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Respiration and Fatty Acid Metabolism

What do plants do when light is not available for photosynthesis?


Respiration [glycolysis, Oxidative Pentose phosphate shunt,
Citric Acid (Krebs) Cycle]
Products for Calvin cycle
NADH, FADH2, and ATP
Lipid metabolism
Respiration
Glucose equivalents (sucrose, triose phosphates, fructans, lipids,
etc.) oxidized to form CO2 and H2O
Glycolysis oxidizes sugars, via hexose phosphates and
triose phosphates, to organic acids (e.g. pyruvate) and yields
NADH and ATP
Oxidative pentose phosphate pathway uses glucose
phosphate to yield ribulose 5-phosphate and NADPH, then
various interconvertible sugars
Citric Acid (Krebs) Cycle breaks pyruvate down to CO2 and
yields NADH and FADH2
Electron transport chain and oxidative phosphorylation
reduces O2 to water, producing ATP and NAD+
12.1 Overview of respiration
12.2 Structures and reactions of major electron-carrying
cofactors (Part 1)

12.2 Structures and reactions of major electron-carrying


cofactors (Part 2)

Reactions of plant glycolysis and fermentation (1)


12.3 Reactions of plant glycolysis and fermentation (Part 2)

12.3 Reactions of plant glycolysis and fermentation (Part 3)

12.4 Reactions of the oxidative pentose phosphate pathway in


higher plants
The Oxidative Pentose phosphate shunt occurs in
the cytosol and especially plastids
-alternative to glycolysis
-Supplies NADPH for redox and respiration
reactions
-produces substrates for biosynthetic pathways
and Calvin cycle
-Primarily occurs in the dark

-Many of the same enzymatic carbon-shuffling


processes and intermediates involved in the
Regeneration phase of the Calvin Cycle
12.5 Structure of plant mitochondria
The Citric acid (TCA) cycle occurs in mitochondria
-pyruvate transported into mitochondrial matrix
12.6 Reactions and enzymes of the plant citric acid cycle
12.7 Malic enzyme and PEP carboxylase provide plants with
metabolic flexibility (Part 1)

12.7 Malic enzyme and PEP carboxylase provide plants with


metabolic flexibility (Part 2)

12.7 Malic enzyme and PEP carboxylase provide plants with


metabolic flexibility (Part 3)

Electron transport
Complex I, NAD(P)H
dehydrogenases, and Complex II
feed electrons to a pool of
ubiquinones
Complex I pumps 4 H+ out of the matrix
for each e- transferred
Ubiquinone passes e- to Complex III
and, via Cytochrome c, on to
Complex IV
Both complexes pump H+ out of the
matrix
Complex IV uses O2 as the e- acceptor
to produce H2O. Poisoned by cyanide.
The alternative oxidase can also accept
e- from ubiquinone to reduce O2 to H2O,
but does not have the accompanying H+
transport. Inhibited by SHAM
ATP synthase (complex V) functions
like the chloroplast ATP synthase,
but is driven more by charge
difference (ΔE) than H+ gradient.
NADH and NADPH are oxidized as the source of e- .
NAD(P)+ reduced by Glycolysis, Krebs cycle, PSI
Figure 12.10 Transmembrane transport in plant mitochondria
Aerobic respiration
Yields approximately 60 ATP per sucrose
52 from oxidative phosphorylation via 20 molecules
NAD(P)H and 4 FADH2 molecules
8 from substrate-level phosphorylation
Represents 52% energy efficiency (vs. 4% from
glycolysis/fermentation)
Plant mitochondria
Genome is much larger than in animals and fungi
RNA splicing occurs on a limited basis
RNA editing occurs (~ C  U)
RNA stability signals differ
Use the universal genetic code (unlike other organisms’
mitochondrial genomes)
Plants can decrease efficiency
Why?
Energy may not be limiting
Plasticity for use of metabolites may be
more important
Alternative oxygenase reduces O2 to
water, siphoning off excess electrons
and producing heat
Stresses that increase Reactive Oxygen
Species (ROS) activate alternative
oxygenase to prevent overreduction
Uncoupling protein relieves the H+
gradient without producing ATP
Non-proton-pumping NADH
dehydrogenases when ADP is
limiting
ADP and Pi are major regulators of
respiration
Figure 12.11 Metabolic interactions between mitochondria and
cytosol
11.11 Metabolic regulation of pyruvate dehydrogenase (PDH)
activity
Regulatory kinases and phosphatases control
Pyruvate dehydrogenase, coupling energy status
and metabolites to activity

Thioredoxins alter S—S bridges to regulate


enzymes according to redox status

11.12 Concept of bottom-up regulation of plant respiration


Allosteric “bottom up” regulation of
pathways by ADP and by metabolic
products allows for fine control of
respiration and use of metabolites for
biosynthetic pathways
Figure 12.14 Glycolysis, the oxidative pentose phosphate shunt,
and the citric acid cycle contribute precursors to many
biosynthetic pathways in plants
Plants respire even during photosynthesis
Metabolism of photorespiratory products (e.g. glycine to serine
conversion, N recovery)
Non-photosynthetic/sink tissues
Provide substrates for numerous biochemical reactions, e.g.
2-oxoglutarate for nitrogen assimilation
NADH production

Lipids in plants
Triacylglycerols
Storage fats and oils
High metabolic energy content
Polar glycerolipids
Membrane components

11.14 Structural features of triacylglycerols and polar


glycerolipids in higher plants
Oils and Fats
Oils often contain unsaturated fatty acid chains
Remain liquid at room temp
Stored in oleosomes with a half lipid bilayer
Fats generally have saturated straight-chain fatty acids
Produced in the ER and accumulate between the bilayer layers
until an oleosome (oil body) buds off
Abundant in seed cotyledons and endosperm

Fatty acid chains

Polar glycerolipids
Primary membrane structural lipids
Glyceroglycolipids
Sugar head group
Abundant in chloroplast membranes
Glycerophospholipids
Phosphate on head group
Sphingolipids, sterols, and other lipids are also membrane
components
Plastoquinones, chlorophylls, carotenoids, tocopherols
involved in photosynthesis and other functions
Very abundant in photosynthetic tissues

11.15 Major polar lipids of plant membranes (Part 1)


11.15 Major polar lipids of plant membranes (Part 2)

Fatty acid metabolism


Sequential addition of 2C acetyl in plastids
Condensation reaction with Malonyl-ACP (acyl-carrier
protein) results in decarboxylation and addition of 2C at each
sequential step
Major products are 16:0- or 18:0-ACP and, after modification by a
desaturase, 18:1-ACP
May be further desaturated to 16:3 and 18:3 fatty acids in
chloroplasts and ER
Fatty acid metabolism
Lipid components alter membrane characteristics
Desaturation maintains fluidity at lower temperatures
Different lipids affect other membrane functions
Lipids and metabolites serve as signals
PIP2 (phosphatidyl inositol 4,5-bisphosphate) breaks down to
IP3 and DAG (diacylglycerol)
11.16 Cycle of fatty acid synthesis in plastids of plant cells
Figure 11.17 The two pathways for glycerolipid synthesis

Fatty acid metabolism


Breakdown of fatty acids yields numerous metabolites including
sugars
Triacylglycerols from lipase action on oils enter glyoxysomes
(specialized peroxisomes)
Serial β-oxidations produce Acetyl-CoA for the glyoxylate
cycle to yield succinates for conversion to malate in the
mitochondria
Malate is converted through oxaloacetate to PEP, and to
sucrose

Figure 12.19 Conversion of fats into sugars during germination in


oil-storing seeds (Part 1)
Figure 12.19 Conversion of fats into sugars during germination in
oil-storing seeds (Part 2)