FINAL EXAM: CARBS Flashcards
plants are versatile
they can:
use energy of sunlight to support biosynthesis
build organic compounds from CO2
move intermediates between cellular compartments
adapt to changing environments
assimilation of CO2 by plants
animal cells: use 3C intermediates (pyruvate, lactate) for synthesis — must eat it
plant cells: make 3C intermediates for further synthesis
using Co2 to make intermediates = carbon assimilation aka carbon fixation
3C intermediates for plants
make glyceraldehyde-3P (GA3P)
made from CO2, H2O, ATP, and NADPH from photosynthesis
plastids
organelles in plants and algae
enclosed by a double membrane
own small genome
can specialize
chloroplasts
photosynthesis
amyloplasts
starch storage
chromoplasts
pigment storage
elaioplats
lipid storage
carbon assimilation
occurs in the stroma of chloroplasts via Calvin cycle
once called dark reaction but runs under light
what does carbon assimilation require
ribulose 1,5-bisphosphate which is constantly regenerated using ATP energy and NADPH
carbon assimilation produces
3-phosphoglycerate, then glyceraldehyde-3P (GA3P) in quillibrium with DHAP
triose phosphates
CO2 is reduced in carbon assimilation
with oxidation of NADPH that was generated in the light reactions of photosynthesis
three stages of calvin cycle
- carbon assimilation
- 3-phosphoglycerate reduction
- regeneration
carbon assimilation stage
3 ribulose 1,5-bisophosphate + 3CO2
6 3-phosphoglycerate
rubisco catalyzes
(6c divided in half)
3-phosphoglycerate reduction stage
6 3-phosphoglycerates converted to 6 triose phosphates (reduction to DHAP and GA3P) using NADPH and ATP from photophosphorylation
5 triose phosphates go to regenerate ribulose 1,5-bisphosphate; 1 utilized for other pathways
3-phosphoglycerate reduction is catalyzed by
phosphoglycerate kinase and glyceraldehyde 3P dehydrogenase
regeneration stage
5 triose phosphates made to 3 ribulose 1,5-bisphosphate
overall calvin cycle reaction
3CO2 + 6NADPH + 5H2O + 9ATP
glyceraldehyde-3P + 6NADP+ + 2H+ + 9ADP + 8Pi
where can the glyceraldehyde 3P go from calvin cycle?
energy production via glycolysis, starch, or sugar synthesis
rubisco
catalyzes carbon assimilation
large Mg2+ containing enzyme
carboxylase and oxygenase functionality
carboxylase functionality in rubisco
adds Co2 to ribulose 1,5-bisphosphate
makes new CC bond
cleaves 6C intermediate into 2 3phosphoglycerates
oxygenase functionality in rubisco
less useful in plants
reacts with O2 instead of CO2 in an inefficient side reaction
rubisco reaction
ribulose 1,5-bisphosphate + H2O + CO2
2 molecules of 3phosphoglycerate
rubisco exits in two major forms: form 1
plants, algae, cyanobacteria
8 large catalytic subunits (encoded by plastid genome) + 8 small subunits (encoded by nucleus
rubisco exists in two major forms: form II
photosynthetic bacteria only
2 catalytic subunits, resemble large plant subunits
rubisco quality
inefficient
low turnover of 3/sec
50% of plant enzymes are rubisco
catalytic role of Mg2+ in rubisco’s carboxylase activity
Mg2+ is held by negatively charged side chains of Glu, Asp, and carbamoylated Lys
brings together the reactants in a correct orientation, stabilizes the negative charge that forms upon the nucleophilic attach of enediolate to CO2
carbamoylated Lys
negative CO2 binds to N on side chain of lysine
rubisco is activated via covalent modification
highly regulated
inactive until Lys-201 is carbamoylated by CO2 (CO2 binds to R group nitrogen; allows binding of Mg2+ to the enzyme which is critical for catalytic activity)
rubisco activase
rubisco activase
ribulose 1,5-bisP bound in active site blocks Lys-201 from being carbamoylated
rubisco activase changes rubisco conformation - causes ribulose-1,5-bisP to leave and exposes Lys-201
reaction requires ATP
triggered by light in some species
rubisco can be inhibited by a nocturnal inhibitor
2-carboxyarabinitol 1-phosphate inhibits carbamoylated rubisco
2-carboxyarabinitol 1P
inhibits carbamoylated rubisco;
transition state analog of beta-keto acid intermediate
synthesized in the dark in some plants
- binds to rubisco, doesn’t work in inappropriate times**
stage 2: 3-phosphoglycerate reduced to glyceraldehyde 3P reaction
6 3PG + 6ATP + 6NADPH + 6H+
6 GA3P + 6ADP + 6NADP+ + 6Pi
3PG reduced to GA3P
requires NADPH and ATP from photosynthesis
similar to gluconeogensis but uses NADPH
enzymes: 3phosphoglycerate kinase; glyceraldehyde 3phosphate dehydrogenase
driven forward by high conc. of NADPH and ATP in chloroplast stroma
enzymes for stage 2 calvin cycle
3PG kinase
GA3P dehydrogenase
phosphoglycerate kinase
3PG + ATP
= 1,3bisphosphoglycerate
glyceraldehyde-3P dehydrogenase
1,3-bisphosphoglycerate + NADPH
glyceraldehyde-3P
redox
fates of glyceraldehyde-3P
5 of 6 recycled to make ribulose 1,5-bisphosphate
remaining:
converted to starch in chloroplast for storage
converted to sucrose in cytosol for export
broken down in glycolysis in cytosol for energy
stage 3: regeneration of ribulose 1,5-bisphosphate reaction
3 GA3P + 2DHAP + 3ATP
3 ribulose 1,5,bisphosphate + 2Pi + 3ADP
5 three carbon sugars converted to 3 five carbon sugars
stoichiometry and energy cost of CO2 assimilation
fixation of 3 CO2 molecules yields one GA3P
9 ATP and 6 NADPH are consumed
light reactions of photosynthesis produce ATP and NADPH at about this same ratio (3/2)
fates of Pi from ATP hydrolysis in stages 2 and 3
8 of 9 Pi: available to combine with ADP to regenerate ATP
9th Pi: incorporated into GA3P and could be moved to cytosol
- requires that Pi be transferred from cytosol back into chloroplast
- uses special Pi-triosephosphate antiporter
Pi-triose phosphate antiporter
needed for sucrose synthesis
sucrose made in cytosol (unlike starch in chloroplast stroma)
- used for transport to distant plant tisuses
inner membrane is impermeable to phosphorylated compounds
antiporter exchanges GA3P or DHAP for one Pi
- sends triose phosphates into cytosol for sucrose synthesis
- sends Pi back into the chloroplast
carbon assimilation is more effective when its light
photosynthesis of one molecule of GA3P (plus recycled ones) requires 24 photons of light
- H+ go from stroma to thylakoid lumen
- creates alkaline conditions in stroma
Mg2+ transport from thylakoid lumen to stroma
enzymes of assimilation more active in alkaline, high Mg2+ conditions of stroma during photophosphorylation