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Ch.5Microbialmetabolism
Metabolismbasics(Fig.5.1)
Breakdownofcarbohydrates,lipidsandproteins
toproducecellularenergy(catabolism)
Redox(reduction/oxidation)reactionscapture,
storeanduseenergyviaelectrontransfers
amongmolecules
Photosynthesiscapturesandstoresenergyfrom
Photosynthesis captures and stores energy from
light
Biosynthesisofmacromoleculesandtheir
precursors(anabolicreactions)
Metabolismisthesumofallbiochemicalreactionsinthe
cell(pathways)
Catabolicpathwaysbreakdownnutrientstoyieldsmaller
moleculesandcapturestoredenergy
Anabolicpathwayssynthesizelargermoleculesfrom
smallerprecursorsanduseenergy
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Basicconceptsofmetabolicprocesses
Redoxreactions(Fig.5.2)
Cellsobtainnutrientsfromphotosynthesisand/or
transportfromenvironment
Nutrientsarecatabolizedtoreleaseandcapture
storedenergyinformofATPorNADHor
electrochemical(chemiosmotic)gradients
Nutrientsareconvertedto12basicprecursorsby
p
y
enzymesinbasicpathways(e.g.,glycolysis,Krebs
cycleandpentosephosphatecycle)
Anabolicreactionsconvertprecursormolecules
(12)intobuildingblocks,buildingblocksinto
macromolecules(polymerization)and
macromoleculesintocellstructurestoforma
newcell.
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Electroncarriers
Fundamentallyallbiochemicalreactionsinvolvethe
transferofelectrons
Reductionisthegainofelectrons;oxidationislossof
electronsandthesearecoupled
Amoleculethathasbeenreducedbecomesrelatively
morenegative,anoxidizedmoleculesbecomesrelatively
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morepositive
ATPproduction(Fig.5.3)
NAD+/NADH,
NADP+/NADPHand
FAD/FADH2areallcellular
electroncarriersusedby
cells
Electronsaretransferred
l
f
d
withprotons(Htransferred)
Manymetabolicpathways
requiretheseelectron
carriers
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Substratelevelphosphorylationinvolvetransferofhigh
energyphosphatetoADPfromanotherphosphorylated
organiccompound
Oxidativephosphorylationusesenergyfromhighenergy
electrontransfersduringrespirationtomakeATPfrom
ADPandPi
Photophosphorylationusesenergyfromlighttomake 6
electrons high energy
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Enzymes(Figs.5.4,5.5)
Enzymesareusually
proteinsthatcatalyze
(speedup)
biochemical reactions
biochemicalreactions
Enzymesoftenrequire
cofactors
Ribozymesare
enzymesmadeofRNA
insteadofprotein
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Enzymaticreactions(Fig.5.7)
Substratebindsto
activesiteofenzyme
Enzymecatalyzes
reaction
Productsleaveactive
site
Enzymaticactivity(Fig.5.8)
Thermaldenaturationofproteins(Fig.5.9)
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IncreasingheatcausesHbondstobreak
MicrobesliveatalltempsatwhichliquidH2Oisfoundand
theoptimumfolding/activityofproteinscoincidewith
optimalgrowthtemperature
pHcauseschargesonproteinstochange,sodenaturation
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occursviabreakingofionicbonds
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Glycolysis catabolismofglucose(Fig.5.13)
Fig.5.13
Glycolysisisametabolicpathwayinwhich
glucose(6Csugar)isoxidizedtopyruvicacid
(2x3C),producingATPandNADH
Foraerobes,pyruvicacidfurtherbrokendownvia
For aerobes pyruvic acid further broken down via
KrebscycletoyieldmoreNADH,FADH2 andATP
ElectronsfromNADHandFADH2 enterelectron
transportsystemandformaH+gradient
H+gradientusedtomakeATP
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Productsofglycolysis(Fig.5.14
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Substratelevelphosphorylation(Fig.5.15)
2NADHand4ATP
produced
ATPproductionvia
substratelevel
phosphorylations
Fermentationoccursif
pyruvicaciddoesnot
py
entertheKrebscycleand
ifelectronsfromglucose
metabolismdontgo
downETS
Fermentationtypically
anaerobicandyieldsacid
biproducts
Thesereactionsare
partofglycolysis
ATPonlyproducedvia
thismechanismduring
hi
h i d i
fermentation
SufficientlevelsofATP
aremadetoallow
growthofmicrobes
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Pentosephosphatepathway(Fig.5.16)
Pentosephosphatepathway(Fig.5.16)
Usedinconjunctionwithglycolysisbymost
microbes
GeneratesNADPHandprecursorsfor
makingbuildingblocks(aminoacids,
nucleotidesugars)
Generatesstartingmaterialfor
photosynthesis,ribulose5 phosphate
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Krebs(akaTCA)cycle(Fig.5.19)
Respiration theelectrontransportsystem
Fig.5.20
MostoftheATPproduced
inmicrobesisproduced
byanelectrontransport
system
Thesystemisaseriesof
membranebound
electroncarriers
Aselectronsflow,energy
isusedtopumpH+sout
ofcell
ManydifferentETSsoccur
inmicrobes
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ATPproductionduringrespiration(Fig.5.21)
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Fermentation(Fig.5.22andTable5.3)
Fermentation
yieldsofATPsmall
relativeto
respiration
Fermentationof
sugarsquicklyuses
i kl
upNAD+,which
needstobe
regeneratedto
keepupATP
production
H+sarepumpedoutaselectronspassthroughvarious
carriers,especiallycytochromes
AsconcentrationofH+builds,theyareforcedbackthrough
membraneboundATPsynthases,makingATP
WhenO2 isfinalelectronacceptor,themostATPisproduced
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RegeneratingNAD+
Somefermentationproducts(Fig.5.23)
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Fig.5.13
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Photosystemsandchlorophylls(Figs.5.26
and5.27)
Photosynthesisoverview
2partsofphotosynthesis,onelight
dependent(photophosphorylation)andthe
othernot(CO2 fixation,CalvinBenson
cycle)
Photophosphorylationuseslightto
Photophosphorylation uses light to
generatehighenergyesthatgenerateATP
inaprocesssimilartorespiration
TheATPfromthelightdependent
reactionsisusedtodrivethereductionof
CO2 toform6Csugarsfrom5Csugars
CO2 fixationalsorequiresNADPH
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Photosystemsarefoundon
specialmembranescalled
thylakoids
Photosystemscontain
chlorophyllmolecules
embedded in sea of light
embeddedinseaoflight
harvestingpigments
Lightenergycollectedand
transferredtochlstoacctivate
electrons
Highenergyestransferred
fromchlsthroughETStp
produceH+gradandthenATP
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Cyclicphotophosphorylation(Fig.5.28a)
Noncyclicphotophosphorylation(Fig.5.28b)
ATPisgeneratedfromprotonmotiveforcegeneratedas
electronsmovethroughETS
Thefinaleacceptoristheinitialdonorinacycle
TheNADPHneededforCO2fixationmustcomefrom
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somewhereelse
2lightdependentstepsareneededtoelevate
lowenergyesfromH2Otohighenoughlevelto
generatePMFandgenerateATP+NADPH
O2 isthebiproductwhenH2Oise donor
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Table5.5
Calvincycle(Fig.5.29)
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CalvinBensoncycle
UsesRuBisCotogenerate2x3Cmolecules
pereachCO2 moleculeaddedtothe5C,
ribulosebisphosphate
Cyclerequires3ATPand2NADPHforeach
CO2 fixed(andreduced)
fixed (and reduced)
EnergyfromtheATPandNADHisstored
andcanlaterberecoveredfromthesugars
produced
G3Pfeedsdirectlyintoglycolysis
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