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.
° 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.

Two groups based on energy source that

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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.
° Wavelengths range from less than a
nanometer (gamma rays) to more than a
kilometer (radio waves).
Entire range of electromagnetic radiation =
electromagnetic spectrum.
(Figure 10.6, Campbell, page 186)

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The most important segment for life is

narrow band between 380 to 750 nm =
visible light.

Light is composed of small particles, or

packets of energy, the 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.
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)

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A spectrophotometer measures ability of a

pigment to absorb various wavelengths of
light. (Figure 10.8, Campbell, page 187)

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° 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)

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Light reaction can perform work with those

wavelengths of light that are absorbed.
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)

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°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

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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.
Photosystem is composed of a reaction
center surrounded by a light-harvesting
complex. (Figure 10.12, Campbell, page 189)

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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 until
it reaches a particular chlorophyll a
molecule, the reaction center.
At the reaction center is a primary
electron acceptor, which accepts an

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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-cyclic Electron Flow

(Non- 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
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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.)
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

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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)

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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.
° 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)

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 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.
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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.
11. NADP is the final electron acceptor
producing NADPH

Figure: Electron Transport and Chemiosmosis during

Photosynthesis

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Light-reaction produces ATP and NADPH on

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

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6.5 Light independent reaction

6.5.1 Calvin cycle/C 3 cycle

Cycle regenerates its starting material after

molecules enter and leave cycle.
Cycle is anabolic - uses energy to build sugar
from smaller molecules.
1. Carbon enters cycle as CO2 and leaves

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as sugar.
2. Cycle uses energy of ATP and
reducing power of electrons carried by
NADPH to make sugar.
3. Actual sugar product of cycle is not

glucose, but glyceraldehyde-3-

phosphate (G3P).
4. Each turn of cycle fixes one carbon.
5. Net synthesis of one G3P molecule,

requires three cycles, fixing three

molecules of CO2.
6. One glucose molecule requires six

cycles and fixation of six CO2

molecules.

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7. 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

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(15C), six molecules of G3P (18C) are formed.

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

carbohydrate. Out of the 6 molecules of
G3P, one will be converted to
sugar/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 C 3 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.
° 2C fragment(phosphoglycolate) is

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exported from chloroplast and degraded

to CO2 by mitochondria and peroxisomes.

O2
Rubisco 3-Phosphoglycerate
+ +
RuBP Phosphoglycolate

CO2

1. Unlike normal respiration, this process

produces no ATP, but consumes ATP.
° 2. 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.
° Inability of active site of rubisco to

exclude O2 would have made little

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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-Slack pathway

Certain plant species have evolved alternate

modes of CO2 fixation to minimize
photorespiration.
C 4 plants first fix CO2 in a 4C compound.
° Example of C4 plant: sugarcane and corn.

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A unique leaf anatomy is correlated with

mechanism of C4 photosynthesis.
(Figure 10.19, Campbell, page 196)

C3 plants

C4 plants

Two types of photosynthetic cells in C4

plants:
(a) bundle-sheath cells
(b) mesophyll cells.
(a) Bundle-sheath cells - arranged into
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tightly packed sheaths around veins

of leaf.
(b) Mesophyll cells - more loosely
arranged cells located between
bundle sheath and leaf surface.

Calvin cycle C3:-

confined to chloroplasts of bundle-sheath
cells.
(Figure 10.19, Campbell, page 196)

Mesophyll Cell :- C4

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

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.

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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 stomata during the day.

 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.

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° In CAM plants, carbon fixation and Calvin

cycle are temporally..pemisahan
masa= siang – malam separated.
Both eventually use Calvin cycle to make
sugar from CO2.

6.6 Factors affecting photosynthesis

rate

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• Rate of photosynthesis may be measured

by quantity of CO2 consumed per unit time.
• or O2 released 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.
° At high light intensity, photosynthetic rate
levels out, not due to light intensity but due

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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.
° So, atmospheric CO2 levels may be a major
limiting factor on photosynthesis when at low
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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.
° Above that temperature, enzymes begin
to denature (as in RUBP carboxylase),

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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.

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

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(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.
° Thus, there is no net output of O2 by the
plant.
° There is nil effective photosynthesis.

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