10: Chloroplast Structure and Function from an inner to a higher/outer orbital shifting to

an excited state
 Heterotrophs are those than depend on an
 In isolated chlorophyll, the solution becomes
external source while autotrophs can
fluorescent because the absorbed energy is re-
manufacture their own nutrients
emitted at longer (low energy) wavelength
 Chemoautotrophs utilize chemical energy in
 In isolated chloroplasts, only a faint fluorescence
inorganic molecules
is observed. Very little of the energy absorbed is
 Photoautotrophs utilize the energy from the
dissipated but are instead transferred to electron
sun
acceptors
 During photosynthesis, low-energy electrons are
removed from a donor compound and converted
into high-energy electrons using the energy Photosynthetic Pigments
absorbed from light  Pigments appear colored because they only
 The switch from sulfur to oxygen was hard since absorb a light at a specific wavelength
sulfur has lower affinity for electrons  Chlorophyll absorbs strongly in the blue and red
 Polypeptides found in modern-day chloroplasts  Chlorophyll structure:
are encoded by both the chloroplast and nuclear o Porphyrin ring with Magnesium for light
genome absorption
 Conjugated (alternating single and
10.1 Chloroplast Structure and Function double bonds)
 Strong absorber of light
 Located predominantly in the mesophyll cells of
 Absorbed energy causes a redistribution
leaves
 Lens-shaped in higher plants, approx. 2-4 x 5-10 of the electron density of the molecule
o Hydrophobic phytol chain that keeps the
µm, 20 per cell
 Arise from fission from preexisting chloroplasts or chlorophyll embedded in the membrane
 Carotenoids absorb well in the blue and green
from non-pigmented precursors called
proplastids spectrum; act as secondary light collectors
 Were identified as the site of photosynthesis in during photosynthesis and draws excess energy
from chlorophyll as heat
1887 by T. Engelmann upon observing that
bacteria would aggregate near the chloroplast
 Contains two membranes separated by a narrow 10.4 Photosynthesis Units and Reaction Centers
space  Not all chlorophyll molecules in a chloroplast are
o Outer membrane contain porins involved in conversion of light into chemical
o Inner membrane is highly impermeable; energy
substances need the help of transporters  Chlorophyll molecules act together as one
 An internal membrane system separate from the photosynthetic unit in which only one member of
bi-layer is organized into sacs called thylakoid, the group, reaction-center chlorophyll
stacked into grana actually transfers electrons to an electron
o Space inside the thylakoid is the lumen acceptor
o Space within the chloroplast is the stroma  These pigment molecules act as an antenna that
 Stroma contains small double-stranded circular absorbs photons of various wavelengths and
DNA and prokaryotic-like ribosomes transfers that energy (excitation energy) to the
 Thylakoid membranes have high protein content reaction-center
and low phospholipid but instead have high  Energy can only be transferred to an equal or
percentage of galactose-containing glycolypids less energy-requiring molecule

10.2 An Overview of Photosynthetic Metabolism Oxygen Formation: PSI & PSII
 Photosynthesis essentially is a redox reaction  Light-absorbing reactions take place in protein-
o H2A (S/O) is an electron donor; reducing complexes called photosystems
agent  Photosystem II (PSII) boosts the electron to a
o CO2 is an oxidizing agent midway point; P680
 Each molecule of O2 is derived from the  Photosystem I (PSI) raises the electrons from a
breakdown of two molecules of H2O midway point to an energy level above NADP+
 Photosynthesis is then the reverse of  After losing electrons, PS becomes positively
mitochondrial respiration charged which attracts electrons
o Mitochondrial respiration reduces oxygen to  Z Electron flow: water  PSII  PSI  NADP+
water  Electron transfer releases energy which is used to
o Photosynthesis oxidizes water to oxygen establish a proton gradient to drive the synthesis
 light-dependent reactions: energy from the sun is of ATP needed for the synthesis of carbohydrates
absorbed and stored as ATP and NADPH
 light-independent reactions: carbohydrates are PSII Operations
synthesized using the stored energy  PSII uses absorbed energy to remove electrons
from water and generate a proton gradient
10.3 The Absorption of Light  D1 & D2 bind the P680 reaction-center with other
 light travels in energy called photons; the energy cofactors
depends on the wavelength of light  Light-harvesting complex II (LHCII) contain the
o The shorted the wavelength, the higher antenna and are situated outside the core of the
energy content photosystem; can migrate along the thylakoid
 When a photon is absorbed by a molecule, an membrane
electron becomes too energetic to be pushed  Formation of a single oxygen requires 4 photons
to be absorbed by PSII

cytochrome b6f translocate a net of 4 protons phosphoglycerate (PGA) 4. Thus. one at a time to PSII 10. the making it the most abundant protein on earth primary electron acceptor creating a separation  Cycle comprises of 3 main steps of charges: P700. 2-3 is repeated forming plastoquinol from the  Movement of protons during electron transport is joined molecules (PQH2). Electrons are passed to chlorophyll a. Electron from PQA is then transferred to another stroma PQB to produce a semi-reduced molecule that o Base: CF0 remains firmly bound to the D1  Protons move from the lumen to the stroma 4. net of 8 electrons are needed  Proton gradient is formed via: o Splitting of water o Oxidation of plastoquinol o Reduction of NADP+ and PQ Practical Application: Killing Weeds  Bind to open Q site.5 Photophosphorylation separation of charge  Conversion of one mole of CO 2 to one mole of  P680+: oxidizing agent carbohydrate requires 3 moles of ATP and 2  Pheo-: reducing agent NADPH 2. 12 GAP are reductase which requires 2 ferredoxin molecules produced o 10 are recycled to form 6 RuBP Not all electron passed to ferredoxin ends up in NADP o 2 are products reduction. Electrons are transferred to plastocyanin  RuBP + CO2  6 Carbon  fragmented into 2 situated on the luminal side which carries molecules of 3-PGA electrons to PSI  Catalyzed by ribulose bisphosphate carboxylase oxygenase (rubisco) which is only capable of PSI Operations fixing three molecules of CO2 per second.  GAP can be: o Exported to the cytosol in exchange of Photosynthetic Electron Transport phosphate ions to synthesize sucrose  Electrons travel from water to NADP+ via the 2 (~glucose) photosystems o Remain in the chloroplast where it is  PSII  produce Oxygen converted to starch (~glycogen)  PSI  produce NADP  Each photosystem requires 4 electrons.  Paraquat competes for electrons with ferredoxin PSII  Platoquinone but also creates free radicals 1. P680 transfers received electron to pheophytin. Some are used for other reductions. derived from the stroma (going in) and thus proton gradient is mainly due to pH only creates a gradient  Non-cyclic phosphorylation: electrons move 6. PQH2 then diffuses into the lipid bilayer linearly from water to NADP+  Cyclic phosphorylation: carried out by PSI Water  PSII independent of PSII  The redox potential of P680 is strong enough for 1. PQH2 donates 2 electrons to cytochrome b6f and C3 plants/Calvin Cycle translocate 2 protons to the lumen  First molecule to be discovered was 3- 3. P680 is reduced back 5. killed cells From PSII to PSI immediately (stopping enzyme activity) and 1. Energy passed to reaction center contain as much as half of its proteins as Rubisco 2. the primary electron acceptor creating a 10.(reducing agent) o Carboxylation of RuBP forming PGA 3.and Ao.6 Carbon Dioxide Synthesis  Melvin Calvin: radioactive Carbon. blocking transport through PSII . This is PSI is made of a core + LHCI the lowest turnover for any enzyme so plants 1. The protons used are neutralized by the movement of other ions. Electron is transferred out via ferredoxin NADPH and ATP o Regeneration of RuBP via ATP NADP+ reduction is done via ferredoxin-NADP+  For every 6 molecules of CO 2 . Electron is routed back to the electron-  Formation of one molecule of oxygen requires 4 deficient reaction-center electrons from 2 molecules of water **cyclic phosphorylation drives enough energy to  Oxygen-evolving complex: 4Mn + 1Ca build the proton gradient necessary** accumulates four oxidizing equivalents by transferring 4 electrons. PSI absorbs light    passed to ferredoxin photolysis of water 2.transfers its electron to plastoquinone  ATP synthase bound near the membrane o Head: CF1 which projects outward to the 3. Ao is passed to phylloquinone and then to 3 o Reduction of PGA to sugar by formation of iron-sulfur clusters glyceraldehyde 3-phosphate (GAP) using 4. PQH2 is mobile and binds to cytochrome b6f analyzed products via chromatography which engages in the Q cycle 2. Pheo.

Redox Control C4 plants  Light dependent  CO2 + phosphoenolpyruvate (PEP) via  Controlled by thioredoxin which reduces phosphoenolpyruvate carboxylase disulfide bridges into sulfhydryl groups in  Tropical grasses that are adapted to hot and arid enzymes causing their activation environments that can close stomata to conserve  Some of the electrons go to thioredoxin instead water but can still maintain photosynthesis. opened at night  Rubisco only has modest preference for carbon  Malate is evident as sour morning taste dioxide vs oxygen making this very easy since neither of the two binds to the active site of the enzyme . of ferredoxin won’t work well in cooler temperatures or in  During the dark. thioredoxin is no longer reduced higher altitudes by ferredoxin and the enymes revert back to an  Continues net synthesis of carbohydrate until CO 2 oxidized state levels have dropped to 1-2 ppm  PEP carboxylase can operate at low levels of CO2 Photorespiration and is not inhibited by oxygen  Glycolate is a product of a reaction catalyzed by  Downside is it requires more ATP and NADPH Rubisco where Oxygen attaches to RuBP forming  Happens in different parts of a leaf 2-phosphoglycolate which is converted to o Mesophyll cells where fixation occurs glycolate o Bundle-sheath cells sealed off from  Glycolate is transferred to the peroxisome where atmospheric gass CO2 is released  Evolved separately from C3 ancestors  Upatake of oxygen and release of carbon dioxide  Waste of plant’s energy CAM plants  Occurs when the concentration of oxygen is high  Also utilize PEP carboxylase but carry out light- as compared to carbon dioxide (when plants dependent reactions in different times of the day close stomata to prevent water loss)  Keep stomata closed at day.