Lecture 12 (2009 web) - Plant Sciences Home 12 (2009).pdf · Lecture 12 • Carbon assimilation in photosynthesis (Calvin Cycle) • Regulation of gluconeogenesis/glycolysis • Synthesis

Lecture 12Lecture 12

• Carbon assimilation in photosynthesis(Calvin CycleCalvin Cycle)

• Regulation Regulation of gluconeogenesis/glycolysis

• Synthesis of other carbohydrates

• The Pentose Phosphate PathwayPentose Phosphate Pathway

CARBOHYDRATES

p. 21

Glucose

Lactate

Ribose-5-P

Oxaloacetate

Amino acids

Ketonebodies

Cholesterol

Fatty acids

ATPNADHPyruvateGlucose-6-PGlycogen Acetyl-CoA

NADPH NADH

LIPIDSPROTEINS

The The ““PowertrainPowertrain”” of Human Metabolism (Overview)of Human Metabolism (Overview)

OtherCarbohydrates

GNGGNG

CARBOHYDRATES

p. 21

Glucose

Lactate

Ribose-5-P

Oxaloacetate

Amino acids

Ketonebodies

Cholesterol

Fatty acids


NADPH NADH

LIPIDSPROTEINS


OtherCarbohydrates

GNGGNG

H

CHO

OH

HHO

OHH

OHH

CH2OH

H

CHO

OH

HHO

OHH

OHH

CH2OPO32-

GLC-6-PGLC

ATP ADP

CH2OH

O

HHO

OHH

OHH

CH2OPO32-

ATP ADP

CH2OPO32-

O

HHO

OHH

OHH

CH2OPO32-

CH2OPO32-

O

CH2OH

H

HC

OH

CH2OPO32-

+

NAD+

NADH

H

C

OH

CH2OPO32-

OOPO3

2-

ATP ADP

H

C

OH

CH2OPO32-

O

H

C

OPO32-

CH2OH

OC

OPO32-

CH2

OC

O

CH3

O

ATP ADP

DHAP

G-3-P

1,3 bisPGA3-PGA2-PGAPEPPYR

1 2 3 4

5

6

78910

H O

OHH

OHH

OH

CH2OHH O

OHH

OHH

OH

CH2OPO32-

ATP ADP

1 2

O3POH2C

OH

CH2OH

HOH H

H OHO

O3POH2C

OH

CH2OPO32-

HOH H

H OHO

ATP ADP

3

Ringform

O

F-1,6-bisP

+ H+

1

2

3

4

5

6

1

2

3

4

5

6

1,6

2,53,4

Haworthprojection

HPO42-

-O-O-O-O

F-6-P

H2O

OH OH

2 Molecules

H

CHO

OH

HHO

OHH

OHH

CH2OH

H

CHO

OH

HHO

OHH

OHH

CH2OPO32-

GLC-6-PGLC

ATP ADP

CH2OH

O

HHO

OHH

OHH

CH2OPO32-

ATP ADP

CH2OPO32-

O

HHO

OHH

OHH

CH2OPO32-

CH2OPO32-

O

CH2OH

H

HC

OH

CH2OPO32-

+

NAD+

NADH

H

C

OH

CH2OPO32-

OOPO3

2-

ATP ADP

H

C

OH

CH2OPO32-

O

H

C

OPO32-

CH2OH

OC

OPO32-

CH2

OC

O

CH3

O

ATP ADP

DHAP

G-3-P

1,3 bisPGA3-PGA2-PGAPEPPYR

1 2 3 4

5

6

78910

H O

OHH

OHH

OH

CH2OHH O

OHH

OHH

OH

CH2OPO32-

ATP ADP

1 2

O3POH2C

OH

CH2OH

HOH H

H OHO

O3POH2C

OH

CH2OPO32-

HOH H

H OHO

ATP ADP

3

Ringform

O

F-1,6-bisP

+ H+

1

2

3

4

5

6

1

2

3

4

5

6

1,6

2,53,4

Haworthprojection

HPO42-

-O-O-O-O

F-6-P

H2O

OH OH

2 Molecules

p. 25

GluconeogenesisGluconeogenesis:: ByBy--pass reactions and enzymespass reactions and enzymes

3. By3. By--passpass 2. By2. By--passpass

1. By1. By--passpass

Go’ = 0.9 kJ mol-1G = - 25 kJ mol-1

Go’ = +32 kJ mol-1G = + 23 kJ mol-1

First By-pass Reaction (Summary)

PyrPyr KinaseKinase

OAAOAA

Pyr PEP

+ATPCO2

+GTPCO2

Does not workDoes not work

PEPPEPCarboxykinaseCarboxykinase

PyruvatePyruvateCarboxylaseCarboxylase

StoichiometryStoichiometry of Glycolysisof Glycolysis ((G ca. G ca. -- 63 kJ mol63 kJ mol--11))

1 1 GlcGlc + 2 ADP + 2 Pi + 2 NAD+ 2 ADP + 2 Pi + 2 NAD++

2 2 PyrPyr + + 2 ATP2 ATP + 2 NADH + 2 H+ 2 NADH + 2 H22OO

2 2 PyrPyr + + 4 ATP4 ATP + + 2 GTP2 GTP + 2 NADH + 6 H+ 2 NADH + 6 H22O O

1 1 GlcGlc + 4 ADP + 2 GDP + 2 NAD+ 4 ADP + 2 GDP + 2 NAD++ + 6 Pi+ 6 Pi

StoichiometryStoichiometry of of GluconeogenesisGluconeogenesis ((G ca. G ca. --16 kJ mol16 kJ mol--11))

Extra Cost of Extra Cost of GluconeogenesisGluconeogenesis: : 4 ATP per glucose4 ATP per glucose

GlucoseGlucose

G6P F6P F1,6BP PEP Pyr

MitochondriaMitochondria

NAD+

NADH

Pyr

OAA

PEP

NAD+

Shared (reversible) reactionsShared (reversible) reactions

GlycolysisGlycolysis--specificspecific

GluconeogenesisGluconeogenesis--specificspecific

LactateLactate

LactateLactateGlucoseGlucose

CytosolCytosol

BloodBlood

FruFru--1,61,6--BPBPPhosphatasePhosphatase

GlcGlc--66--PPPhosphatasePhosphatase

LiverLiver

PyruvatePyruvate

Acetyl-CoA

Succinyl-CoA

-KetoglutarateFumarate

IsocitrateMalate

CitrateOxalaoacetate

(+) ADP(-) ATP, NADH, Citrate, Succinyl-CoA

(+) ADP, NAD+

(-) ATP, NADH

(+) AMP(-) ATP, NADH, Succinyl-CoA

(+) AMP, NAD+, CoA(-) ATP, NADH, Acetyl-CoA

PEPPEP

LactateLactate

GlucoseGlucose

TCA Cycle

Reciprocal Regulation of Reciprocal Regulation of GlycolysisGlycolysis and and GluconeogenesisGluconeogenesis

Pyr Kinase

PC

PEP-CK

(+) allosteric activation(-) allosteric inhibition

PyruvatePyruvate

Acetyl-CoA

Succinyl-CoA


IsocitrateMalate



(+) ADP, NAD+

(-) ATP, NADH



(+) Acetyl(+) Acetyl--CoACoA((--) ADP) ADP

ActivtationActivtation of of GluconeogenesisGluconeogenesis at at ““First ByFirst By--passpass””

PEPPEP

LactateLactate

((--) ADP) ADP

GlucoseGlucose

TCA Cycle


PyruvatePyruvate

Acetyl-CoA

Succinyl-CoA


IsocitrateMalate



(+) ADP, NAD+

(-) ATP, NADH



p. 49

PEPPEP

GlucoseGlucose

TCA Cycle

Activation of GlycolysisGlycolysis at “Pyruvate Kinase Reaction”

Pyr Kinase

(+) F1,6BP(+) F1,6BP((--) ATP) ATP((--) ) AlanineAlanine


PyruvatePyruvate

Acetyl-CoA

Succinyl-CoA


IsocitrateMalate



(+) ADP, NAD+

(-) ATP, NADH



PEPPEP

LactateLactate

GlucoseGlucose

TCA Cycle

Reciprocal Regulation of Reciprocal Regulation of GlycolysisGlycolysis and and GluconeogenesisGluconeogenesis

Pyr Kinase

PC

PEP-CK


p. 63

Phosphoenolpyruvate Pyruvate

ATPADP + HPO43-

Pyr kinase

(+) Acetyl CoA (-) ADP

(+) F1,6BP(-) ATP(-) Alanine

OAAPEP CK PC

(-) ADP

Phosphoenolpyruvate Pyruvate

ATPADP + HPO43-

Pyr kinase

(+) Acetyl CoA (-) ADP

(+) F1,6BP(-) ATP(-) Alanine

OAAPEP CK PC

(-) ADP

Reciprocal RegulationReciprocal Regulation of Glycolysis and of Glycolysis and GluconeogenesisGluconeogenesis((FirstFirst byby--pass)pass)

GlycolysisGlycolysis

GluconeogenesisGluconeogenesis


p. 63



Reciprocal RegulationReciprocal Regulation of Glycolysis and of Glycolysis and GluconeogenesisGluconeogenesis((SecondSecond byby--pass)pass)

Fructose-6-P Fructose-1,6-BP

ATP

H2OHPO43-

ADP + HPO43-

PFK-1

FBPase

(+) Citrate (-) AMP(-) F2,6BP

(+) F2,6BP(+) AMP(-) ATP(-) Citrate

Glucose Fructose-6-P Fructose-1,6-BP

ATP

H2OHPO43-

ADP + HPO43-

PFK-1

FBPase



Glucose


H

CHO

OH

HHO

OHH

OHH

CH2OH

H

CHO

OH

HHO

OHH

OHH

CH2OPO32-

GLC-6-PGLC

ATP ADP

CH2OH

O

HHO

OHH

OHH

CH2OPO32-

ATP ADP

CH2OPO32-

O

HHO

OHH

OHH

CH2OPO32-

CH2OPO32-

O

CH2OH

H

HC

OH

CH2OPO32-

+

DHAP

G-3-P

1 2 3 4

5

Ringform

O

F-1,6-bisP

1

2

3

4

5

6

1

2

3

4

5

6Haworthprojection

F-6-P

H

CHO

OH

HHO

OHH

OHH

CH2OH

H

CHO

OH

HHO

OHH

OHH

CH2OPO32-

GLC-6-PGLC

ATP ADP

CH2OH

O

HHO

OHH

OHH

CH2OPO32-

ATP ADP

CH2OPO32-

O

HHO

OHH

OHH

CH2OPO32-

CH2OPO32-

O

CH2OH

H

HC

OH

CH2OPO32-

+

DHAP

G-3-P

1 2 3 4

5

Ringform

O

F-1,6-bisP

1

2

3

4

5

6

1

2

3

4

5

6Haworthprojection

F-6-P


FF--2,62,6--bisPbisP

FF--2,62,6--bisP functions only as an allosteric effector!bisP functions only as an allosteric effector!

ATP

ADP H2O

Pi

Activity of Activity of bifunctionalbifunctional enzyme is reciprocally controlled enzyme is reciprocally controlled (hormones, metabolites)(hormones, metabolites)

Bifunctional Enzyme FF--2,62,6--bisPbisPPhosphatasePhosphatase

FF--66--P 2P 2--kinasekinase(PFK(PFK--2)2)

“Dead-end” metabolite

F6P Concentration

PFK

-1 A

ctiv

ity No allostericeffector

High Citrate or high ATP

+ F2,6BPHigh AMP + F2,6BP

Phosphofructokinase-1

F6P Concentration

PFK

-1 A

ctiv

ity No allostericeffector

High Citrate or high ATP

+ F2,6BPHigh AMP + F2,6BP

Phosphofructokinase-1

p. 64

p. 64

F1,6BP Concentration

FBP

ase

Act

ivity No allosteric

effector

High AMP or high F2,6BP

High Citrate

Fructose Bisphosphatase


FBP

ase

Act

ivity No allosteric

effector


High Citrate


FBP

ase

Act

ivity No allosteric

effector


High Citrate

Fructose Bisphosphatase

p. 63



Reciprocal RegulationReciprocal Regulation of Glycolysis and of Glycolysis and GluconeogenesisGluconeogenesis((SecondSecond byby--pass)pass)


ATP

H2OHPO43-

ADP + HPO43-

PFK-1

FBPase



Glucose Fructose-6-P Fructose-1,6-BP

ATP

H2OHPO43-

ADP + HPO43-

PFK-1

FBPase



Glucose


PFK-1

FBPase

(+) Citrate(-) AMP(-) F2,6BP

(+) F2,6BP(+) AMP(-) Citrate

Glucose-6-P

Flux Control by Substrate CyclesSubstrate Cycles


PFK-1

FBPase

(+) Citrate(-) AMP(-) F2,6BP

(+) F2,6BP(+) AMP(-) Citrate

Glucose-6-P

+ 80%

- 80%

Flux Control by Substrate CyclesSubstrate Cycles

GlucoseGlucose

G6P F6P F1,6BP PEP Pyr Pyr

OAA

PEP

F1,6BPaseF1,6BPase

G6PaseG6Pase

PyrPyr KinaseKinase

PCPC

PEPPEP--CKCK

TCATCACycleCycle

AcetylAcetyl--CoACoAHexokinaseHexokinase

PFKPFK--11

F2,6BP


ReciprocalReciprocal regulation by:regulation by:

•• F2,6BP (F2,6BP (activactiv. . // inhibinhib..))•• ATP (ATP (inhibinhib.. / / activactiv..))•• Citrate (Citrate (inhibinhib.. / / activactiv..))•• OtherOther


Reciprocal Reciprocal regulation by:regulation by:

•• ATP (ATP (inhibinhib.. / / activactiv..))

3. By-pass

Hexokinase(Cytosol)

G6Pase(Endoplasmicreticulum)

Regulation

Feedback Control (“Acceptor Control”)

Glucose

Lactate


ADPPFK-1PK PDH

CSIDH

KGA-DH ETC

Allosteric ActivationAllosteric Activation

Allosteric InhibitionAllosteric Inhibition

CARBOHYDRATES

p. 21

Glucose

Lactate

Ribose-5-P

Oxaloacetate

Amino acids

Ketonebodies

Cholesterol

Fatty acids


NADPH NADH

LIPIDSPROTEINS

Synthesis of Other CarbohydratesSynthesis of Other Carbohydrates

OtherCarbohydrates

GNGGNG

Activation of Sugars for BiosynthesisActivation of Sugars for Biosynthesis

1. Sugar phosphate + NTP NDP-Sugar + Pyrophosphate (PPi)

HOWEVERHOWEVER, the first reaction is sequentially coupled to the hydrolysis o, the first reaction is sequentially coupled to the hydrolysis of f PPiPPi2. Pyrophosphate (2. Pyrophosphate (PPiPPi) + H) + H22O O 2 Phosphate (2 Pi)2 Phosphate (2 Pi)

GlucoseGlucose--11--P + UTP P + UTP UDPUDP--Glucose + Glucose + PPiPPi (see page 37)

NDPNDP--Group:Group: -- NNucleoside ucleoside DDiiPPhosphatehosphate is typically UDP or ADPis typically UDP or ADP

-- NDPNDP--sugar formation is sugar formation is irreversible irreversible ((PPiPPi hydrolysis)hydrolysis)

-- Serves as a Serves as a ““recognition tagrecognition tag”” for biosynthesesfor biosyntheses

-- Facilitates Facilitates substrate bindingsubstrate binding

-- Excellent Excellent leaving groupleaving group in in nucleophilicnucleophilic reactionsreactions

O

OH OPO32-

OH

OH

CH2OH

O O

OHOHH

HN

N

O

O

OPO

O-PO

O-

O O

OHOHH

HN

N

O

O

OPO

O-PO

OO-

O

OHOH

OH

CH2OH

O O-P

O

O-

P

O

-O

O-

Pyrophosphate (PPi) UDP-Glc

+

+

-D-Glc-1-P

OPO

-OO-

25

UTP

26

2 x Phosphate (Pi)

H2O

p. 37

Source of UDPSource of UDP--GlucoseGlucose

p. 65

O O

OHOHH

HNN

O

OOP

O

O-PO

OO-

OOH

OHOHCH2OH

H O HH

OHH

OH

CH2OHH O H

H

OHH

OH

CH2OHH O H

H

OHH

OH

CH2OH

HO O O O

Reducing "end"

14

Primer

UDP-Glucose

O

OHHO

HO

CH2OH

H O

OH

HH

OHH

OH

CH2OHH O H

H

OHH

OH

CH2OHH O H

H

OHH

OH

CH2OHH O

OH

OHH

OH

CH2OH

O O O14

Amylose

UDP

H+

Glycogen and Starch BiosynthesisGlycogen and Starch Biosynthesis

Branching Enzyme

1234567

123

4567

Glycogen or Starch (Amylopectin)

p. 65

Glycogen and Starch BiosynthesisGlycogen and Starch Biosynthesis

GlucoseGlucose GlcGlc--66--PP

GlucoseGlucose

CytosolCytosol

BloodBlood

Glc-1-P

UDPUDP-Glc

GlycogenGlycogen

Glycogen Glycogen SynthaseSynthase

To GlycolysisTo Glycolysis

Glycogen Glycogen PhosphorylasePhosphorylase

Skeletal MuscleSkeletal Muscle LiverLiver

UTPUTP

PPiPPi2Pi2Pi

p. 66

Cellulose BiosynthesisCellulose Biosynthesis(1(14) 4) glycosidicglycosidic bondsbonds

p. 66

Cellulose BiosynthesisCellulose Biosynthesis

p. 66

Cellulose Biosynthesis

Cellulose Biosynthesis

O O

OHOH

HN

N

O

O

OPO

O-PO

OO-

OHO

OH

OH

CH2OH

OH

O

HO

HO

OH

CH2OH

H O

H

OHH

OH

CH2OH

O

OH O

OH

OH

CH2OH

glucosyl

galactosyl OH

Glucose

UDP-Galactose

UDP

Lactose

+

+

p. 67

Lactose BiosynthesisLactose Biosynthesis

p. 67

Sucrose BiosynthesisSucrose Biosynthesis

H O

OH

O

H

OHH

OH

CH2OH

2-O3POH2C

CH2OHHOH H

H OHO

H O

OH

O

H

OHH

OH

CH2OH

HOH2C

CH2OHHOH H

H OHO

glucosyl

fructosyl

glucose (1 2) fructose

Sucrose Phosphate Sucrose

O O

OHOHH

HN

N

O

O

OPO

O-PO

OO-

O

OHOH

OH

CH2OH

UDP-Glucose

2-O3POH2C

CH2OHHOH H

H OHO

H2O

UDP

Fructose-6-P

CARBOHYDRATES

p. 21

Glucose

Lactate

Ribose-5-P

Oxaloacetate

Amino acids

Ketonebodies

Cholesterol

Fatty acids


NADPH NADH

LIPIDSPROTEINS


OtherCarbohydrates

PPPPPP

The PentosePentose--Phosphate PathwayPhosphate Pathway (PPP)

or

The HexoseHexose--monophosphatemonophosphate ShuntShunt

The Pentose Phosphate Pathway: The Pentose Phosphate Pathway: FunctionsFunctions

1. Production of NAD1. Production of NADPPHH

3. Biosynthesis/degradation of 3. Biosynthesis/degradation of other sugarsother sugars

(C(C33 CC44 CC55 CC66 CC7 7 sugars) sugars)

2. Biosynthesis and degradation of 2. Biosynthesis and degradation of riboseribose

Determined by cell-specific needs for NADPH, ATP, sugars

Roles for NADRoles for NADPPHH

1.1. ReductantReductant in in biosyntheticbiosynthetic reactionsreactions(e.g., fatty acid synthesis, steroid synthesis)(e.g., fatty acid synthesis, steroid synthesis)

4. Production of reactive oxygen species (ROS) 4. Production of reactive oxygen species (ROS) (e.g., innate immune response)(e.g., innate immune response)

3. Regulation of 3. Regulation of redoxredox homeostasishomeostasis(e.g., erythrocytes, detoxification reactions)(e.g., erythrocytes, detoxification reactions)

2. Separation of catabolic and anabolic steps2. Separation of catabolic and anabolic steps(e.g., in (e.g., in cytosolcytosol: : low NADHlow NADH, but , but high NADhigh NADPPHH))

The Pentose Phosphate Pathway: The Pentose Phosphate Pathway: ReactionsReactions(Cytosol)

1. Oxidation Phase 1. Oxidation Phase (C(C66 CC55 + CO+ CO22))

2. 2. IsomerizationIsomerization Phase Phase (C(C55 CC55))

3. Rearrangement Phase 3. Rearrangement Phase (xC(xC55 yCyC66))

p. 68

Phase 1: Oxidative ReactionsPhase 1: Oxidative Reactions

H O

OHH

OHH

OH

CH2OPO32-

H O

OHH

OHH

OH

CH2OPO32-

O

HCOO-

OHHHOOHHOHH

CH2OPO32-

HCOO-

OHOOHHOHH

CH2OPO32-

HHC OH

OOHHOHH

CH2OPO32-

NADP+ NADPH + H+

+

Glc-6-P 6-P-Glucono--lactone 6-P-Gluconate

Ribulose-5-P (Ru5P)

OH

H2O

CO2

NADP+ NADPH + H+

Phase I

Phase II

1 32

GlcGlc--66--P P DehydrogenaseDehydrogenase

LactonaseLactonase

66--PhosphogluconatePhosphogluconateDehydrogenaseDehydrogenase

6 Glc6 Glc--66--PP 6 Ru6 Ru--55--PP + 6 CO2 + 12 NADPH12 NADPH

Phase 1: Oxidative Reactions (Summary)Phase 1: Oxidative Reactions (Summary)

6 Ru6 Ru--55--PP

6 Pentoses

(30 Carbons)(30 Carbons)

5 Glc5 Glc--66--PP

5 Hexoses

(30 Carbons)(30 Carbons)

TrioseTriose

TetroseTetrose

PentosePentose

HexoseHexose

HeptoseHeptose

Phases 2 and 3: Phases 2 and 3: IsomerizationIsomerization and Rearrangement Reactions (Preview)and Rearrangement Reactions (Preview)

p. 68

Phase 2: Phase 2: IsomerizationIsomerization ReactionsReactions

Ribulose-5-P (Ru5P)

HHC OH

OOHHOHH

CH2OPO32-

HC OOHOHHOHH

CH2OPO32-

HHC OH

OOHHOHH

CH2OPO32-

HHC OH

OHHOOHH

CH2OPO32-

H

Ribose-5-P Xylulose-5-P (Xu5P)Ru5P Ru5P

Phase II

4 5

PhosphopentosePhosphopentoseIsoIsomerasemerase

PhosphopentosePhosphopentoseEpiEpimerasemerase

TransketolaseTransketolase

p. 68

Phase 3: CarbonPhase 3: Carbon--transfer or Rearrangement Reactionstransfer or Rearrangement Reactions

TransketolaseTransketolase

TransaldolaseTransaldolaseH

HC OH

OHHOOHH

CH2OPO32-

Xu5P R5P

HHC OH

OHHOOHH

Sedoheptulose-7-P

+ + C OHOHH

CH2OPO32-

GA3P

CH2OH

O

HHO

OHH

OHH

CH2OPO32-

OOHHOHH

CH2OPO32-

H

Erythrose-4-P (E4P)

+

OOHHOHH

CH2OPO32-

H

+

HHC OH

OHHOOHH

CH2OPO32-

Xu5P

CH2OH

O

HHO

OHH

OHH

CH2OPO32-

+C OHOHH

CH2OPO32-

HC OOHOHHOHH

CH2OPO32-

H

OHOHH

H

CH2OPO32-

6 7

6

GA3PE4P F6P

Phase III

F6P

CC33

CC55

CC55

CC55

CC66

CC66

Phase 3: CarbonPhase 3: Carbon--transfer Reactions (Summary)transfer Reactions (Summary)

2 Xu5P2 Xu5P + R5PR5P 2 F6P2 F6P + GA3PGA3P

4 Xu5P4 Xu5P + 2 R5P2 R5P 4 F6P4 F6P + 2 GA3P2 GA3P

2 Rounds

5 Glucose5 Glucose--66--PP

GluconeogensisGluconeogensis

Overall Stoichiometry of Pentose Phosphate Pathway

1 Glc-6-P + 12 NADP+ + 7H2O 6 CO2 + 12 NADPH + 12H+ + Pi

Overall Stoichiometry of Glycolysis/PDH/TCA Cycle

1 Glc + 6 H2O + 10 NAD+ + 2 FAD 6 CO2 + 10 NADH + 10H+ + 2 FADH2

(plus 4 ATP from substrate-level phosphorylation)

6 Glc-6-P + 12 NADP+ + 6H2O 6 CO2 + 12 NADPH + 12H+ + 4 F6P + 2 GA3P

6 Glc-6-P + 12 NADP+ + 7H2O 6 CO2 + 12 NADPH + 12H+ + Pi + 5 Glc-6-P

Documents

Lecture 12 (2009 web) - Plant Sciences Home 12 (2009).pdf · Lecture 12 • Carbon assimilation in photosynthesis (Calvin Cycle) • Regulation of gluconeogenesis/glycolysis • Synthesis