Chapter 8:
Photosynthesis
Photosynthesis…
■ …theconversion of light to
chemical energy
auto – trophic = “self feed”
■ ultimate source of organic
compounds
■ PRODUCERS
plants
multicellular alga
unicellular protist
cyanobacteria
hetero – troph = “other feed”
■ live on compounds produced by
other organisms
■ CONSUMERS
■ subtle forms: decomposers, etc.
CHLOROPLASTS
■ all plant parts have chloroplasts,
but they are most abundant in
leaves
■ color comes from chlorophyll (or
another pigment)
■ light energy absorbed drives the
synthesis of food molecules
CHLOROPLASTS
■ found mainly in cells of
mesophyll
(tissue in interior of leaf)
■ microscopic pores (stomata)
control flow of gases
■ water is absorbed by roots
CHLOROPLASTS
■ STRUCTURE
■ double membrane
■ FLUID = stroma
■ thylakoid = “coins”
■ grana = “stacks”
CHLOROPLASTS
PHOTOSYNTHESIS
EQUATION
6CO2 + 12H2O + light energy →
C6H12O6 + 6O2 + 6H2O
■ net equation would only show water as a
reactant
■ net equation is reverse of respiration
(both processes occur in plants)
PHOTOSYNTHESIS
EQUATION
CO2 + H2O + light energy →
CH2O + O2 (simplified form)
■ synthesis of carbohydrate one carbon at
a time
PHOTOSYNTHESIS
EQUATION
■ MECHANISM: O2 released from H2O
(not CO2)
Old View:
Step 1: CO2 → C + O2
Step 2: C + H2O → CH2O
PHOTOSYNTHESIS
EQUATION
■ Van Niel: O2 released from H2O
CO2 + H2S → CH2O + H2O + 2S
PHOTOSYNTHESIS
EQUATION
■ All photosynthetic organisms require a
hydrogen source.
General:
CO2 + 2H2X → CH2O + H2O + 2X
Sulfur bacteria:
2H2X → 2S
Plants:
2H2O → O2
PHOTOSYNTHESIS
EQUATION
■ The splitting of water was confirmed
using a radioactive source.
(1.)
CO2 + 2H2O → CH2O + H2O + O2
(2.)
CO2 + 2H2O → CH2O + H2O + O2
PHOTOSYNTHESIS
EQUATION
■ The splitting of water was confirmed
using a radioactive source.
(1.)
Water is the source of H in products.
(2.)
Water is responsible for the release of O2
PHOTOSYNTHESIS
EQUATION
■ Photosynthesis is also a redox reaction,
but e- flow is in the opposite direction.
■ electrons increase PE as they move
from H2O to sugar
■ energy boost is provided by light
2 STAGES OF
PHOTOSYNTHESIS
■ PURPOSE: convert solar energy to chemical
energy
■ light reactions = thylakoid membrane
■ Calvin cycle = stroma
--incorporates CO2 from air into organic
compounds (CARBON FIXATION)
--fixed C reduced to CH2O
(reducing power provided by NADPH)
LIGHT REACTIONS
■ OVERVIEW:
■ light absorbed by chlorophyll drives transfer of
e- from H2O to NADP+ (e-
acceptor…temporarily stores energized e-)
■ water is split, resulting in O2 as a product
■ ATP is also generated in
photophosphorylation
LIGHT REACTIONS
■ SUNLIGHT:
■ light = electromagnetic radiation; travels
in waves (disturbances in electric and
magnetic fields)
■ visible light is most important portion of
EM spectrum to living things
electromagnetic spectrum
LIGHT REACTIONS
■ SUNLIGHT:
■ light also behaves as particles, called
photons
■ amount of energy is inversely related to
wavelength
■ light of visible spectrum drives
photosynthesis
location and structure of chlorophyll
LIGHT REACTIONS
■ PIGMENTS:
■ absorb visible light
■ black absorbs all wavelengths
■ as light meets matter, it may be
reflected, transmitted, or absorbed
excitation of chlorophyll
LIGHT REACTIONS
■ PIGMENTS:
■ spectrophotometer can measure
ability of pigment to absorb various
wavelengths
■ measure absorption vs. wavelength
(absorption = fraction of light not
transmitted or reflected)
LIGHT REACTIONS
■ PIGMENTS:
■ absorption spectrum underestimates
effectiveness of certain wavelengths
■ only cholorphyll a can participate in
light reactions, but other pigments can
absorb light and transfer the energy to
chlorophyll a
LIGHT REACTIONS
■ ACCESSORY PIGMENTS:
■ chlorophyll b—slight structural difference
leads to different absorption spectrum
■ carotenoids—function of some seems to be
photoprotection; instead of transmitting light
to chlorphyll a, they absorb and dissipate
excessive light that would otherwise damage
chlorophyll a
absorption spectra
LIGHT REACTIONS
■ EXCITATION:
■ after absorbing a photon of light, one of
the molecules’ electrons are elevated to
an orbital where it has more PE
■ photons are absorbed by clusters of
pigment molecules
PHOTOSYSTEMS
■ consist of light-gathering complex of
chlorophyll a, chlorophyll b, carotenoids
■ enable harvesting of light over wide
surface area and wide spectrum of light
■ energy is transmitted between
chlorophyll a molecules, then eventually
to reaction center ( a specific chlorophyll
a molecule)
how a photosystem harvests light
PHOTOSYSTEMS
■ Photosystem I (P700)
■ Photosystem II (P680)
■ these are actually identical chlorophyll a
molecules, but have slightly different
proteins
PHOTOSYSTEMS
■ Noncyclic electron flow
■ light drives the synthesis of NADPH and
ATP by energizing the photosystems
noncyclic electron flow
PHOTOSYSTEMS
■ Cyclic electron flow
■ uses Photosystem I, but not photosystem II
(short circuit)
■ electrons cycle back to Fd from primary e-
acceptor
--no production of NADPH
--no release of O2
-- ATP is generated = cyclical photo-
phosphorylation
cyclic electron flow
PHOTOSYSTEMS
■ Cyclic electron flow
■ What is the purpose?
■ Calvin cycle uses more ATP than
NADPH
■ cyclic photophosphorylation makes up
the difference
PHOTOSYSTEMS
■ Cyclic electron flow
■ Concentration of NADPH may regulate flow
■ if ATP runs low, NADPH will accumulate
because photosynthesis slows down
■ a rise in NADPH may lead to a shift to cyclic
electron flow
CHEMIOSMOSIS
■ Mitochondria: food (CE) → ATP
H+ driven from innermembrane space to
matrix
(OUT → IN)
■ Chloroplast: light → chemical energy
H+ driven from thylakoid space to stroma
(IN → OUT)
--ATP forms in stroma
--pH in thylakoid space rapidly drops when
illuminated
chemiosmosis:
mitochondria vs. chloroplasts
light reactions and chemiosmosis
Melvin Calvin
CALVIN CYCLE
■ similar to Krebs cycle in that starting
material is regenerated
■ CARBON: enters as CO2, leaves as
carbohydrate (G3P)
--requires 3 cycles
CALVIN CYCLE
■ PHASE 1= Carbon fixation
■ ribulose bisphosphate catalyzes
reaction
■ rubisco is most abundant protein in
chloroplasts (likely the most abundant
protein on Earth)
CALVIN CYCLE
■ PHASE 1= Carbon fixation
■ CO2 + RuBP → intermediate
(1 C) (5 C) (6 C)
■ (6 C) → 2 3-phosphoglycerate (PGA)
(3 C) (3 C)
CALVIN CYCLE
■ PHASE 2= Reduction
■ 3-phosphoglycerate (PGA) + P →
1,3-bisphosphoglycerate (4-C)
CALVIN CYCLE
■ PHASE 2= Reduction
■ 1,3-bisphosphoglycerate + 2e- →
G3P (same sugar is formed when
glucose is split)
CALVIN CYCLE
■ PHASE 3= Regeneration of CO2
acceptor (RuBP)
CALVIN CYCLE
METABOLIC ADAPTATIONS
FOR PLANT SURVIVAL
■ compromise between photosynthesis and
excessive water loss from the plant
■ on a hot, dry day, most plants will close
stomata to conserve water
– even with stomata partially closed, [CO2]
decreases in air spaces
– [O2] released from photosynthesis begins to
increase
– under these conditions, photorespiration is favored
C3 PLANTS
■ most plants use the Calvin cycle to fix C
into 3-C compounds
– 3-C compound= 3-phosphoglycerate
– rubisco is enzyme
■ Ex. rice, soy, wheat, bean
■ produce less food when stomata close
on hot dry days
PHOTORESPIRATION
■ photo = occurs in light
■ respiration = consumes O2
■ rubisco can accept O2 instead of CO2
■ photorespiration decreases photosynthetic
output by siphoning organic materials from
the Calvin cycle
– generates no ATP
– produces no food
■ not known how process is beneficial (drains
up to 50% of C from Calvin cycle in soy
beans)
PHOTORESPIRATION
■ conditions that lead to photorespiration:
– bright, hot, dry days
– leads plants to close stomata
■ C4 and CAM pathways minimize water
loss and photorespiration
C4 PLANTS
■ preface Calvin cycle by forming 4-C
compounds
■ Ex. corn, sugar cane
■ have unique anatomy:
– bundle-sheath cells: arranged in tightly
packed sheaths around leaf veins
• site of Calvin cycle in chloroplasts
– mesophyll cells: more loosely arranged
around bundle-sheath cells
• CO2 incorporated into organic compounds
C4 PLANTS
C4 PLANTS
[in mesophyll cells]
1. CO2 + PEP (3-C) → oxaloacetate (4-C)
enzyme: PEP carboxylase
(has high affinity for CO2 compared to rubisco)
(can fix CO2 efficiently when rubisco can’t)
2. oxaloacetate (4-C) → malate (4-C)
[in bundle-sheath cells]
3. malate (4-C) → pyruvate (3-C) → PEP
CAM PLANTS
■ Ex. cactus, pineapple
■ open stomata at night, close during the
day
■ mesophyll cells use vacuoles to store
organic acids made during the night
■ stomata close in the morning
■ during the day, CO2 is released from
organic acids, allowing ATP and NADPH
to run the Calvin cycle
C4 and CAM plants
photosynthesis overview