Chapter 10 - stomata  microscopic pores where CO2 enters and O exits

Photosynthesis - stroma
 dense fluid inside the chloroplast
PHOTOSYNTHESIS  enclosed by 2 membranes
- the conversion of light energy to chemical energy stored in - thylakoid membrane
sugar and other organic molecules  separates stroma from thylakoid space/lumen
 where chlorophyll resides
- grana  stacks of thylakoid sacs
PHOTOSYNTHESIS IN NATURE

THE PATHWAYS OF PHOTOSYNTHESIS

2 Types of Nutrition
1. Autotrophic
THE SPLITTING OF WATER
2. Heterotrophic
- oxygen given off by plants was derived from water and not
from CO2
AUTOTROPHS
- most impt result of shuffling of atoms during photosynthesis 
- self-feeders in the sense that they sustain themselves extraction of H from H2O and its incorporation into sugar
without eating other organisms or substances derived from - O2
other organisms  waste product of photosynthesis
- produce their organic molecules from CO2 and other  restores the atmospheric oxygen consumed during cellular
inorganic raw materials obtained from env respiration
- ultimate sources of organic cpds for all nonautotrophic org.
- producers
PHOTOSYNTHESIS AS A REDOX PROCESS

Photoautotrophs - during respiration, energy is released from sugar when

electrons associated with H are transported by carriers to
- Plants oxygen forming water as by-product
 uses light as source of energy for to synthesize - reverses the direction of e- flow
organic substances - water is split  e- are transferred along with H ions from the
 raw materials: CO2 from air, water and minerals from water to CO2  reducing it to sugar
soil
- algae
STAGES OF PHOTOSYNTHESIS
- prokaryotes
1. light reactions  “photo” part
2. Calvin cycle/dark reactions  “synthesis” part
Chemoautotrophs
- a rarer form of self-feeding
OVERVIEW OF PHOTOSYNTHESIS
- energy is obtained by oxidizing inorganic substances such
as ammonia and sulfur
LIGHT REACTIONS
HETEROTROPHS
- Convert solar energy to chemical energy
- live on compounds produced by other organisms - Light absorbed by chlorophyll drives a transfer of e- and
- consumers hydrogen from water to NADP+
- other-feeding  animal eats plants or other animals  an acceptor
- almost all heterotrophs are completely dependent on  nicotinamide adenine dinucleotide phosphate
photoautotrophs for food and oxygen (by-product of  temporarily stores the energized e-
photosynthesis) - water is split in the process
- by-product  O2
Decomposers - use solar power to reduce NADP+ to NADPH by adding 2 e-
and H+
- feed on carcasses, feces, and fallen leaves - generate ATP by powering the addition of a phosphate group
- fungi and some bacteria to ADP
- photophosphorylation  process by which ATP is generated
- light energy is converted to chemical energy in the form of the
CHLOROPLASTS
2 compounds
- sites of photosynthesis in plants  NADPH  a source of energized e-/reducing power
- all green parts of a plant have chloroplasts  ATP  energy currency of cells
- contains chlorophyll - NOTE: doesn‟t produce sugar!!!
 green pigment
 gives leaf its green color
CALVIN CYCLE
- found mainly in the cells of the mesophyll tissue in the
interior of the leaf - Carbon fixation  Incorporating air CO2 into organic
molecules already present in chloroplast
- Reduces fixed carbon to carbohydrate by addition of e-
 Reducing power  NADPH from light rxns
 Needs ATP
- Makes sugar but only does so with the help of NADPH
and ATP from light rxns
- Also referred to as dark reactions because light is not
directly needed
- Usually occurs in daylight for most plants
CHLOROPLAST
- Uses light energy to make sugar by coordinating 2 stages
of photosynthesis
- Light reactions  occur in thylakoids
- Calvin  stroma
- As NADP+ and ADP molecules bump into the thylakoid
membrane, they pick up e- and phosphate and transfer
their high-energy cargo to Calvin cycle
LIGHT REACTIONS
Review: Waves and Light (see book)
- Photon  light behaves as though it consists of discrete
particles
- Visible light  radiation that drives photosynthesis
PHOTOSYNTHETIC PIGMENTS
- Substances that absorb visible light
- Diff pigments absorb light of different wavelengths
- Wavelengths that are absorbed disappear
- Chlorophyll  green because it absorbs red and blue
while transmitting and reflecting green light
- Light can only perform work in chloroplasts only if it is
absorbed
- Blue and red light works best for photosynthesis
- Green  least effective color
Chlorophyll a
- Not the only photosynthetically impt pigment
chloroplasts
- Only one that can participate directly in light reactions
- Converts solar to chemical energy
- Blue-green
- Ground state: e- is in its normal orbital
- a molecule absorbs a photon
- one of molecule‟s e- is elevated to an orbital with greater PE 
higher energy
- pigment molecule is said to be in an excited state
- photons are absorbed by clusters of pigment molecules
embedded in thylakoid membrane
- energy of absorbed photon is converted to PE of an e- raised
from ground to excited state
- fluorescence
 e- jumps to an excited state and back to ground state 
photon is given off  the afterglow is fluorescence
 if chlorophyll is isolated, this is what happens  no e-
acceptor
PHOTOSYSTEMS
- light-harvesting complexes of the thylakoid membrane
- chlorophyll, proteins, other kinds of organic molecules
- antenna complex
 gathers light
 consists of cluster of chlorophyll a, b, and carotenoid
molecules
- no. and variety of pigments enable a photosystem to harvest
light over a larger surface and larger portion of the spectrum
- antenna absorbs a photon
- energy is transmitted from pigment to pigment until it reaches
chlorophyll a
- chlorophyll a is located in the reaction center
 contains chlorophyll and primary e- acceptor
 where the first light-driven chemical reaction of
photosynthesis occurs
- light excites the e-  chlorophyll a loses 1 e-  primary e-
acceptor traps the high energy e-
Photosystem I
- has a characteristic rxn center  chlor. a associated with
specific proteins
- chlorophyll a  P700  pigment is best at absorbing light with
wavelength of 700 nm
in Photosystem II
- has a characteristic rxn center  chlor. a associated with
specific proteins
- P680

Chlorophyll b 2 Possible Routes of Electron Flow

- Another form of chlorophyll 1. noncyclic
- Almost structurally similar to chlorophyll a 2. cyclic
- Yellow-green
- Absorbs sunlight and transfers it to chlorophyll a
NONCYCLIC ELECTRON FLOW
- Predominant route
Carotenoids
- Hydrocarbons that are various shades of yellow and
Steps
orange
- May broaden the spectrum of colors that drive 1. photosystem II absorbs light
photosynthesis  e- is excited
- Photoprotection  absorb and dissipate excessive light  e- is captured by primary e- acceptor
energy that would otherwise damage chlorophyll 2. enzyme extracts e- from H2O
 supplies the e- to P680  replaces e- lost
EXCITATION OF CHLOROPHYLL BY LIGHT  rxn splits a H2O molecule into 2 H+ and 1 O atom
  releases O2
3. each photoexcited e- passes from primary e- acceptor of
PII to PI via electron transport chain
 e- carrier  plastoquinone (Pq)
 complex of 2 cytochromes  iron-containing protein
 plastocyanin (Pc)  copper-containing protein
4. e- passes through chain  fall to lower energy level
 harnessed by thylakoid membrane
 ATP is produced  will eventually provide chem
energy for production of sugar in Calvin cycle
 photophosphorylation  ATP synthesis
 since noncyclic  noncyclic photophosphorylation
 chemiosmosis  mechanism for photophos…
5. e- reaches bottom of ETC
 fills e- hole in PI
 hole resulted from excitation of e-  accepted by PI‟s
primary e- acceptor
6. PI‟s primary e- acceptor passes the excited e-s to another
ETC
 transmits ferredoxin (Fd)  iron containing protein
 NADP+ reductase  enzyme that transfers the e-„s
from Fd to NADP+
 This is the redox rxn that stores high-energy e- in
NADPH  provides reducing power for synthesis of
sugar in Calvin cycle
CYCLIC ELECTRON FLOW
- Only uses PI
- No production of NADPH and no release of oxygen
- Cyclic photophosphorylation  Generation of ATP
- Fcn  to replenish lacking ATP
 Noncyclic EF produces ~ = amounts of ATP and
NADPH
 Calvin cycle consumes more ATP than NADPH
- Cyclic vs noncyclic  regulated by concentration of
NADPH
Steps
1. excitation of e-  e- is trapped by PI‟s primary e- acceptor
2. e- passes through Fd, cytochrome complex, and Pc
3. e- goes back to P700 chlorophyll a
CHEMIOSMOSIS: MITOCHONDRIA VS. CHLOROPLAST
- both generate ATP by this mechanism
- ATP synthase complexes of both are alike
- Mitochondria transfer chemical energy from food
molecules to ATP
- Chloroplasts transform light energy into chemical energy
- ATP is formed in stroma
- NADPH is produced on the side of the membrane facing
the stroma
Summary of Light Reactions
- Noncyclic electron flow pushes e- from water (low state of
PE) to NADPH (stored at high PE)
- Generates ATP
- Equipment of thylakoid membrane converts light energy
into chemical energy stored in NADPH and ATP
- Oxygen is a by-product
CALVIN CYCLE
- Starting material is regenerated after molecules enter and
leave the cycle
- Carbon enters in form of CO2 and leaves in form of sugar
- Spends ATP as energy source and NADPH as reducing power
- Product: glyceraldehyde-3-phosphate (G3P)
 3 carbon sugar
 carbohydrate
 for the net synthesis of 1 molecule of G3P, cycle must
take place 3x, fixing 3 molecules of CO2, and spending 9
ATP & 6 NADPH molecules
3 PHASES OF THE CALVIN CYCLE
1. Carbon Fixation
 CO2 is incorporated by attaching 1 CO2 to ribulose
biphosphate
 Ribulose biphosphate  RuBP  5-C sugar
 Rubisco/RuBP carboxylase catalyzes this step  most
abundant protein in chloroplasts and probably on earth
 Product  6-C intermediate that splits into 2 molecules of
3-phosphoglycerate (for each CO2)
2. Reduction
 each molecule of 3-phosphoglycerate becomes 1,3-
biphosphoglycerate  by receiving a phosphate grp
from ATP
 2 e- from NADPH reduces 1,3-biphosphoglycerate to G3P
NOTE:
 G3P is a sugar that stores more PE
 for every 3 molecules of CO2 there are 6 molecules of G3P
 Net gain G3P = 1  because 1 G3P exits cycle to be used while
other 5 must be recycled to regenerate the 3 RuBP molecules
3. Regeneration of CO2 acceptor (RuBP)
 carbon skeletons of 5 G3P molecules are rearranged into
3 RuBP molecules
 for this step, the cycle spends 3 ATP molecules
- G3P becomes the starting material for metabolic pathways that
synthesize other organic compounds, including glucose and
other carbohydrates
ALTERNATIVE MECHANISMS OF CARBON FIXATION
- Developed in hot, arid climates
Compromise between photosynthesis and excessive H2O loss
- CO2 for photosynthesis enters via stomata
- Stomata are also main venues for transpiration  evaporative
loss of water from leaves
- Hot dry day  close stomata
 conserve water
 reduces photosynthetic yield by limiting access to CO2
PHOTORESPIRATION
C3 plants
- for most plants, the initial fixation of CO2 occurs via rubisco
- first organic product of carbon fixation is a 3 carbon cpd  3-
phosphoglycerate
- ex. Rice, wheat, soy - during the day, light reactions can supply ATP and NADPH for
Calvin cycle
- CO2 is released from the organic acids made the night before
Steps
to become incorporated into sugar in the chloroplasts
- declining level of CO2 starves Calvin cycle - NOTE: the 2 steps occur at separate times
- rubisco can accept O2 instead of CO2
- product splits REVIEW
- 1 pc, 2-C cpd is exported from chloroplast
- mitochondria and peroxisomes break the 2-C cpd to CO2 - Sugar made in chloroplasts supplies the entire plant with
- called as such because it occurs in the light and chemical energy and carbon skeletons to synthesize all major
consumes O2 organic molecules of the cells
- generates no ATP - About 50% of organic material is consumed for cellular
- produces no food respiration in mitochondria
- decreases photosynthetic output by siphoning organic - Technically the green cells are the only autotrophic parts of the
material away from Calvin cycle plant
- fostered by hot, dry days - The rest of the plant depends on organic molecules exported
from the leaves via veins
- Carbo is transported out of the leaves as sucrose
C4 PHOTOSYNTHESIS
 dissacharide
- preface Calvin cycle with alternate mode of carbon fixation  provides raw material for cellular respiration and a
- first product  4-C compound multitude of anabolic pathways that synthesize proteins,
- ex. Sugarcane and corn lipids, and other products
- its mechanism is correlated with a unique leaf anatomy - cellulose
 glucose linked together
 polysaccharide
2 Types of Photosynthetic Cells
 main ingredient of cell walls
1. bundle-sheath cells  most abundant organic molecule in the plant and probably
 arranged into tightly-packed sheaths around the veins on the planet
of the leaf - extra sugar is stockpiled by synthesizing starch
 where Calvin cycle is confined  some are stored in chloroplasts
2. mesophyll cells  storage cells of roots, tubers, seeds and fruits
 between bundle sheath and leaf surface
 more loosely arranged
 where the CO2 is incorporated into organic cpds

Steps
- addition of CO2 to phosphoenolpyruvate (PEP)
- oxaloacetate  4-C cpd is formed
- PEP carboxylase  the enzyme fixes CO2 efficiently
when rubisco cannot because of its high affinity for CO2
- Mesophyll exports their 4-C products to bundle-sheath
cells through plasmodesmata
- 4-C cpds release CO2 in the bundle-sheath cells
- CO2 is reassimilated into organic material by rubisco and
Calvin cycle
- Mesophyll cells pump CO2 into the bundle-sheath,
keeping the CO2 concentration high enough for rubisco to
accept CO2 rather than O2
- C4 photosynthesis minimizes photorespiration and
enhances sugar production
- NOTE: initial steps of carbon fixation are separated
structurally from Calvin cycle

CAM PLANTS
- Evolved in water-storing plants
- Ex. Cacti and pineapples
- Open their stomata during night and close them during the
day
- Crassulacean acid metabolism
 plants take up CO2 during the night and incorporate it
into a variety of organic acids
- the mesophyll cells store organic acids they make during
the night in their vacuoles until morning