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Transcript
Bil 255 Spring
photosynthesis
LITTLE GREEN MEN
Why do metazoan cells not photosynthesize ???
• chlorophyll vs. hemoglobin…
Bil 255 – Spring semester
–
–
–
–
–
(leghemoglobins)
mutant…… where hemoglobins can capture light energy
current photosynthetic rates = 20 mg hexose/dm2/ hr
average human surface area = 170 dm2
thus mutant hexose productivity = 40.8 gm/d
energy calculation… 1 mole glucose = 183 gm = 686Kc/mol
• thus 41 gm = 153 Kc/mol
• BMR = 2,000 Kc/d, or only about 8.5 % of what is needed
photosynthesis
Evolve = increase surface area (2000 dm2),
remain sessile, peristalsis becomes vestigal,
circulation replaced by “xylem & phloem” …
We are a plant
photosynthesis
255 Spring - Mallery
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Light driven phosphorylation production of ATP via photo-phosphorylation
Cellular process - bacteria, blue-green, and
eucaryotic cells with chloroplasts
Capture of light energy by pigments chlorophylls & accessory pigments
3
1st autotrophic cells probably used H2S as e- source
purple-sulfur bacteria of today
CO2 + H2S --> CH2O + H2O + 2S
- oxygenic photosynthetic procaryotes
CO2 + H2O --> CH2O + H2O + O2
van Neil equation [gs@SU] Phts is a REDOX reaction
CO2 + H2A --> CH2O + H2O + 2A
Mallery
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• reduction of NADP to NADPH via an ETS
DARK Reactions
(thermo-chemical reactions)
• CO2 fixation (reduction) stages
photosynthesis
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Morphological Basis of Photosynthesis…
Evolutionary Basis of Photosynthesis
photosynthesis
LIGHT Reactions (photo-chemical reactions)
• molecular excitation chl by light... charge separation
• generation of proton motive force (H+ gradient)
6CO2 + 12H2O* --> C6H12O6 + 6H2O + 6O*2
Reduction of CO2 to CH2O
cyanobacteria
2
– carboxylation CO2 RuBP --> 2 PGA
– reduction PGA + NADPH --> PGAL
– regeneration of RuBP via HMP path ---> RuBP
Capture e’s as reducing power in NADPH
255 Spring - Mallery
255 Spring - Mallery
2 Fundamental Reaction Mechanisms
PHOTOSYNTHESIS...
photosynthesis
photosynthesis
5
PLASTIDS - double unit membrane bound organelles
classified by pigment content (a functional classification)
PROPLASTIDS
the progenitor plastid found in MERISTEMATIC cells
1 to 3 µmeter dia; often 20+/cell; spherical to ellipsoid;
few primary thylakoids; does contain lipid droplets.
divide by fission-like process... give rise to all plastids
LEUCOPLASTS - defined by their storage function
amyloplasts - synthsize & store starch
aleuoplasts - contain stored protein (crystals)
[figure]
elaioplasts contain oil & fat globules - fat biosynthesis
CHROMOPLASTS - found in flower petals, ripe fruit, scenescent
leaves hold many water soluble anthocyanin pigments
of unknown function
photosynthesis
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1
Bil 255 Spring
photosynthesis
CHLOROPLAST
ubiquitious to all green plants within mesophyll cells *
defined by its containing chlorophyll…
SHAPE - variable (elipsoid, lenticular, stellate, convex; oblate spheroid)
SIZE - 2 to 3 um dia by 5 to 10 um long fig 14.31*
NUMBER - 15 to 20 per mesophyll cell [can be up to 400,000/cc]
VOLUME - often much larger than mitochondria - (apprx 30 um3)
CHLOROPLASM - (Stroma)
pyrenoids - which are starch coated protein granules [fig]
70s (prokaryotic size) ribosomes
naked DNA - 2 to 10fg DNA/chlp (equal to bacterial cell DNA,
highly supercoiled & repetitive - up to6 copies)
enzymes of CO2 fixation and lipid droplets
THYLAKOID* Membranes
(see fig's fig 14.29b*, 14.29c*, EM* & fig 14.30*)
photosynthesis
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Molecular Excitation of Chlorophyll
• Absorption of Light Energy
– blue light [440nm] = 71.5 Kc/einstein
– red light [700nm] = 40.9 Kc/einstein
• Ground State – paired e's with opposite spin = stability
– absorption moves non-bounded e's to higher
orbitals
• 1st excited singlet state
• 2nd excited singlet state
• 1st long-lived state
photosynthesis
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Photosynthetic Electron Flow
• Photosystem - chlorophylls, reaction center, primary acceptor
– PS 1 and
PS 2
– path of e- flow (cyclic vs. non-cyclic)
– release of O2
PIGMENTS
Accessory Pigments
Carotenoids- carotenes, xanthophylls
Phycobilins- chromophore + a protein
• phycoerythrin & phycocyanin
Chlorophylls - a,b,c,d, etc...... [side chain differences]
magnesium porphyrins with a phytol chain [polyene molecule]
• ABSORPTION SPECTRA plot of the amount of light
absorbed by a solution
vs. the wavelength of light
• ACTION SPECTRA - (Engelman)
plot of physiological activity
(e.g., O2 evolution, AP’s)
vs. the wavelength of light
photosynthesis
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FATES of Absorbed Light Energy
1. Re-radiated as vibrational heat
2. Re-radiated as fluorescence
emission of light of longer wavelength
700 nm --> 710nm in time frame 10-9sec less energetic
3. Re-radiated as phosphorescence
emission of light much longer wavelength
700nm --> 720nm in real time (1sec)
4. Induced resonance vibrational e excitation inducing like vibrations
in adjacent molecules causing their excitation
5. Photoionization enters into the photochmical reactions
loses electron to acceptor = ionized chl+
photosynthesis
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Dark Reactions of Photosynthesis
• occur in stroma (chloroplasm)
• consume ATP and NADPH made in light reactions
• reduces (fixes) CO2 into CH2O (sugars)
– capture of e- into coenzyme NADP+ ---> NADPH
• ATPase makes ATP (just like in mitochondria)
– chemiosmosis & its location in chloroplasts
3 different pathways to make sugar
C3 - CALVIN cycle
–
–
–
–
1 CO2 + 5C RuBP ---> (2) 3C sugars (PGA)
(2) 3C sugars combine ---> 1 net glucose
RuBP carboxylase [50% of leaf protein]
Photo-respiration
• inhibition by O2
photosynthesis
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photosynthesis
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2
Bil 255 Spring
photosynthesis
PHOTORESPRATION... RUBISCO catalyzes two different reactions:
a carboxylase activity… adding CO2 to ribulose bisphosphate an oxygenase activity… adding O2 to ribulose bisphosphate Karp fig 6.19*
C4 - Hatch & Slack pathway
– 4C acid ---> 3C + CO2 in bundle sheath
– 1 CO2 + 3C PGA --> 4C acid (mesophyll cells)
– CO2 into Clavin cycle (as above)
which one predominates depends on the relative
concentrations of O2 & CO2 with
high CO2 : low O2 favoring the carboxylase action
high O2 : low CO2 favoring the oxygenase action
The light reactions liberate O2 & more O2 dissolves in the cytosol of the cell
at higher temps. Thus, high light intensities & high temps (above ~ 30°C) favor
the oxygenase second reaction. The uptake
of O2 by RUBISCO forms two 3carbon molecules:
[reaction]
1) one is 3-phosphoglyceric acid [3PGA] just as in the Calvin cycle
2) the other is 2P-glycolate.
and involves 3 organelles: chloroplast, peroxisome, and mitochondria.
photosynthesis
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CAM Pathway (C4- Crassulacean Acid Metabolism)
photosynthesis
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- CAM plants...
are also C4 plants but do not separate C4 & C3 pathways in different parts
of the leaf (spatial), but rather separate them in time instead. CAM was 1st
studied in members of the plant family Crassulaceae.
At night,
CAM plants take in CO2 through open stomata at night, when the
succulents are in a cool environment. The CO2 joins with PEP to form the 4carbon oxaloacetic acid. OAA is converted to 4-carbon malic acid that
accumulates during the night in the central vacuole.
In the morning,
stomata close (thus conserving moisture as well as reducing the inward
diffusion of oxygen). Accumulated malic acid leaves the vacuole and is broken
down to release CO2 that is taken up into the Calvin (C3) cycle.
photosynthesis
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