Homework questions to be emailed to
biovcc@yahoo.com
0
List the wavelengths of light that are most effective for photosynthesis
2 Compare
cyclic and noncyclic electron flow and explain the relationship between these
components of the light reaction
3 Describe
the role of ATP and NADPH in the Calvin cycle
4 Describe
what happens to rubisco when the O2 concentration is much higher than CO2
5
Describe two important photosynthetic adaptations that minimize photorespiration
LECTURE Notes Photosynthesis and Energy
Photosynthesis
WHY STUDY PHOTOSYNTHESIS?
•Photosynthesis is arguably
the most important biological process on earth.
• By liberating oxygen and
consuming carbon dioxide, it has transformed the world into the hospitable
environment we know today.
•.
Directly or
indirectly, photosynthesis fills all of our food requirements and many of our
needs for fiber and building materials.
•The energy stored in
petroleum, natural gas and coal all came from the sun via photosynthesis, as
does the energy in firewood, which is a major fuel in many parts of the world.
. This being the case,
scientific research into photosynthesis is vitally important.
If we can understand and
control the intricacies of the photosynthetic process, we can learn how to
increase crop yields of food, fiber, wood, and fuel, and how to better use our
lands.
. The energy-harvesting
secrets of plants can be adapted to man-made systems which provide new,
efficient ways to collect and use solar energy.
Respiration is an Oxidation-Reduction process
–
Loss of electrons from one substance = oxidation.
–
Addition of electrons to a substance = reduction.
–
Oxidizing agent - accepts electrons.
–
Reducing agent - gives up electrons.
•
Oxygen - very strong oxidizing agent (hence: “oxidizing” or
“oxidation”)
Redox
reactions
E.g.
Na
+
Cl
->
Na+
+
Cl-
According to the first law
of thermodynamics, energy cannot be created or destroyed, but it can be
transferred from one place to another and transformed from one form to another.
During photosynthesis, energy in the form of light is transferred from the
sun, some 92 million miles away, to a pigment molecule in a photosynthetic
organism such as a plant. What follows is an interesting series of energy
transformations in which light energy is transformed into electrochemical energy
and then into energy stored within chemical bonds.
Albert Einstein formulated
the photon theory of light in which he proposed that light is composed of
discrete particles called photons—massless particles traveling in a wavelike
pattern and moving at the speed of light. Each photon contains a specific amount
of energy. An important difference between the various types of electromagnetic
radiation is the amount of energy found in the photons.
Shorter wavelength
radiation carries more energy per unit of time than longer wavelength radiation.
For example, the photons of gamma rays carry more energy than those of radio
waves.
The photons found in gamma
rays, X-rays, and UV rays have very high energy. When molecules in cells absorb
such energy, the effects can be devastating. Such types of radiation can cause
mutations in DNA and even lead to cancer. By comparison, the energy of photons
found in visible light is much milder. Molecules can absorb this energy in a way
that does not cause permanent harm. Next, we will consider how molecules in
living cells absorb the energy within visible light.
Pigments Absorb Light
Energy
When light strikes an
object, one of three things will happen.
•
First, light may simply pass through the object.
•
Second, the object may change the path of light toward a different direction.
•
A third possibility is that the object may absorb the light.
The term pigment is used
to describe a molecule that can absorb light energy. When light strikes a
pigment, some of the wavelengths of light energy are absorbed, while others are
reflected.
For example, leaves look
green to us because they reflect radiant energy of the green wavelength. Various
pigments in the leaves absorb the other light energy wavelengths. At the
extremes of color reflection are white and black.
A white object reflects
nearly all of the visible light energy falling on it, whereas a black object
absorbs nearly all of the light energy. This is why it’s coolest to wear white
clothes on a sunny, hot day.
What do we mean when we
say that light energy is absorbed?
In the visible spectrum,
light energy is usually absorbed by boosting electrons to higher energy levels .
Recall from Chapter 2 that
electrons are located around the nucleus of an atom. The location in which an
electron is likely to be found is called its orbital. Electrons in different
orbitals possess different amounts of energy.
Figure 2.9 Energy levels
of an atom’s electrons
For an electron to absorb
light energy and be boosted to an orbital with a higher energy, it must overcome
the difference in energy between the orbital it is in and the orbital to which
it is going.
For this to happen, an
electron must absorb a photon that contains precisely that amount of energy.
In the case of
photosynthetic pigments, however, a different event happens that is critical for
the process of photosynthesis.
Rather than releasing
energy, an excited electron in a photosynthetic pigment is removed from that
molecule and transferred to another molecule where the electron is more stable.
When this occurs, the energy in the electron is said to be “captured,” because
the electron does not readily drop down to a lower energy level and release heat
or light.
The role of the reaction
center is to quickly remove the high energy electron from P680 and transfer it
to another molecule, where the electron will be more stable. This molecule is
called the primary electron acceptor.
The transfer of the
electron from P680 to the primary electron acceptor is remarkably fast. It
occurs in less than a few picoseconds! (One picosecond equals one-trillionth of
a second, also noted as 10-12 s.)
Because this occurs so
quickly, the excited electron does not have much time to release its energy in
the form of heat or light.
After the primary electron
acceptor has received this high energy electron, the light energy has been
captured and can be used to perform cellular work.
As discussed earlier, the
work it performs is to synthesize the energy intermediates ATP and NADPH.
The Splitting of
Water
•
Chloroplasts split water into
–
Hydrogen and oxygen, incorporating the electrons of hydrogen into sugar
molecules
•
The light reactions produce three chemical products: ATP, NADPH, and O2.
•
ATP and NADPH are energy intermediates that provide the needed energy and
electrons to drive the Calvin cycle.
•
Like NADH, NADPH (nicotinamide adenine dinucleotide phosphate) is an electron
carrier that can accept two electrons. Its structure differs from NADH by the
presence of an additional phosphate group.
•
As we have just seen, the Calvin cycle begins by using carbon from an inorganic
source, that is, CO2, and ends with organic molecules that will be used by the
plant to make other compounds.
•
You may be wondering why CO2 molecules cannot be directly linked to form these
larger molecules.
•
The answer lies in the number of electrons that orbit carbon atoms.
•
In CO2, the carbon atom is considered electron poor.
•
Oxygen is a very electronegative atom that monopolizes the electrons it shares
with other atoms.
•
In a covalent bond between carbon and oxygen, the shared electrons are closer to
the oxygen atom.
•
By comparison, in an organic molecule, the carbon atom is electron rich.
•
During the Calvin cycle, ATP provides energy and NADPH donates high-energy
electrons, so the carbon originally in CO2 has been reduced.
•
The Calvin cycle combines less electronegative atoms with carbon atoms so that
C—H and C—C bonds are formed.
•
This allows the eventual synthesis of larger organic molecules including
glucose, amino acids, and so on.
•
In addition, the covalent bonds within these molecules are capable of storing
large amounts of energy.
The
Calvin cycle proceeds in three stages: carboxylation, reduction, and
regeneration
•
The Calvin cycle
Carbon mobilization in vascular plants
We learned that rubisco
functions as a carboxylase because it adds a CO2 molecule to RuBP, an organic
molecule, to create two molecules of 3 phosphoglycerate (3PG).
RuBP +CO2 → 2 3PG
For most species of
plants, the incorporation of CO2 into RuBP is the only way for carbon fixation
to occur. Because 3PG is a three-carbon molecule, these plants are called C3
plants. Examples of C3 plants include wheat and oak trees.
About 90% of the plant
species on Earth are C3 plants. Researchers have discovered that the active site
of rubisco can also function as an oxygenase, although its affinity for CO2 is
over 10-fold better than that for O2. Even so, when O2 levels are high and CO2
levels are low, rubisco adds an O2 molecule
to RuBP. This creates only
one molecule of 3-phosphoglycerate and a two-carbon molecule called
phosphoglycolate. The phosphoglycolate is then dephosphorylated to glycolate
and released from the chloroplast.
In a series of several
steps, the two-carbon glycolate is eventually oxidized in other organelles to
produce an organic molecule plus a molecule of CO2.
RuBP O2 →
3-phosphoglycerate
Phosphoglycolate →
Glycolate + An organic molecule CO2
This process, called
photorespiration, uses O2 and liberates CO2.
Photorespiration is
considered wasteful because it reverses the effects of photosynthesis.
This reduces the ability
of a plant to make carbohydrates and thereby limits plant growth.
Photorespiration is more
likely to occur when plants are exposed to a hot and dry environment.
Under these conditions,the
stomata of the leaves close, inhibiting the uptake of CO2 from the air and
trapping the O2 that is produced by photosynthesis.
When the level of CO2 is
low and O2 is high, photorespiration is favored.
When rubisco first evolved
some 3 billion years ago, the atmospheric oxygen level was low, so
photorespiration would not have been a problem.
Another view is that
photorespiration may have a protective advantage. On hot and dry days when the
stomata are closed, CO2 levels within the leaves will fall, and O2 levels will
rise. Under these conditions, highly toxic oxygen-containing molecules such as
free radicals may be produced that could damage the
plant. ??????
. Many plants that evolved
in arid or hot environments display an adaptation known as C4 photosynthesis,
which improves productivity by aiding CO2 absorption. An amazing 30,000 plant
species utilize C4 photosynthesis, which is thought to have evolved on about 70
separate occasions. In addition, 7,500 other plant species possess a variation
of C4 photosynthesis known as CAM (crassulacean acid metabolism; see Chapter 8).
However, many other plants that grow in hot environments lack C4 photosynthesis.
Rice, a staple crop for much of the world’s population, is a prominent example.
Agricultural scientists envision using genetic engineering techniques to endow
rice with C4 photosynthesis with the goal of increasing this crop’s
productivity.
•
The CAM pathway is similar to the C4 pathway
8.14
Crassulacean acid metabolism (CAM) (Part 1)
8.14
Crassulacean acid metabolism (CAM) (Part 2)
•
other
I. Photosynthesis: Light is important to
plants
II. Properties of Light
- A. Light is part of the electromagnetic
spectrum -- Wave properties
- 1. Wavelength= the distance between crests
of the wave
The
Spectrum
- a. TV waves are very long
wavelengths -> Infra-red (IR) (appear black)
b. Ultraviolet (looks black) -> X-rays wavelengths are very
short
c. Visible light are the colors you see (each color has a
different wavelength)
X-ray---UV--380nm-------------------------760nm---IR---TV
appears black violet blue green yellow orange red black
tanning heat lamps
(nanometer - nm = 1 billionth of a meter)
- B. Particle nature of radiation
III. Pigments
- A. Light
[REQUIRED READING] must first be absorbed in order to produce a
biological effect
- 1. absorbed by pigments
(photoreceptors)
2. example - in photosynthesis, chlorophyll & carotenoids
B. Absorption
spectrum - how much light of each wavelength is absorbed by a
particular pigment - see right ---> [from MIT's Photosynthesis
Hypertextbook]
- 1. Example - absorption spectrum of
chlorophyll (green pigment => absorbs red and blue)
C. Action spectrum - how much of a
physiological process occurs at each particular wavelength of light
- 1. Similar to absorption spectrum but
not identical
IV. Summary
of Photosynthesis
- A. General
information about photosynthesis [REQUIRED READING]
- 1. Green plants, algae (seaweeds), bacteria
2. Importance - required for the existence of other life forms (food
chain) and source of atmospheric oxygen
B. Definition - synthesis of organic compounds
from water and carbon dioxide using energy absorbed by pigments from
sunlight
6 CO2 + 12 H2O ----------> C6H12O6 + 6 02 + 6 H2O
^ sugar
light (glucose)
pigments
enzymes
C.Chloroplasts - in eukaryotic cells
granum w/chlorophyll
(round stacks of plates)
stroma w/enzymes for Calvin
(jello)
double membrane
- Structure
of a chloroplast
- Structure
of a thylakoid
- See
image of chloroplasts in a leaf of Elodea
- See
chloroplasts in an algal (Zygnema), x.s.
D. Two stages of photosynthesis
- 1. Light reactions (light dependent
reactions) - in grana of chloroplasts
- a. High energy compounds involved in
light reaction
1) ADP + Pi + energy <-> ATP
adenosine inorganic
diphosphate phosphate
2) NADP+ + 2e- + 2H+ <-> NADPH2 + H+
(nicotinamide adenine dinucleotide phosphate)
b. OVERALL - use of light energy to generate two high-energy
compounds, ATP and NADPH2
- 1) ATP
- a) When light is absorbed by
chlorophyll, some of its electrons become excited and leap
out of the chlorophyll molecule, grabbed by energy
receptors.
b) The energy of these electrons is used to make ATP from
ADP + Pi
2) NADPH2
- a) When light is absorbed by
chlorophyll, some of its electrons become excited and leap
out of the chlorophyll molecule, grabbed by energy
receptors.
b) These electrons are then used to convert NADP+ to NADPH2
3) The lost electrons in
chlorophyll are replaced from electrons of oxygen in water; When
e- are removed from water, oxygen is produced as a by-product of
photosynthesis, water is split -> 2H+ (protons) + 2e- + 1/2 O2
(gas)
(Note - NADP+ + 2e- + 2H+ <-> NADPH2)
2. Dark reactions (Calvin Cycle or light
independent reactions), in stroma of chloroplasts - occur same time as
light reaction but does not require light
- a. OVERALL - using CO2 to make
carbohydrate (sugar)
b. Requires energy of ATP and NADPH2 (from
light reactions)
c. Uses enzymes located in the stroma of the chloroplasts; enzyme
speeds up chemical reaction
d. Formula for Calvin Cycle (named for Melvin
Calvin [1911-1997] who died in January of 1997)
6CO2 + 12NADPH2 + 18ATP ---> 1 C6H12O6 + 12NADP + 18ADP + 18Pi + 6H2O
^ glucose
enzymes
E. Summary of some key points in
photosynthesis
- 1. Photosynthesis is the major
energy-storing process of life (light energy stored as chemical energy
in organic compounds)
2. CO2 and H2O are raw materials
3. Products are sugar and oxygen
4. Light energy is absorbed by pigments and drives the reactions of
photosynthesis
5. ATP and NADPH2 are formed during the light
reactions
6. Oxygen of water is liberated as a gas
7. Steps of Calvin Cycle are controlled by enzymes
8. Light reactions occur in the grana
Dark reactions occur in the stroma
Links relating to photosynthesis
Why
Study Photosynthesis: An essay on the significance of photosynthesis on
living organism, especially humans
Photosynthesis
from Newton's Apple
Photosynthesis:
An excellent review
Photosynthetic
Antennas: Advanced information
How
a rainbow is formed
The Light Reaction:
Detailed but most useful
The Dark Reaction:
Detailed but most useful


