptidehormonen ................................................................................................... 154
Peptidehormonen ........................................................................................................................... 154
TRH .............................................................................................................................................. 154
Insuline ........................................................................................................................................ 155
POMC tot alfa-MSH ..................................................................................................................... 155
7
, Renine-angiotensine.................................................................................................................... 156
Prohormoon convertasen ........................................................................................................... 156
Werking gezonde bètacellen ........................................................................................................... 156
Regeling door glucose ................................................................................................................. 157
Versterking door incretine .......................................................................................................... 157
Remmen door noradrenaline en adrenaline ............................................................................... 158
Werking gezonde alfacellen ............................................................................................................ 158
Stimulatie door noradrenaline en adrenaline ............................................................................. 158
Hyperglycemie ................................................................................................................................. 158
Diabetes type I............................................................................................................................. 158
Diabetes type II............................................................................................................................ 159
Hypoglycemie .................................................................................................................................. 159
Bijnier .............................................................................................................................................. 160
Syndroom van crushing ............................................................................................................... 160
Schildklier ........................................................................................................................................ 161
Hypothyroïdie .............................................................................................................................. 161
Hyperthyroïdie............................................................................................................................. 161
Inleiding hormonen ......................................................................................................................... 162
Endocrien systeem .......................................................................................................................... 163
Wateroplosbare peptidehormonen ............................................................................................ 164
Wateronoplosbare steroïdhormonen ......................................................................................... 164
Expressie van hormoonreceptoren op doelwitcel is dynamisch en regelbaar ........................... 164
Adrenaline gaat tegelijk insulinesecretie inhiberen en glucagonsecretie stimuleren ................ 165
POMC (pro-opio melanocortine) ................................................................................................. 165
Insulinegevoeligheid .................................................................................................................... 165
Hoofdstuk 15: energie-uitgave ............................................................................................................ 166
Inleiding ........................................................................................................................................... 166
3 grote energie-uitgaven ................................................................................................................. 166
Rendement .................................................................................................................................. 167
ATP............................................................................................................................................... 167
Spiervezels ....................................................................................................................................... 167
Vetcellen .......................................................................................................................................... 169
Hoofdstuk 16: detoxificatie ................................................................................................................. 170
Cholesterol .................................................................................................................................. 170
8
,Detoxificatie van nutteloze of schadelijke stoffen in de voeding ................................................... 170
Fase 1: detoxificatie via cytochroom P450 systeem ................................................................... 171
Fase 2: conjugatie aan een polaire groep ................................................................................... 173
Detoxificatie van alcohol ................................................................................................................. 174
Ethylisme ..................................................................................................................................... 174
Nutritionele intoxicatie ................................................................................................................... 176
9
, Samenvatting metabolisme
Hoofdstuk 1: biomoleculen
Inleiding
Metabolisme of stofwisseling = verzamelnaam voor alle chemische reacties die het leven nodig heeft
om zichzelf in stand te houden, te groeien, zich als een soort te verdedigen en nakomelingen te
produceren
4 belangrijke klassen van biomoleculen:
- Koolhydraten
- Lipiden
- Proteïnen
- Nucleïnezuren
Koolhydraten (suikers)
- Eigenschappen
o Bestaan uit monosachariden
o Glycosidebindingen → vorming van meerdere
▪ Disachariden
▪ Oligosachariden
▪ Polysachariden
o Belangrijk in de stereochemie
o Belangrijk in voeding en metabolisme
o Niet gecodeerd in het genoom
o Polymeren hebben geen uniek grootte → molmassa kan variëren
o Meest voorkomende biomoleculen op Aarde
- Belangrijke koolhydraten
o Glyceraldehyde en dihydroxyaceton
▪ Functie: metabolisme
o Ribose en deoxyribose
▪ Bouwstenen van RNA, DNA en nucleotiden
o Glucose, galactose en fructose
▪ Belangrijkste brandstoffen voor metabolisme
o Sucrose, lactose
▪ Voedingsstoffen (tafelsuiker en melksuiker)
o Maltose
▪ Splitsingsproduct van zetmeel
o Mannose, fucose, siaalzuur, glucuronzuur, N-acetylgalactose, N-acetylglucosamine
▪ Gebonden aan serine-, threonine- of asparaginezijketens van eiwitten =
glycoproteïnen
▪ Gebonden aan serine-, threonine- of asparaginezijketens van ceramide =
glycolipiden
10
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