ENERGY METABOLISM: PHOTOSYNTHESIS

Living organisms are highly ordered, structured systems. y SUNLIGHT (effects): 1) Directly photosynthesis 2) Indirectly respiration Photoautotrophs gather energy from sunlight, using CO2 in building materials. Ex. Photosynthetic (all green) plants & (some) bacteria, algae, cyanobacteria Heterotrophs sources of energy are food (CH20, proteins & fats) through respiration, use food to build materials. Ex. Animals, fungi, most bacteria, parasitic plants, non-photosynthetic plants, protozoa Tissues and organs are either photoautotrophic/ heterotrophic. y PHOTOAUTOTROPHIC: Chlorophyllous leaves & stems; young seedlings (emerge in sunlight) HETEROTROPHIC: roots, wood & flowers, young seedlings (germinating) Energy enters biological world by photosynthesis. Sunlight is captured by pigments (of plants), using the energy in chemical reactions (energized pigments can enter in only 2 chemical reactions). WAYS OF MOVING ENERGY FROM ENERGIZED PIGMENTS INTO ENDERGONIC REACTIONS: 1.) Allow pigments to enter into every important reaction. Energized pigments They are large molecules, not very mobile, never move across membranes, hard to control, can react with almost anything. 2.) Allow pigments to make one/ several smaller, less energetic intermediates (that can be moved & controlled easily). ATP an essential, extraordinary molecule produced in photosynthetic reactions (form tiny fraction of plant) y Converted to ADP & phosphate by metabolic reactions. Guanosine triphosphate relative of ATP

Immature fruits green & photosynthetic. In last stages of maturation, chloroplasts became chromoplasts Chromoplasts carry other pigments than chlorophyll Photosynthesis complex process by w/c CO2 is converted to carbohydrate involving endergonic reactions driven by ATP (light to chemical energy) Chloroplast most impt. (in photosynthetic prokaryote) CO2 + H2O (CH2O)n + O2

METHODS BY w/c ADP can be PHOSPHORYLATED TO ATP 1.) Phosphorylation involves light energy in photosynthesis. (chloroplasts) 2.) Substrate-level phosphorylation compounds w/ high-energy phosphate groups are produced, forcing their phosphate onto ADP ATP. It doesnt involve oxygen (site is cytosol) 3.) Oxidative phosphorylation oxidations with oxygen (site mitochondria) Oxidized an atom doesnt carry as many electrons Reduction reaction reduces positive charge Oxidation reaction increases positive charge Oxidized compounds often carry great deal of O Reduced compounds contain Hydrogen Oxygen pull electrons away from an atom, increasing atoms partial + charge. Hydrogen stable when giving up electrons, reducing + charge.

NADPH nicotinamide adenine dinucleotide phosphate ATP & NADPH formed in highly endergonic reactions driven by light energy.

Redox reaction full reaction; oxidationreduction reaction Reducing power Its the ability to force electrons onto compounds. NAD+ (nicotinamide adenine dinucleotide) & NADP+ (nicotinamide adenine dinucleotide phosphate) are used mostly, picking up e and + p , reduced to NADH & NADPH. Oxidizing agents: NAD+ & NADP+ (take away e from molecules) Reducing agents: NADH & NADPH (place eonto molecules, reducing them and become oxidized themselves. Redox potential Its the tendency to accept/ donate e- varying greatly. ELECTRON CARRIERS 1.) Cytochromes small intrinsic membrane proteins containing cofactor Heme (has iron atom). Like cytochromes, 2.) Plastoquinones transport e- over short distances w/in a membrane, picking up 2 e , + bind 2 p . Their hydrocarbon tail makes them hydrophobic. 3.) Plastocyanins carry copper atom (+2 when oxidized, +1 when reduced). They can move a short distance along surface. H2O & CO2 are excellent for they diffuse into plants automatically. (Both are very stable and have little chemical energy). CH2O It stores energy, stable, chemically unreactive. 6CO2 + 6H2O + energy C6H12O6 + 6O2

Stroma reactions (dark reactions) endergonic ATP + NADPH + CO2 CH2O

Electromagnetic radiation spectrum include gamma, cosmic rays, UV (short WL), X-rays; infrared light, microwaves, radar, radio waves (long WL) & visible light Quantum/ photons set of particles Radiation can be thought of & treated either as quanta (long) or set of waves (short, < 1nm). Short wavelengths large amounts of energy, more heat in each quantum Long wavelengths little energy amounts, cant boost electrons energy, make molecule warmer, not useful in chemical synthesis Quantum energy wavelengths, distinguished in colors. y Animals see 350 to 760 nm. y Red is 760 nm, violet is 390 nm. Pigment any material absorbing certain WL specifically & has distinctive color. It should absorb high-energy radiation instead of fairly weak visible light.

Photosynthetic pigments transfer absorbed light energy to e entering chemical reactions. Its black when its theoretically ideal, absorbing & using ALL LIGHT. Chlorophyll a absorbs some red & blue light. Its tail has H and methyl groups, thus, hydrophobic. Its essential in all plants. Chlorophyll doesnt use high-energy quanta for it has too much energy. Its green for most green light passes through it. Conjugated double bonds all of its bonds are DOUBLE bonds alternating with SINGLE bonds; excellent for absorbing quanta. Visible light has right amount of energy. When 1 of quanta is absorbed by pigment, an e- is activated. If quantum has wrong WL, it passes through pigments bonding orbitals w/o being absorbed. If correct WL, its absorbed.

Carbon in CO2 is +4, while in CH2O is +0. Products of photosynthesis are non-toxic. Carbon is more stable in oxidized state. CH2O can release energy, but CO2 cant. Water electron source; light energy source Light-dependent reactions H2O & light create intermediates ATP & NADPH, result in a Chemiosmotic gradient.

Electron & molecule go from ground state excited state (can be used in chem. reactions). Fluorescence release of light by a pigment Absorption spectrum Its a graph showing which WLs are most strongly absorbed. Action spectrum It shows which WLs are most effective at powering photosynthesis. Accessory pigments molecules that absorb WLs strongly that chlorophyll a doesnt. They overcome narrow absorption of a & broaden action spectrum. Chlorophyll a & b large, flat with almost same porphyrin ring structures and hydrophobic phytol tails. Theyre different in terms of substituent Carbon #17. Reaction centers special a molecules Resonance Its the transfer of absorbed energy by chlorophyll to another in a different part of complex. Allows chlorophyll b absorb WLs that chlorophyll a misses & transfer energy to a for chemical reactions.

PHOTOSYSTEM I reaction 1.) When light strikes any pigment of Photosystem I, energy is moved to reaction center & absorbed by a pair of P700 molecules (absorb red light of 700 nm well). 2.) Energy excites an e- of P700, then absorbed by Fx (no orbital formed). *Fx = Fe4S4. 3.) The transferred e- is unstable, & reduced Fx passes it onto ferredoxin (protein with 1 electron & a site having 2 iron atoms bound to 2 sulfur atoms). 4.) E are passed to ferredoxin-NADP reductase (enzyme with 2 electrons) 5.) The enzyme transfers 2 e to NADP , reducing it to NADPH + H+ 2 PHASES/ CYCLES OF PHOTOSYNTHESIS 1.) Light reaction involves membrane where pigments are found. (Hills reaction) H2O (energy carried given to ADP) oxygen
+ +

2.) Dark reaction involves stroma where carriers/ coenzymes are found. (biochemical) CO2 + solar energy sugar

Carotenoids Theyre important in absorbing excess light, protecting chlorophylls from excessive sunlight. - Xanthophylls have oxygen - Carotenes lack oxygen When light excites an e- in chlorophyll a, its unstable, wasting energy. Thylakoid membranes They are sites where chlorophyll molecules & e- carriers (in photosynthesis) must be bound. It has ferredoxin. Photosynthetic unit granule resulted as a pack of all pigments and carriers working together. (It has 300 molecules of a & b) Photosystem I more a, less b (P700) Photosystem II more b, less a or b = a; reduces P700 (P680), working backward from P700.

PARTS OF A PIGMENT 1.) Head made up of porphyrin (in center has +2 Mg (absorbs e ), when it gets e reduced form) y Porphyrin on top of membrane like soldiers, with 4 phytol rings (chlorophyll). y NADH oxidized form of NAD 2.) Tail PHOTOSYSTEM II Reaction Chlorophyll a (absorb light) Cytochrome b6/f complex Plastocyanin Phaeophytin (no Mg) Plastoquinone Q

chlorophyll a (P680) P700 NADP+

P680 gets new e from H2O, not from Plastocyanin. H2O breaks down into H+ (plant uses) & O2 (discarded through stomata) PHOTOSYSTEM I & II 1.) E- are passed H2O P680 2.) Energys boosted by light, moving through electron transport chain P700 3.) Energys boosted by light again nd 4.) Energy passes through a short 2 ETC + reduced NADP , NADPH. 5.) Protons + NADP+ ETC between P680 & P700 = ATP y Plastoquinone moves p+ from stroma to thylakoid lumen = increased p+ concentration in TL & decreased one in stroma.

Cyclic Electron Transport chloroplasts make extra ATP without making NADPH ATP : NADPH (proper ratios) - Flow of e- from P700 back to plastoquinone, such that that theres p+ pumping but no NADPH synthesis. Calvin/ Benson cycle C3 cycle; CO2 CH2O (In stroma reactions) *product is PGAL 1.) RuBP (ribulose-1, 5-biphosphate) accepts CO2 forming 2 similar molecules with 3 carbons: 3-phosphoglycerate (PGA). 2.) RuBP carboxylase (RUBISCO) carries out the reaction. 480,000 Daltons, most abundant protein on Earth; low affinity on CO2 3.) The enzyme puts O rather than CO2 onto RuBP (amino acid sequence is identical). st Carboxylation 1 step of stroma reactions donates energy to PGA ATP 1, 3-diphosphoglycerate NADPH reduces it 3-phosphoglyceraldehyde (PGAL amazing versatile molecule; basis of animal metabolism) RuBP (after stroma reactions) Plants needs large amount of RuBP to intake CO2. As PGAL is formed reconverted RuBP by stroma reactions, its exported to cytoplasm. Anabolism consists of anabolic reactions, construction from simple to complex molecules in cytoplasm. 2 IMPORTANT PATHWAYS 1.) Synthetic pathways of polysaccharides 2.) Fats (storage forms of energy & carbon) ATP & NADPH excellent sources of energy, cant be stored for a short time. (Reactive & unstable break down) Short-term storage: ATP & NADPH Intermediate-term: simple sugar glucose & disaccharide sucrose Long-term storage: starch (polymer of glucose, too large to be moved)

Z scheme shape/ other name for this reaction NADPH produced in light-dependent reactions. Chemiosmotic phosphorylation synthesis of ATP from ADP & phosphate using energy of an osmotic gradient & a gradient of electric charge (occurs in chloroplasts & mitochondria). Frets thylakoids lying between grana Thylakoid lumen Its an important compartment where enzymes & e- carriers are embedded (facing lumen or stroma). Lumen side/ Granum areas Its where reactions break down H2O & produce O & protons Stroma side The site where ferredoxin-NADP reductase generate NADPH. y As protons are absorbed = decrease in their stroma concentration. ATP synthetase entire complex creating ATP from ADP & phosphate CF0-CF1 complex ATP synthetase of chloroplasts + y CF0 where actual p channel is. y CF1 phosphorylates ADP ATP + y P flow thru ATP synthetase powers phosphorylation of ADP to ATP Noncyclic electron transport when e- flows from H2O NADPH (release of O), without this, theres no H2O breakdown, O making.

Gluconeogenesis Its the anabolic synthesis of y glucose in chloroplasts, amyloplasts or cytosol. Dihydroxyacetone phosphate product of conversion of PGAL exported to cytoplasm Fructose-1, 6-biphosphate sugar formed when 1 molecule of DHP condenses with 1 molecule of unconverted PGA. (Versatile entering many metabolic pathways) Fructose-6-phosphate when FBP loses a phosphate (Versatile entering many metabolic pathways) Glucose-6-phosphate part of F6P is rearranged. In plants, its polymerized into polysaccharides (amylase, amylopectin or cellulose).

If plants of hot weather, o they keep stomata closed = starve, o leaf cells packed, H2O loss is reduced, o hard to dissolve CO2 from air to cytoplasm = slow photosynthesis Cylindrical leaves minimize H2O loss while maintaining photosynthesis reduce external surface. Photosynthesis reduced = CO2 absorption slowed Open stomata permit CO2, loss of H2O (day) Closed stomata CO2 cant be used at night, H2O retained DAY: CO2 cant enter if soils dry & H2O isnt free C4 METABOLISM (evolution of compartmentalization of RuBP carboxylase in CO2 high concentration & oxygen low concentration Theres a low ratio on: Each molecule CO2 absorption = H2O loss CO2 diffuses faster air concentration is HIGHER or leaf protoplasm concentration is LOWER CO2 concentration drops = enzyme-substrate binding slows

ENVIRONMENTAL & INTERNAL FACTORS 1.) Quality of sunlight colors/ WLs it contains. Sunlight is pure white (entire visible spectrum). y Red-enriched light formed at sunset/ sunrise, little effect on photosynthesis y Blue-enriched light formed at noon, allow efficient photosynthesis. 2.) Quantity of light light intensity/ brightness; greatest at noon in midsummer y More light = more CO2 y Under normal levels of CO2, light is limiting. y Photosynthesis is faster on brighter days. y Light compensation point level of illumination at which photosynthetic fixation of CO2 matches respiratory loss. y Cutin absorb short WLs y Understory plants are adapted to low light. y Lamina face exposed fully in mornings & afternoons 3.) Duration - # of hours per day sunlights available. y Starch sugar at night LEAF STRUCTURE: standard (palisade parenchyma above & spongy mesophyll below) ideal for absorbing CO2, scarce in conserving H2O

RuBP carboxylase binds O instead of CO2, acting as an oxygenase & producing 1 PGA & 1 of phosphoglycolate From chloroplast to peroxisomes & mitochondria
converted

Amino acids (serine & glycine) Photorespiration Its the breakdown, energywasting process, extremely exergonic, not used in providing power to endergonic reactions. Protects plant whenever RuBP carboxylase (no mutation) picks up oxygen (since phosphoglycolate can be toxic)

C4 CO2 absorption, transportation & concentration in leaf, oxygen is kept way from RuBP carboxylase (w/ Kranz anatomy) PEP (phosphoenolpyruvate) carboxylase in mesophyll cells, high affinity & specifity for CO2. Never picks up oxygen H2O loss: CO2 low ratio

Adds CO2 to PEP = oxaloacetic acid (4 Cs)

reduced by NADPH

SUMMARY Anabolism from raw materials to compound (polysaccharide photosynthesis) Catabolism large molecule CO2 & H2O (respiration breakdown of compounds) Chemoautotroph chemicals as source Prokaryote ingested by eukaryote RESPIRATION (backward reaction) (CH2O)n + O2 CO2 + H2O

Malic acid (malate)

release CO2, moves to bundle sheath

Pyruvate forming NADPH Result: CO2 & reducing power transfer (by malate) Bundle sheath chloroplasts carry cyclic e+ transport, pump p , making ATP, where RuBP carboxylase is located BUNDLE SHEATH: Malate Pyruvate (CO2 release) Light is needed to make ATP to convert: Pyruvate PEP CAM (Crassulacean Acid Metabolism) For H2O conservation during photosynthesis 1st discovered in Crassulaceae Plant cant store ATP & NADPH during day for night use. Stomata close at dawn. Not particularly efficient, small amount of CO2

PHOTOCHEMICAL REACTION 1.) Photoexcitation of chlorophyll 2.) Photolysis of H2O 3.) Photophosphorylation (ATP production) + 4.) Photoreduction of NADP Chlorophyll a blue-green Chlorophyll b yellowish-green Emersons effect movement of quantum/ energy from 1 molecule to another

NOTES: y C3 plants cant survive in dry climate. y C4 met. an adjunct/ addition to C3 y CAM plants are selectively advantageous in dry climate. They grow & reproduce more than C3 & C4. y CAM plants arent selectively advantageous under moist conditions. y Bph bacteriopheophythin y Bacteriochlorophylls - pigments of purple & green bacteria C3
Ultimate carb. Adjunct Metab. Adjunct carb. Photorespiration Stomata open
RuBP carb.

C4 Same CO2 transfer PEP carb. Low Day

CAM Same CO2 storage PEP carb. Mod. Night

High Day