HMM/SCM1414_Biology 1

CHAPTER 6

PHOTOSYNTHESIS
• Life on Earth is solar powered.
• Chloroplasts capture light energy from sun
and convert it to chemical energy stored in
sugars and other organic molecules.

6.1 Types of Nutrition

• Photosynthesis nourishes almost all the

living world directly or indirectly.
° All organisms use organic compounds for
energy and for carbon skeletons
synthesis of organic compounds.
° Obtain organic compounds by autotrophic
or heterotrophic nutrition.

6.1.1 Autotrophic Nutrition

• Autotrophs produce organic molecules from

CO2 and other inorganic raw materials
obtained from environment.
° Autotrophs are ultimate sources of
organic compounds for all heterotrophic
organisms.
° Producers of the biosphere.

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• Two groups based on energy source that

drives their metabolism.
° Photoautotrophs use light as energy
source to synthesize organic compounds.

Example, plants, algae, some other
protists, and some prokaryotes.
° Chemoautotrophs harvest energy from
oxidizing inorganic substances, such as
sulfur and ammonia.

Unique to prokaryotes.

6.1.2 Heterotrophic Nutrition

• Heterotrophs live on organic compounds

produced by other organisms.
° Consumers of the biosphere.
° Most feeds on other organisms.

Example, animals.
° Others decompose and feed on dead
organisms or organic litter, like feces and
fallen leaves.

Example, most fungi and many
prokaryotes.
° Almost all heterotrophs completely
dependent on photoautotrophs for food
and for oxygen, a by-product of
photosynthesis.

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6.2 The biophysics of light

• Thylakoids convert light energy into

chemical energy of ATP and NADPH.
• Light is a form of electromagnetic
radiation.
• Light travels in rhythmic waves.
• Distance between crests of
electromagnetic waves = wavelength.
wavelength.
° Wavelengths range from less than a
nanometer (gamma rays) to more than a
kilometer (radio waves).
• Entire range of electromagnetic radiation =
electromagnetic spectrum.
spectrum.
(Figure 10.6, Campbell, page 186)
• The most important segment for life is
narrow band between 380 to 750 nm =
light.
visible light.

• Light is composed of small particles, or

packets of energy, the photons
photons.
° Photons have fixed quantities of energy.
• Amount of energy packaged in a photon is
inversely related to its wavelength.
° The shorter the wavelengths, the more
the energy.

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• Atmosphere selectively screens out most

wavelengths, allowing only visible light to
pass in significant quantities.
° Visible light drives photosynthesis.

• When light meets matter, it may be

reflected, transmitted, or absorbed.
° Different pigments absorb photons of
different wavelengths.
° Wavelengths that are absorbed
disappear.
° A leaf looks green because chlorophyll
absorbs red and blue light, while
transmitting and reflecting green light.
(Figure 10.7, Campbell, page 186)
• A spectrophotometer measures ability of a
pigment to absorb various wavelengths of
light. (Figure 10.8, Campbell, page 187)
° It beams narrow wavelengths of light
through solution containing pigment and
measures fraction of light transmitted at
each wavelength.
° An absorption spectrum plots a pigment’s
light absorption versus wavelength.
(Figure 10.9 (a), Campbell, page 187)
• Light reaction can perform work with those
wavelengths of light that are absorbed.

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• Thylakoid contains several pigments that

differ in their absorption spectra.
° Chlorophyll a, the dominant pigment,
absorbs best in red and violet-blue
wavelengths and least in green.
° Other pigments have different absorption
spectra.
• Collectively, these photosynthetic

pigments determine an overall action

spectrum for photosynthesis.
(Figure 10.9 (b), Campbell, page 187)
° Action spectrum measures changes in
some measure of photosynthetic activity
(for example, O2 release) as wavelength
is varied.
• Action spectrum of photosynthesis was
first demonstrated by Thomas Engelmann
(1883):
° Different segments of filamentous alga
were exposed to different wavelengths of
light.
° Areas receiving wavelengths favorable to
photosynthesis produced excess O2.
° Most aerobic bacteria clustered along
segment of algal filament emitting most
O2, i.e., segments in red and blue portion
of spectrum. (Figure 10.9 (c), Campbell, page 187)

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• Action spectrum of photosynthesis does

not match exactly absorption spectrum of
any one photosynthetic pigment, including
chlorophyll a.
• Only chlorophyll a participates directly in
the light reaction, but accessory
photosynthetic pigments absorb light and
transfer energy to chlorophyll a.
° Chlorophyll b has slightly different
absorption spectrum and funnels energy
from these wavelengths to chlorophyll a.
° Carotenoids funnel energy from other
wavelengths to chlorophyll a and also
participate in photoprotection against
excessive light:
 Absorb and dissipate excessive light

energy that would damage chlorophyll.

 Also interact with oxygen to form

reactive oxidative molecules that could

damage the cell.

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6.3 How the two photosystems of plants

work together

• When a molecule absorbs a photon, one of

its electrons is elevated to an orbital with
more potential energy.
° The electron moves from its ground state
to an excited state.
• Excited electrons are unstable.
• Generally, they drop to their ground state in
a billionth of a second, releasing heat
energy.

• In the thylakoid membrane, chlorophyll is

organized with proteins and smaller organic
molecules into photosystems.
photosystems.
• Photosystem is composed of a reaction
center surrounded by a light-
light-harvesting
complex.
complex (Figure 10.12, Campbell, page 189)
• Each light-harvesting complex consists of
pigment molecules (chlorophyll a,
chlorophyll b, and carotenoid molecules)
bound to particular proteins.
• Light-harvesting complexes act like light-
gathering “antenna complexes” for reaction
center.
• When antenna molecule absorbs photon, it
is transmitted from molecule to molecule
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until it reaches a particular chlorophyll a

molecule, the reaction center..
• At the reaction center is a primary electron
acceptor,
acceptor, which accepts an excited
electron from chlorophyll a.
° The solar-powered transfer of an electron
from chlorophyll a to the primary
electron acceptor is the first step of light
reactions.

• Two types of photosystems in thylakoid

membrane.
1. Photosystem I (PS I) has a reaction
center chlorophyll a that has absorption
peak at 700 nm.
2. Photosystem II (PS II) has a reaction
center chlorophyll a that has absorption
peak at 680 nm.
° The differences between these reaction
centers (and their absorption spectra) lie
not in the chlorophyll molecules, but in
the proteins associated with each
reaction center.
° Both photosystems work together to use
light energy to generate ATP and NADPH.

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6.4 Light dependent reaction

• Two possible routes for electron flow:

cyclic and non-cyclic.

6.4.1 Non
Non--cyclic Electron Flow (Cyclic
Photophosphorylation)
(Figure 10.13, Campbell, page 190)

• The predominant route - produces both ATP

and NADPH:
1. Photosystem II absorbs a photon of light.
One of the electrons of P680 is excited to
a higher energy state.
2. Electron captured by primary electron
acceptor, leaving reaction center
oxidized.
3. An enzyme extracts electrons from water
and supplies them to oxidized reaction
center. Water split into two H+ and an
oxygen atom that combines with another
oxygen atom to form O2.
4. Photoexcited electrons pass along an
electron transport chain before ending up
at an oxidized photosystem I reaction
center. (Electron transport chain made up
of plastoquinine, Pq, a cytochrome
complex, and plastocyanin, Pc.)

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5. As these electrons “fall” to a lower

energy level, their energy is used to
produce ATP.
6. Meanwhile, light energy excited an
electron of PSI’s P700 reaction center.
7. Photoexcited electron is captured by
PSI’s primary electron acceptor, creating
an electron “hole” in P700.
8. Hole is filled by electron from PS II.
9. Photoexcited electrons are passed from
PSI’s primary electron acceptor down a
second electron transport chain through
the protein ferredoxin (Fd).
10. NADP+ reductase transfers electrons from
Fd to NADP+.
11. Two electrons are required for NADP+’s
reduction to NADPH.
12. NADPH will carry reducing power of these
high-energy electrons to Calvin cycle.

• Light reactions use solar power of photons

absorbed by both photosystem I and
photosystem II to provide
(i) chemical energy – ATP; and
(ii) reducing power - in the form of
electrons carried by NADPH.

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6.4.2 Cyclic Electron Flow

(Figure 10.15, Campbell, page 101)

• Under certain conditions, photoexcited

electrons from photosystem I can take an
alternative pathway, cyclic electron flow.
° Excited electrons cycle from reaction
center to a primary acceptor, along an
electron transport chain, and return to
the oxidized P700 chlorophyll.
° As electrons flow along electron
transport chain, they generate ATP by
cyclic photophosphorylation.
photophosphorylation.
° No NADPH produced and no release of
oxygen.

What is the function of cyclic electron flow?

• Noncyclic electron flow produces ATP and

NADPH in roughly equal quantities.
• However, Calvin cycle consumes more ATP
than NADPH.
• Cyclic electron flow allows the chloroplast
to generate enough surplus ATP to satisfy
the higher demand for ATP in Calvin cycle.

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6.4.3 Chemiosmosis
(Figure 10.17, Campbell, page 193)

 Formation of ATP in light reaction as a

result of a pH gradient across thylakoid
membrane.

1. Photon strikes pigment molecule in

PSII.
2. Excited electron is passed along
electron carriers.
3. Water is split – O2 is released and H+
remains in thylakoid space.
4. Energy released as electron passes
through electron carriers between PSII
and PSI is used by proton pump to pump
H+ into thylakoid space.
5. A pH gradient develops between inside
and outside of thylakoid sac.
6. Inside high H+ concentration (lower pH).
7. A strong diffusion gradient is set up.
8. H+ cross back across membrane
through ATPase synthetase complex.
9. Energy released as H+ flow down their
gradient is used for synthesis of ATP
from ADP and P.
10. 3 H+ pass through ATPase synthetase
complex to make 1 ATP.

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11. NADP is the final electron acceptor

producing NADPH

Figure: Electron Transport and Chemiosmosis during

Photosynthesis

• Light-reaction produces ATP and NADPH on

stroma side of thylakoid, where Calvin
cycle reactions take place.

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CHAPTER 6- 2nd part.

PHOTOSYNTHESIS
6.5 Light independent reaction

6.5.1 Calvin cycle/C3 cycle

• Cycle regenerates its starting material

after molecules enter and leave cycle.
• Cycle is anabolic - uses energy to build
sugar from smaller molecules.
• Carbon enters cycle as CO2 and leaves as
sugar.
• Cycle uses energy of ATP and reducing
power of electrons carried by NADPH to
make sugar.
• Actual sugar product of cycle is not
glucose, but glyceraldehyde
glyceraldehyde-
ehyde-3-phosphate
(G3P).
• Each turn of cycle fixes one carbon.
• Net synthesis of one G3P molecule,
requires three cycles, fixing three
molecules of CO2.
• One glucose molecule requires six cycles
and fixation of six CO2 molecules.

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• Three phases.
(Figure 10.18, Campbell, page 194)

Phase 1: Carbon fixation

• CO2 is attached to a 5C sugar, ribulose

bisphosphate (RuBP).
° Catalyzed by RuBP carboxylase or
rubisco.
 Most abundant protein in chloroplasts
and on Earth.
° 6C intermediate formed is unstable and
splits in half to form two 3C molecules of
3-phosphoglycerate for each CO2.

Phase 2: Reduction

• Each 3-phosphoglycerate receives another

phosphate group from ATP to form 1,3-
bisphosphoglycerate.
• A pair of electrons from NADPH reduces
each 1,3-bisphosphoglycerate to G3P (3C).
° Electrons reduce a carboxyl group to
aldehyde group of G3P, which stores
more potential energy.
• For every three CO2 (3C) and three RuBP
(15C), six molecules of G3P (18C) are
formed.

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° One of these six G3P (3C) is a net gain of

carbohydrate.
 This molecule exit cycle to be

used by plant cell.

Phase 3: Regeneration

• The other five G3P (15C) remain in cycle to

regenerate three RuBP. Carbon skeletons
of five molecules of G3P are rearranged in a
series of reactions to regenerate three
molecules of RuBP.
• Three ATP used.

• For net synthesis of one G3P molecule, the

Calvin cycle consumes nine ATP and six
NADPH.
• G3P from Calvin cycle is the starting
material for metabolic pathways that
synthesize other organic compounds,
including glucose and other carbohydrates.

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6.5.2 Photorespiration

• One major problem facing terrestrial plants

is dehydration.
• Stomata - major route for gas exchange
(CO2 in; O2 out), and for evaporative loss of
water.
• On hot, dry days, plants close stomata to
conserve water.

• In C3 plants, initial fixation of CO2 occurs

via rubisco, forming 3C compound, 3-
phosphoglycerate.
° C3 plants - rice, wheat, and soybeans.
• When stomata partially close on a hot, dry
day, CO2 levels drop as CO2 is used in
Calvin cycle.
• At same time, O2 levels rise as light
reaction converts light to chemical energy.
• Rubisco normally accepts CO.
• But, when O2:CO2 ratio increases (on a hot,
dry day with closed stomata), rubisco can
add O2 to RuBP.
• RuBP then splits into a 3C piece (3-
phosphoglycerate) and a 2C piece
(phosphoglycolate) in a process called
photorespiration.

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° 2C fragment is exported from chloroplast

and degraded to CO2 by mitochondria and
peroxisomes.

O2 Rubisco 3-Phosphoglycerate
+
+
RuBP Phosphoglycolate

CO2

° Unlike normal respiration, this process

produces no ATP, but consumes ATP.
° Unlike photosynthesis, photorespiration
does not produce organic molecules, but
decreases photosynthetic output by
siphoning organic material from Calvin
cycle.

• Hypothesis for the existence of

photorespiration – metabolic relic from a
much earlier time:
° When rubisco first evolved, atmosphere
had far less O2 and more CO2 than it does
today.

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° Inability of active site of rubisco to

exclude O2 would have made little
difference.
• Today it does make a difference.
° Photorespiration can drain away as much
as 50% of carbon fixed by Calvin cycle on
a hot, dry day.

6.4.3 C4 / Hatch-
Hatch-Slack pathway

• Certain plant species have evolved

alternate modes of CO2 fixation to minimize
photorespiration.
• C4 plants first fix CO2 in a 4C compound.
° Example of C4 plant: sugarcane and corn.
• A unique leaf anatomy is correlated with
mechanism of C4 photosynthesis.
(Figure 10.19, Campbell, page 196)

C3 plants

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C4 plants

• Two types of photosynthetic cells in C4

plants: bundle-sheath cells and mesophyll
cells.
(a) Bundle-
Bundle-sheath cells - arranged into
tightly packed sheaths around veins of
leaf.
(b) Mesophyll cells - more loosely arranged
cells located between bundle sheath
and leaf surface.
• Calvin cycle confined to chloroplasts of
bundle-sheath cells.
(Figure 10.19, Campbell, page 196)

• Phosphoenolpyruvate carboxylase (pepco),

adds CO2 to phosphoenolpyruvate (PEP) to
form oxaloacetate (OAA) in mesophyll cells.
° PEP carboxylase has very high affinity for
CO2 and can fix CO2 efficiently when

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rubisco cannot (i.e., on hot, dry days

when stomata are closed).
• OAA converted to malate (4C).
• Malate exported into bundle-sheath cells.
° Malate is decarboxylated forming
pyruvate, releasing CO2.
° Rubisco starts Calvin cycle, using
abundant supply of CO2.
° Pyruvate returns to mesophyll cells and
regenerated to PEP. ATP is used.

• By pumping CO2 into bundle sheath cells,

CO2 levels are kept high for rubisco to
accept CO2 and not O2.
• Photorespiration is minimized; sugar
production is enhanced.
• C4 plants thrive in hot regions with intense
sunlight.

6.5.4 Crassulacean Acid /CAM pathway

(Figure 10.20, Campbell, page 197)

• CAM plants - cacti, pineapples, and several

other plant families.
° CAM - crassulacean acid metabolism.
° Open stomata at night and close during
the day.

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 Temperatures lower at night, and

humidity is higher.
° Night - plants fix CO2 into a variety of
organic acids in mesophyll cells.
 Organic acids stored in vacuoles.

° Day - light reactions supply ATP and

NADPH; CO2 is released from the organic
acids and enters Calvin cycle.
• Both C4 and CAM plants add CO2 into
organic intermediates before it enters
Calvin cycle.
° In C4 plants, carbon fixation and Calvin
cycle are spatially separated.
° In CAM plants, carbon fixation and Calvin
cycle are temporally separated.
• Both eventually use Calvin cycle to make
sugar from CO2.

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6.6 Factors affecting photosynthesis

photosynthesis rate

• Rate of photosynthesis may be measured

by quantity of CO2 or O2 consumed per unit
time.

(1) Light intensity

° As light intensity increases,

photosynthetic rate increases until a
point is reached where rate begins to
level off.
° At low light intensity, photosynthesis
occurs slowly because only a small
quantity of ATP and NADPH is created
by the light dependent reactions.
° As light intensity increases, more ATP
and NADPH are created, thus increasing
the photosynthetic rate.
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° At high light intensity, photosynthetic

rate levels out, not due to light intensity
but due to other limiting factors, e.g.,
competition between O2 & CO2 for
active site on RUBP carboxylase.

(2) CO2 concentration

° As CO2 concentration increases, rate of

photosynthesis increases.
° At high concentrations, rate of
photosynthesis begins to level out due
to factors not related to carbon dioxide
concentration.
 Example, some enzymes of

photosynthesis might be working at

their maximum rate.
° In general, CO2 is found in low
concentration in atmosphere.
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° So, atmospheric CO2 levels may be a

major limiting factor on photosynthesis
when at low levels.

(3) Temperature

° As temperature increases above

freezing, rate of photosynthesis
increases.
 This occurs because molecules are

moving more quickly and there is a

greater chance of a collision resulting
in a chemical reaction.
° At some point, a temperature is
reached that is an optimum
temperature.
° Photosynthetic reaction rate is at its
quickest rate at this point.

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° Above that temperature, enzymes begin

to denature (as in RUBP carboxylase),
slowing rate of photosynthesis until a
temperature is reached where
photosynthesis does not occur at all.

(4) Light duration

° Photosynthesis is only affected by light
duration in that it only occurs during
periods of light.

(5) Level of air pollution

° Low levels of O3 and SO2 are very
damaging to some plant leaves.
° Soot can block stomata and prevent
light from reaching chloroplasts by
coating leaf.

(6) Oxygen
° Competes with CO2 for active site of
RuBP carboxylase.
° Relatively high concentrations of O2, for
example the 21% in our atmosphere,
inhibit photosynthesis.
° O2 does not inhibit CO2 fixation in C4
plants.

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(7) Water
° Slight lack of water can lead to severe
loss of carbohydrate yield.

(8) Chlorophyll concentration

° Can become a limiting factor if
chlorophyll levels are abnormally low.
° May be caused by disease, mineral
deficiency, and senescence.
° Iron, magnesium, nitrogen, and sunlight
are necessary for chlorophyll
production - lack of any one of these
can lead to yellowing of leaves.

Compensation Point

 Point during photosynthesis where rate of

photosynthesis exactly matches rate of
respiration, i.e., input of CO2 or O2 is same
as output of the other by a plant.
• Point is reached during early mornings and

late evenings.
• Summary of compensation point:

° Rate of photosynthesis = rate of

respiration.
° Amount of O2 produced by plant is equal
to amount used up at that point in time.

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° Thus, there is no net output of O2 by the

plant.
° There is nil effective photosynthesis.

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