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39 Pages
Lect13
Course: CHEM 114B, Spring 2012School: UCSD
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Word Count: 757
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13 Lecture Pentose Phosphate Pathway 1 Important concepts 1. Memorize the intermediates of pentose phosphate pathway Understand how to make 6C and 3C from 5C And understand the mechanism Balance sheet 2. Understand regulation of pentose phosphate pathway Regulation on oxidation (NADP+ activates G6PD) Regulated by the need of the intermediates 3. Understand how NADPH removes reactive oxygen species...
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13 Lecture Pentose Phosphate Pathway
1
Important concepts
1. Memorize the intermediates of pentose phosphate pathway
Understand how to make 6C and 3C from 5C
And understand the mechanism
Balance sheet
2. Understand regulation of pentose phosphate pathway
Regulation on oxidation (NADP+ activates G6PD)
Regulated by the need of the intermediates
3. Understand how NADPH removes reactive oxygen species (ROS)
2
Pentose Phosphate Pathway
Required for generation of NADPH Also produces Ribose-5-phosphate (R5P) for nucleotide biosynthesis
3
Compartments
Pentose phosphate Pathway -- cytoplasm
4
NAD+ Capture energy from metabolism For ATP generation through oxidative phosphorylation NAD+/NADH 1000/1(favors metabolite oxidation) Selective dehydrogenases
NADP+ Capture energy from metabolism For reductive biosynthesis (fatty acids and cholesterol) NADP+/NADPH 1/100 (favors reductive biosynthesis) Selective dehydrogenases
5
6
Pentose Phosphate Pathway
Where does glucose 6phosphate come from?
7
First step of glycolysis -- Hexokinase
ATP
+
PO32-
+ ADP + H+ Glucose Glucose-6phosphate (G6P)
First step: Getting a phosphate on carbon #6.
8
Instead of second step of glycolysis, Phosphoglucose Isomerase Use Glucose-6phosphate dehydrogenase
9 9
Pentose Phosphate Pathway: Phase 1
Oxidation: electron acceptor NADP+ Glucose 6-posphate dehydrogenase specific for NADP+
10
Pentose Phosphate Pathway: Phase 1
Oxidation: electron acceptor NADP+ Hydrolysis
11
Pentose Phosphate Pathway: Phase 1
Oxidation: electron acceptor NADP+ Hydrolysis Oxidative decarboxylation: electron acceptor NADP+
5C
12
Problem now is how to get back to glycolysis
C5 (Ribulose-5- P)
C6 (Glucose-6-P )
?
+ CO2
Glycolysis
Glycolysis good at dealing with 6 carbon and 3 carbon compounds, but left with a 5 carbon here.
13
Convert Ribulose 5-phosphate
Same as Calvin cycle
Nucleotide biosynthesis
14
15
Convert Ribulose 5-phosphate
How to convert 5C to 6C + 3C? transketolase
5C
5C
3C
7C
16
Convert Ribulose 5-phosphate
How to convert 5C to 6C + 3C? transketolase + transaldolase
3C
7C
6C
4C
17
Compare with Calvin cycle:
How to convert 6C and 3C to 5C? transketolase + aldolase
4C
3C
7C
7C
18
Compare with Glycolysis: 4. Aldolase: Breaking up
Fourth step: Break 6 carbon sugar to 3 carbon sugars.19
Convert Ribulose 5-phosphate
How to convert 5C to 6C + 3C? transketolase + transaldolase + transketolase
4C
5C
6C
3C
20
Balance sheet: Input: 3 glucose 6-phosphate 6 NADP+ Output: 2 fructose 6-phosphate 1 GAP 6 NADPH 3 CO2
3
3
21
Enzymes
22
Important concepts
1. Memorize the intermediates of pentose phosphate pathway
Understand how to make 6C and 3C from 5C
And understand the mechanism
Balance sheet
2. Understand regulation of pentose phosphate pathway
Regulation on oxidation (NADP+ activates G6PD)
Regulated by the need of the intermediates
3. Understand how NADPH removes reactive oxygen species (ROS)
23
Regulation pentose of phosphate pathway
G6PD is activated by NADP+: Consume NADPH-> [NADP+] rises
Glucose 6-phosphate dehydrogenase Irreversible Rate-limiting
24
Regulation of pentose phosphate pathway
Much more ribose 5-phosphate needed than NADPH (e.g. cell division) => use Ribulose 5-phosphate from pentose phosphate. Also reverse transketolase/transaldolase to make ribose 5phosphate from F6P and GAP
25
Balance sheet: Input: 3 glucose 6-phosphate 6 NADP+ Output: 2 fructose 6-phosphate 1 GAP 6 NADPH 3 CO2
3
3
26
Regulation of pentose phosphate pathway
Balanced need for ribose 5-phosphate and NADPH => Generate 2 NADPH and 1 ribose 5-phosphate
27
Regulation of pentose phosphate pathway
Need NADPH more than ribose 5-phosphate (e.g. adipose tissue synthesizing fatty acids) Oxidization of glucose 6-phosphate Consumption of ribose 5-phosphate Regeneration of glucose 6-phosphate
28
Regulation of pentose phosphate pathway
Need NADPH and ATP Oxidization of glucose 6-phosphate Consumption of ribose 5-phosphate Glycolysis to generate ATP
30
Glycolysis vs. Pentose Phosphate
Glycolysis
3 Glucose
6 Pyruvate + 6 ATP + 6 NADH
Pentose Phosphate (if continued through glycolysis)
3 Glucose-6-Phosphate
Cost 3 ATP
1 GAP 2 F6P 3 CO2
2 ATP, 1 NADH, 1 pyruvate 2 x (3 ATP, 2 NADH, 2 pyruvate)
31 5 Pyruvate + 5 ATP + 5 NADH + 6 NADPH
Important concepts
1. Memorize the intermediates of pentose phosphate pathway
Understand how to make 6C and 3C from 5C
And understand the mechanism
Balance sheet
2. Understand regulation of pentose phosphate pathway
Regulation on oxidation (NADP+ activates G6PD)
Regulated by the need of the intermediates
3. Understand how NADPH removes reactive oxygen species (ROS)
32
Maintaining reduced state of molecules, salvage reactive oxygen species (ROS)
GSH
33
Hydroperoxide reduced
Glutathione oxidized
Hydrooperoxide damage hemoglobin and cleavage c-c bonds in the phospholipid tails of cell membrane
34
NADPH required to maintain reduced state
Electron transfer through bound FAD
Glucose-6-phosphate dehydrogenase deficiency
Particularly for red blood cells under oxidative stress, without mitochondria thus no alternative reductant
35
Glucose-6-phosphate dehydrogenase deficiency
Inherited on X chromosome Affects 400 million people worldwide Highest prevalence rates (5-25% gene frequency) found in tropical Africa, Middle East, tropical and subtropical Asia, some areas of the Mediterranean, and Papua New Guinea ~300 different variants, mostly amino acid substitutions Neonatal jaundice, acute hemolytic anemia
36
Reference slides
37
Mechanism of transketolase
Thiamine pyrophosphate ionization Attack ketose
Break C-C bond, Free aldose, Attach 2C
Release ketose
Attacks aldose substrate
38
Mechanism of transaldolase
Form schiff base protonate schiff base
Break C-C bond, Attach 3C
hydrolyze schiff base
deprotonate schiff base Attacks aldose substrate 39
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