Uniqueness of Plants
- Harnessing radiant energy for food
production
Photosynthesis
- CO2 + 6H20 + (energy) → C6H12O6 + 6O2
Carbon dioxide + water + energy from
light produces glucose and oxygen
- 13 x 1023 cal/year reaches the Earth
o 30% reflected back to the space
as light
o 20% absorbed by atmosphere
o 50% absorbed by earth itself,
converted into heat
o <1% used by plant during
photosynthesis
Photosynthesis
- Photos - light
- Synthesis – put together
- Light energy is converted to chemical
energy
- Life on earth depends on this process
- Not only does it provide food, it also
supplies oxygen
Basic Concepts
- Occurs in chloroplast in the presence of
light, CO2 is assimilated to form
carbohydrate and oxygen
- Forms carbohydrates and oxygen
- Process that converts electromagnetic
energy in visible regions of the light
spectrum into chemical energy (NADPH
and ATP) utilized in enzymatic reactions
that convert CO2 into organic molecule
Overall reaction
- Oxidation-redox reaction
o Oxidation of H2O
 2 H2O → 4 e- + 4 protons +O2
o Reduction of CO2 to form organic
compounds
 Through the transfer of e- from
donor to acceptor molecule
Leaf Structure
- Raw materials of PS enter the cells of the
leaf
o Water and CO2
- Products of photosynthesis exit leaf
o Sugar and oxygen
Nature of Light
- Visible light – small portion of EM
spectrum
- Plant use light energy from visible light
range for photosynthesis
Three possibilities when light hits an object
- Absorbed by object
- Reflected off object
- Transmitted through object
Colors we see are actually light reflected from
an object
- Plants reflect green, but absorb other
wavelengths for use in PS
Red and blue wavelengths are important for
photosynthesis
- Captured and absorbed by chloroplasts
used to initiate photosynthetic reactions
Action spectrum
- spectrum of electromagnetic radiation
most effective for photosynthesis
Leaf
- Major organ for photosynthesis in
vascular plants
Photosynthetic pigments
- pigments absorb photons of light
- electrons get excited
 Chlorophylls
- must be present so that plants can
undergo photosynthesis
- main pigment for photosynthesis
- absorbs light primarily in the blue and red
regions of the visible spectrum
 o a - Light-independent stage does not occur
 primarily absorbs light in the blue in the dark
and red regions of the visible
Light-dependent reactions
spectrum
 initiates light-dependent reactions - In the first stage of photosynthesis, light
 bright green in color energy drives electrons out of
 has methyl group (CH3) photosystems
o b - The electrons may be used in a cyclic or
 accessory pigment that is yellow- non-cyclic pathway of ATP formation
green in color
 has carbonyl group (CHO) Capturing Energy for Photosynthesis
 carotenoids – pigments that are yellow and - Photons boost e- in pigments to higher
orange in color and expand the spectrum energy
of light that provides energy for the whole - Light-harvesting complexes absorb
photosynthetic process, functions as energy
antenna/accessory pigments and - Electrons are released from special pairs
photoprotection of chlorophyll a molecules in
o carotenes photosystems
o xanthophylls
 collectively, chlorophyll and accessory Photosystems I and II
pigments absorb most wavelengths of - Photosynthetic units, include chlorophyll,
visible light accessory pigments
 certain electrons in pigment molecules - Organized with pigment-binding proteins
absorb photons of light energy, boosting into antenna complexes
electrons to a higher energy level
 energy is captured and used for Reactions centers
photosynthesis
- Special pair of chlorophyll a molecules
Light-dependent release energized electrons to acceptors
o P700 – reaction center for PS 1
- light energy is transferred to ATP and o P680 – reaction center for PS II
NADPH (study what these stand for lolz)
- H2O molecules split, release O2, Photosystem I
important for replenishing electrons
- NADPH formation
needed for the process
- 700 nm
Light-independent
Photosystem II
- Utilizing energy in ATP and NADPH to
- O2 evolution
drive the synthesis of glucose
- 680 nm
- Energy in ATP and NADPH drives synthesis
of glucose and other carbohydrates Character PS I PS II
from CO2 and water Reaction P700 P600
center
Misconceptions Best 700 nm (far- 600 nm (red)
- Triose phosphate – net product of the absorbed red)
wavelength
carbon dioxide fixation process (Calvin
Primary e- Ao Pheophytin
cycle); used to form sucrose and/or
acceptor
starch
- Glucose is NOT the product of Calvin
cycle
 Thylakoid Stroma Appressed Energy flow in photosystem
location thylakoids grana
and non- regions - Energy flow in the light-dependent
apressed reaction is an example of how organisms
grana harvest energy from their environment
regions
Harvesting e- energy
Responsible NADPH O2 evolution
for formation - Light energy is converted into chemical
Electron flow Cyclic Non-cyclic energy
o Plants harvest radiant energy
Replacing lost e- - Entry of electrons from a photosystem
into the electron transport chain is the
- e- lost from PS II are replaced by first step in light-dependent reactions
photolysis of water molecules, which - ATP forms in the stroma
dissociate into hydrogen ions and o Electron energy is used to build
oxygen up a H+ gradient across the
o photolysis – process by which light membrane
energy breaks down a molecule o H+ flows through ATP synthase,
such as water which attaches phosphate group
to ADP
Photophosphorylation
Electron flow in noncyclic pathway
- light-driven reaction that attaches
phosphate group to a molecule - Electrons lost from a photosystem enter
an electron transfer chain in the
Cyclic photophosphorylation
thylakoid membrane
- e- cycle within photosystem I o Electron transfer chains –
- output: ATP organized arrays of enzymes,
- No NADPH coenzymes, and other proteins
that accept and donate
Non-cyclic photophosphorylation
electrons in a series
- e- move from water to PS II, to PS I, to - Non-cyclic electron transport
NADPH o Light-dependent reactions
- O2 is released  Form ATP and NADPH
- Forms ATP and NADPH - Outputs: ATP, O2, NADPH

A comparison of Noncyclic and Cyclic Electron flow in a cyclic pathway

Electron Transport
- When NADPH accumulates in the
Non-cyclic Cyclic stroma, the non-cyclic pathway stalls
Electron H2O None – - A cyclic pathway runs in the type I
source electrons photosystems to make ATP; electrons are
cycle through cycled back to photosystem I and
the system NADPH does not form
Oxygen Yes No - Output: ATP (produced from
released? (from H2O) chemiosmosis)
Terminal e- ATP (by ATP (by - No oxygen generated
acceptor chemiosmosis chemiosmosis
) )
Photosyste PS I (P700) PS I (P700
m required and PS II (680) only)
Key concepts in the making of ATP and NADPH
- Photosynthesis proceeds through 2
stages in the chloroplasts in plants and
protists
- First stage: sunlight energy is converted
to chemical energy to form ATP and
NADPH
- The coenzyme NADPH forms in a
pathway that also releases oxygen
Light-independent reactions: Sugar Factory
- Cyclic, light-independent
- “synthesis” part
- a.k.a. calvin cycle, “carbon-fixation
stages”
- enzyme-mediated reactions that build
sugars in the stroma of chloroplast
Calvin Cycle (C3 pathway)
- carbon fixation reactions
- 3 phases:
o CO2 uptake
o Carbon reduction
o RUBP regeneration
Carbon fixation
- Extraction carbon from inorganic sources
(atmosphere) and incorporating them
into an organic molecule
- Builds triose phosphate from CO2
- Uses bond energy formed in light-
dependent reactions (ATP, NADPH)
1. CO2 uptake phase
o Enzyme rubisco (ribulose
biphosphate
carboxylase/oxygenase)
combines CO2 with RuBP (ribulose
biphosphate), 5-C sugar
o Forms 3-carbon PGA
2. Carbon reduction
o Energy of ATP and NADPH
 Convert PGA molecule to G3P
(glyceraldehyde-3-
phosphate)
o For each 6 CO2 fixed
 12 G3P produced
 2 G3P leave to form 1 glucose
3. RuBP regeneration phase
o Remaining G3P molecules are
modified to regenerate RUBP
Photorespiration
- Biological function: to dissipate excess
ATP and reducing power under
conditions of high light intensity, thus
preventing damage to chloroplast
- Rubisco acts as an oxygenase, catalyzes
reaction of RUBP and O2 forming
phosphoglycolate (2-C compound) in
addition to PGA in the chloroplast
- Phosphoglycolate is rapidly converted
into glycolate
- A light-induced respiration occurring in
green tissues as a response to low
CO2/O2 ratio
- Involves cooperative interaction of
chloroplast, peroxisomes, and
mitochondria
o Chloroplast: RuBP is oxygenated
to form phosphoglycolate →
glycolate
o Peroxisome: glycolate is
oxidized/converted to glycine
o Mitochondria: glycine (2
molecules) is metabolized into
serine (releasing ammonia and
CO2)
- Occurs at high temperature and high
irradiance due to:
o Substrate specificity of rubisco
shifts in favour of O2 as
temperature increases;
o Leaf of CO2 concentration
becomes very low at high
irradiance largely due to rapid
photosynthesis; the low [CO2]
favors oxygenation
Importance of Photorespiration
- Internal cellular recycling of 3 gases:
CO2, O2, NH3
- Glycolate synthesis in the leaf decrease
toxic levels of O2
- Physiological defense against high
irradiance and thermal load (dissipation
of excess photochemical energy)
 - Contribution to cell amino acid pool o It reduces photorespiration and water
(Gly; Ser) loss
- Response to niche diversification scheme  CAM (Crassulacean Acid Metabolism)
o Similar to C4
Criteria Photorespiration Respiration
o PEP carboxylase fixes carbon at night in
mesophyll cells
Reactants RuBP + O2 CH2O + O2
o Calvin cycle occurs during the day
Products CO2, NH3, PGA CO2, H2O, ATP
o e.g. succulent, cactus, pineapple
o temporal isolation
Organelle(s) Chloroplast, Mitochondrion
involved peroxisome, CAM plants
mitochondrion
Energy Requiring Releasing - stomata opens at night
ATP None 36 ATP - stomata closes during the day when
formation transpiration is high
O2 > 21% O2 21% O2 - leaves have low surface to volume ratio
saturation in leaves
- cells have larger vacuoles, deeper
stomates, and denser guard cells
CO2 fixation pathways
- form malic acid (low pH) at night which
 C3 pathway (Calvin-Benson Cycle) disappear during daytime
o Fundamental carboxylating mechanism - succulent
in plants resulting to a net fixation of CO2
CAM pathway
– all plants undergo C3
o Examples: rice, tobacco, soybean - similar to C4 pathway except for
o Name was derived from the fact that o lacking specialized leaf anatomy
PGA is a 3-C compound (Krantz anatomy)
o Each turn of cycle: fix one molecule of o formation of C4 acids is temporal
CO2 (time), not spatial (space),
o Input: 3 ATP and 2 NADPH separation from decarboxylation
o Product: G3P/Triose phosphate and re-fixation via the C3 cycle
o 1 TP = 3 turns (3 CO2) - an ecological adaptation that evolved
o 6 CO2 = hexose in habitats where water shortage was
 C4 pathway or Hatch-Slack Pathway the dominant factor for plant survival
o CO2 assimilation pathway exhibiting CO2 - can be:
concentrating mechanism o obligate CAM - cactus
o Example: corn, sugarcane o facultative (inducible) CAM –
o Substrate: HCO3-, instead of CO2 pineapple
o First stable product: 4-C oxaloacetate
Three Major Plant Groups Based on CO2
o Spatial separation of steps
Fixation Pathway
o Releases CO2 in bundle sheath cell
when rubisco is present Characteri C3 C4 CAM
o Involves two cell types: stics
 Mesophyll cell: HCO3- fixation Leaf Mesophyll Mesophyll Mesophyl
anatomy with no with l with
 HCO3- / PEP → oxaloacetate →
distinct distinct large
malate or aspartate bundle bundle vacuoles
 Bundle sheath cell: Krantz anatomy sheath sheath
o C4 cycle increases the concentration of Carboxyla Rubisco PEPCases PEPcase
CO2 in bundle sheath cell where rubisco ting and and
is present enzyme Rubisco Rubisco
 (separation (separati o Plant dry weight can increase
in space) on in only when the irradiance is
time)
above the LC
Initial C3 acids: C4 acids: C4 acids:
producs of phosphogl oxaloacet oxaloace
o Differences in LC are caused by
CO2 ycerate ate, tate, differences in respiration rates:
fixation malate, malate, when RD is low, leaf requires less
aspartate aspartat light for the level of PS that will
e balance the lost CO2, thus LC is
Photorespi High Very low or Very low
also low
ration absent
Sun plant vs. Shade plant
Single Cell C4 Photosynthesis Criteria Sun plant Shade plant
Photosynthesis High Low irradiance
- ability to perform C4 photosynthesis saturation irradiance
within a single chlorechyma cell by High High Photoinhibition
intracellular partitioning of enzymes and irradiance photosynthesis (low PS)
organelles in two compartments Leaf Thicker Larger in area
morphology but thinner
Product Synthesis Total Less More
chl/reaction
 sucrose reaction
o main form of carbohydrate translocated center
in plants Chl b to a ratio Low High
o synthesized outside the chloroplast Stroma and More Less
thylakoid
 cytosol
proteins
 starch
o synthesized in the chloroplast stroma
where it is built up during the day and Carbon dioxide
used as energy source at night
- PS is enhance by higher [CO2] when
o produced and stored in amyloplast
stomates are partly closed
o “both are regulated by Pi
- Higher [CO2] favors CO2 concentration
concentration”
- The CO2 concentrating mechanisms in
Factors affecting C4 and CAM plants allow the leaf to
maintain high PS rate at lower
- light
intercellular CO2 values
- CO2 availability
- CO2 compensation point
- Temperature
o The [CO2] at which net
- Water
photosynthesis just balances
- Nutrients
respiratory loss (net CO2
- Leaf age
exchange is zero).
- Carbohydrate translocation
Temperature
Light irradiance
- Photosynthetic responses to temperature
- Light compensation point (LC)
depend on the species and growth
o Irradiance at which
condition
photosynthesis (CO2 uptake) just
- Optimum temp. for PS is usually defined
balances respiration (CO2
as the point at which the capacities of
evolved), where the net CO2
various steps of PS are balanced
exchange is zero
- PS is slow at cooler temperature and
increases to an optimum as temp. rises
 - C4 have higher optimum than C3
- C4 is also inefficient at low temp due to
membrane damage and chlorophyll
synthesis inhibition
Water
- Water as a photosynthetic reactants it
the:
o Source of e- to form NADPH
o Source of protons for ATP
synthesis
o Source of O2 for people to
breathe
- For every mole of CO2 absorbed by the
leaf, there is a corresponding amount of
water lost by the plant
- Transpiration ratio
o Amount of water lost per mole of
CO2 fixed
o CAM is most efficient in water
Nutrients
- The effect of nutrients in PS depends on
the specific photosynthetic step which a
specific nutrient affects
o Nitrogen – structural component
of chlorophyll and protein
o Phosphate – component of ATP
and needed to export/import TP
to and from the chloroplast
o Manganese, chlorine, and
calcium – needed in O2 evolution
during the photolysis of water
o Iron (in Fe-S center) – specific
effects on electron transport
o Magnesium – chlorophyll
component; needed for for
photoprotection
Internal Factors
- Leaf age
o As leaves grow, their ability to
photosynthesize increases until
they are fully expanded, then it
begins to decrease slowly until
the leaves senesce
- Carbohydrate translocation
o Effective transport of sucrose
from leaves to various sink organs
maintains rapid carbon dioxide
fixation
Signal Transduction Pathway
- One or more sequences of biochemical
reactions connecting the stimulus with a
cellular response
- Coordinate gene expression through
transcription factors (e.g. WRKY) with
environmental conditions and
development
- Involves secondary messengers,
hormones, protein kinases,
phosphatases, etc.
o Amplification of initial signal
Plant hormones
- Regulates growth and development
based on biological and environmental
influences
- Plant growth regulators (PGRs) –
synthesized in the lab
- Classics: auxin, cytokinin, gibberellin,
abscissic acid, ethylene
o Jasmonic acid (JA),
brassinosteroids
Auxin
- Auxein – to grow, to increase
- Apical dominance
- Cell elongation
- Rooting
- Acid-induced extrusion
Cytokinin
- Cytokinesis – cell division
- Xylem development
- Lateral growth of shoots and for stem
development
- Open stomata
Gibberellins
- Found I Gibberella fujikuroi
- Cell elongation
- Flowering and seed germination (more
fruits, no seeds)
Abscisic acid
 - Leaf senescence
- Maintains seed dormancy
- Closes stomata

Ethylene

- Fruit ripening
- Stress response
- Inhibits growth in the dark

Jasmonic acid

- In jasmine oil first identified

- Response to biotic stress
o Wounding; microbial and fungal
- Similar to auxin for actions

Brassinosteroids

- Stress responses
- Stem elongation
- Seed germination
- Pollen tube growth
- Cell differentation