1/23/2011. Grapevine Anatomy & Physiology. What is Light? WSU Viticulture Certificate Program. Photosynthesis & Respiration.

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1 WSU Viticulture Certificate Program Grapevine Anatomy & Physiology & Respiration Markus Keller PHOTOS: Converts sunlight to chemical energy SYNTHESIS: Uses energy to convert inorganic compounds to organic compounds in the plant What is Light? Electromagnetic radiation moving at c = 300,000 km/s Waves: wavelength (λ), frequency (ν) Particles: quanta (photons), which contain energy Energy: E = h*ν ν = h*c/λ Visible light: PAR photosynthetically active radiation nm 1

2 2 Stages inside chloroplasts 1. Thylakoid membranes : Sunlight ATP & NADPH (adenosine-5 -triphosphate -triphosphate & nicotinamide adenine dinucleotide phosphate) 2. Chloroplast stroma: Enzymes: + H 2 O sugar (carbohydrate) Light Interception e - excited chlorophyll Unstable!: 10-9 s M g The Z Scheme PSII PSI 2

3 Energy Generation Photophosphorylation Antenna pigments & Reaction center ph difference proton gradient Waste product In and Out: Uptake Diffusion rate: driving force (= concentration gradient) & resistance (stomata + membranes!) [ ] outside = 0.036% [ ] inside > 0% [H 2 O] outside = 1.25% (50% rh) [H 2 O] inside = 2.5% (99% rh) > H 2 O Dilemma: stomata open in H 2 O out Calvin cycle in stroma Consumes ATP & NADPH in sugar out 13 Enzyme steps in 3 stages The Production Line H 2 O C 6 H 12 O 6 + 6O 2 + 6H 2 O ΔG = +2.8MJ/mol Rubisco = key enzyme 3 Turns per triose phosphate 5 Triose phosphates recycled & 1 available for sugar production Rubisco 3

4 Photorespiration Problem 1: [ ] in atmosphere 0.036%, but [O 2 ] 21% Problem 2: rubisco is carboxylase/oxygenase conflict of interest! Carboxylase: in O 2 out & fixed C available for sugar production Oxygenase: O 2 in out & fixed C lost to sugar production Partial recovery through glycolate pathway across 3 cell organelles Complex, costs energy, wastes heat Releases NH 4+ detoxification Reduces photosynthetic efficiency (25% lost) Evolutionary hangover? Products Photosynthate: organic products of photosynthesis Building blocks: sugar (triose) phosphates Production sites: chloroplast & cytosol Transport: phosphate translocator (triose phosphate out P i in) Carbohydrate: (CH 2 O) n, n = number of units Hexose: glucose, fructose (monosaccharide, n = 6: C 6 H 12 O 6 ) Export product: sucrose (disaccharide, n = 12) Storage product: starch (glucose polysaccharide) Structural product: cellulose (glucose polysaccharide) Export vs. Storage Sucrose Main photosynthesis product (glucose + fructose) Produced in leaf cell cytosol for export or storage in vacuole Buffer (short-term) against photosynthesis fluctuations Osmotically active attracts water pressure! Feedback inhibition with P i as regulator: low [P i ] = traffic jam Starch Surplus photosynthesis product (1000s of glucoses) Amylose (linear) and Amylopectin (branched) Produced in leaf cell chloroplast for storage as grains (growth rings) Osmotically inactive Overflow mechanism for rapid photosynthesis or stuck sucrose export Buffer (medium-term) against stuck photosynthesis (night!) 4

5 The Power Plant C 6 H 12 O 6 + 6O 2 + 6H 2 O H 2 O ΔG = -2.8MJ/mol Respiration burns sugar (fuel: sucrose or starch) to and H 2 O Releases energy stored in sugar & incorporates it into usable form (ATP, NADH) ATP equivalents per sucrose Glycolysis (cytosol): glucose (or fructose) pyruvate + ATP + NADH TCA cycle (mitochondria): pyruvate + H 2 O + NADH Recycling of oxaloacetate for next round Oxidative phosphorylation (mitochondrion membrane): e - transport chain for ADP + P i ATP Modifies sugar (building blocks: sucrose or starch) Produces carbon skeletons as building blocks for metabolism Supplies material for production of cellulose, DNA, amino acids, proteins, phenolics, lignin, lipids, chlorophyll, carotenoids, terpenes, hormones Respiration for All From Cells to Plants Respiration = growth respiration + maintenance respiration Growth respiration: sugar as fuel for new biomass production (amino acids ) Maintenance respiration: energy to keep cells alive Respiration consumes carbon from photosynthesis th Compromise between energy generation and carbon gain Leaves respire at 5-10% of photosynthesis, but other tissues respire 24/7 Young vines respire 30% of daily photosynthesis, older vines more Highest respiration in tissues with most metabolic activity (buds, expanding leaves, growing root tips, flowers) 5

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