bioenergetics Flashcards
Bioenergetics
Bioenergetics is the quantitative study of energy relationships and energy conversions in biological systems. Biological energy transformations obey the laws of thermodynamics.
ATP a Chemical Link
With the emergence of photosynthesis on earth, molecular oxygen began to accumulate slowly in the atmosphere. The presence of free oxygen made possible the evolution of respiration. Respiration releases great deal of energy, and couples some of this energy to the formation of adenosine triphosphate (ATP) molecules. ATP is a kind of chemical link between catabolism and anabolism.
Photosynthesis
The process of photosynthesis helps understand some of the principles of energy transformation (Bioenergetics) in living systems. Photosynthetic organisms (higher land plants for instance) use solar energy to synthesize organic compounds (such as carbohydrates) that can not be formed without the input of energy.
glycolysis and respiration are the processes where
by the energy stores in carbohydrate is released in a controlled manner. So the photosynthesis acts as an energy-capturing while respiration as an energy releasing process.
Photosynthesis
Photosynthesis can be defined as the process in which energy-poor inorganic oxidized compounds of carbon (i.e. CO2) and hydrogen (i.e. mainly water) are reduced to energy-rich carbohydrate (i.e. sugar-glucose) using the light energy that is absorbed and converted into chemical energy by chlorophyll and some other photosynthetic pigments. The process of photosynthesis in green plants can be summarized as:
6CO2 + 12H2O + Light Chlorophyll⎯⎯⎯⎯⎯→ C6 H12 O6 + 6O2 + 6H2O
This is almost exactly opposite to the overall equation of aerobic respiration
C6 H12 O6 + 6O2 ⎯→ 6CO2 + 6H2O + energy
Photosynthesis uses the products of respiration and respiration uses the products of photosynthesis. There is another important difference between the two processes: Photosynthesis occurs only during day time, whereas respiration goes on day and night.
During darkness in photosynthesis
During darkness leaves (and other actively metabolizing cells) respire and utilize oxygen and release carbon dioxide
At dawn and dusk in photosynthesis
At dawn and dusk, when light intensity is low, the rate of photosynthesis and respiration may, for a short time, equal one another. Thus the oxygen released from photosynthesis is just the amount required for cellular respiration. Also, the carbon dioxide released by respiration just equals the quantity required by photosynthesizing cells. At this moment there is no net gas exchange between the leaves and the atmosphere. This is termed as compensation point.
During day time in photosynthesis
As the light intensity increases, so does the rate of photosynthesis and hence the requirement for more carbon dioxide increases which respiration alone cannot supply. Similarly, the oxygen produced during photosynthesis is more than the need of the respiring cells, so the result is the net release of oxygen coupled with the uptake of carbon dioxide.
Neil’s Hypothesis
(Water and Photosynthesis)
Oxygen released during photosynthesis comes from water, and is an important source of atmospheric oxygen which most organisms need for aerobic respiration and thus for obtaining energy to live.
Neil’s Hypothesis
(Van Neil’s Hypothesis)
In 1930s, Van Neil’s hypothesized that plants split water as a source of hydrogen, releasing oxygen as a by-product. Neil’s hypothesis was based on his investigations on photosynthesis in bacteria that make carbohydrate from carbon dioxide, but do not release oxygen.
Neil’s Hypothesis
(Experimental verification)
Neil’s hypothesis that source of oxygen released during photosynthesis is water and not carbon dioxide, was later confirmed by scientists during 1940s when first use of an isotopic tracer (O18) in biological research was made. Water and carbon dioxide containing heavy-oxygen isotope O18 were prepared in the laboratory. Experimental green plants in one group were supplied with H2O containing O18and with CO2 containing common oxygen O16. Plants in the second group were supplied with H2O containing common oxygen O16but with CO2 containing O18.
It was found that plants of first group produced O18 but the plants of second group did not.
Group-1 Plants: CO2 + 2H2O18 ⎯→ CH2O + H2O + O182
Group-2 Plants: CO182 + 2H2O ⎯→ CH2O18 +H2O18 + O2
Neil’s Hypothesis
(Confirmation of Neil’s hypothesis)
Water is thus one of the raw materials of photosynthesis, other being carbon dioxide. Hydrogen produced by splitting of water reduces NADP to NADPH2 (NADPH + H+).
NADPH2 is the “reducing power” which, along with ATP also formed during ‘light reaction’, is used to reduce CO2 to form sugar during ‘dark reaction’.
Chloroplasts − The Sites of Photosynthesis in Plants
All green parts of a plant have chloroplasts, but the leaves are the major sites of photosynthesis in most plants.
Number of Chloroplast
Chloroplasts are present in very large number, about half a million per square
millimeter of leaf surface. Chloroplasts are present mainly in the cells of mesophyll
tissue inside the leaf. Each mesophyll cell has about 20-100 chloroplasts.
Structure of chloroplast
Chloroplast has a double membrane envelope that encloses dense fluid-filled
region, the stroma which contains most of the enzymes required to produce
carbohydrate molecules. Another system of membranes is suspended in the stroma.
Thylakoid and Grana
These membranes form an elaborate interconnected set of flat, disc like sacs
called thylakoids. The thylakoid membrane encloses a fluid-filled ‘thylakoid interior-
space’ or lumen, which is separated from the stroma by thylakoid membrane. In some
places, thylakoid sacs are stacked in columns called grana (sing granum).
Role of Thylakoid
Chlorophyll (and other photosynthetic pigments) are found embedded in the
thylakoid membranes and impart green colour to the plant. Electron acceptors of
photosynthetic ‘Electron Transport Chain’ are also parts of these membranes. Thylakoid
membranes are thus involved in ATP synthesis by chemiosmosis.
Chlorophyll (and other pigments) absorb light energy, which is converted into chemical energy of ATP and NADPH, the products which are used to synthesize sugar in the stroma of chloroplast.
Imp points 1
- Photosynthetic prokaryotes lack
chloroplasts but they do have unstacked photosynthetic membranes which work like thylakoids.
2.
Photosynthetic Pigments
Pigments
Light can work in chloroplasts only if it is absorbed. Pigments are the substances that absorb visible light (380-750) nm in wave length). Different pigments absorb light of different wave lengths (colours), and the wave lengths that are absorbed disappear.
Spectrophotometer
An instrument called Spectrophotometer is used to measure relative abilities of different pigments to absorb different wavelengths of light.
Absorption spectrum:
A graph plotting absorption of light of different wave lengths by a pigment is called absorption spectrum of the pigment.
Kinds of Photosynthetic Pigments
Thylakoid membranes contain several kinds of pigments, but chlorophylls are the main photosynthetic pigments.
Other, accessory photosynthetic pigments present in the chloroplasts include yellow and red to orange carotenoids; carotenes are mostly red to orange and xanthophylls are yellow to orange. These broaden the absorption and utilization of light energy.
Chlorophylls
There are known many different kinds of chlorophylls. Chlorophyll a, b, c and d
are found in eukaryotic photosynthetic plants are algae, while the other are found in photosynthetic bacteria and are known as bacteriochlorophylls.
Absorption range
Chlorophylls absorb mainly violet-blue and orange-red wave lengths. Green,
yellow and indigo wave lengths are least absorbed by chlorophylls and are transmitted or reflected, although the yellows are often masked by darker green colour. Hence plants appear green, unless masked by other pigments
Structure of chlorophyll
A chlorophyll molecule has two
main parts.
Porphyrin head
One flat, square, light absorbing
hydrophilic head and the other long,
anchoring, hydrophobic hydrocarbon tail.
The head is complex porphyrin ring which is made up of 4 joined smaller pyrrole rings composed of carbon and nitrogen atoms.
An atom of magnesium is present in the center of porphyrin ring and is coordinated with the nitrogen of each pyrrole ring That is why magnesium deficiency
causes yellowing in plants.
Haem portion of haemoglobin is
also a porphyrin ring but containing an iron atom instead of magnesium atom in the center.
Phytol tail:
Long hydrocarbon tail which attached to one of the pyrrole rings is
phytol (C20 H39). The chlorophyll molecule is embedded in the hydrophobic core of thylakoid membrane by this tail
Difference between chlorophyll a and b
Chlorophyll a and chlorophyll b differ from each other in only one of the
functional groups bonded to the porphyrin; the methyl group (-CH3) in chlorophyll a is replaced by a terminal carbonyl group (-CHO) in chlorophyll b.
The molecular formulae for chlorophyll a and b are:
Chlorophyll a : C55 H72 O5 N4 Mg
Chlorophyll b : C55 H70 O6 N4 Mg
Due to this slight difference in their structure, the two chlorophylls show slightly different absorption spectra and hence different colours. Some wave lengths not absorbed by chlorophyll a are very effectively absorbed by chlorophyll b and vice-versa.
Most abundant and important
Of all the chlorophylls, chlorophyll a is the most abundant and the most
important photosynthetic pigment as it takes part directly in the light-dependent reactions which convert solar energy into chemical energy. It is found in all
photosynthetic organisms except photosynthetic bacteria. Chlorophyll a itself exists in several forms differing slightly in their red absorbing peaks e.g. at 670, 680, 690, 700 nm.
Chlorophyll b is found along with chlorophyll a in all green plants (embryophytes) and green algae.
Chlorophylls are insoluble in water but soluble in organic solvents, such as carbon tetrachloride, alcohol etc.
Carotenoids-accessory pigments
Carotenoids are yellow and red to orange pigments that absorb strongly the
blue-violet range, different wave lengths than the chlorophyll absorbs. So they broaden
the spectrum of light that provides energy for photosynthesis.
Role of Accessory Pigments
These and chlorophyll b are called accessory pigments because they absorb light
and transfer the energy to chlorophyll a, which then initiates the light reactions. It is
generally believed that the order of transfer of energy is:
Carotenoids ⎯→ Chlorophyll b ⎯→ Chlorophyll a
Some carotenoids protect chlorophyll from intense light by absorbing and
dissipating excessive light energy, rather than transferring energy to chlorophyll. (Similar
carotenoids may be protecting humans eye).
Light − The Driving Energy
Light is a form of energy called electromagnetic energy or radiations. Light
behaves as waves as well as sort of particles called photons. The radiations most
important to life are the visible light that ranges from about 380 to 750 nm in wave
length.
It is the sunlight energy that is absorbed by chlorophyll, converted into chemical energy, and drives the photosynthetic process. Not all the light falling on the leaves is absorbed. Only about one percent (1%) of the light falling on the leaf surface is absorbed, the rest is reflected or transmitted.Absorption spectrum
Absorption spectrum for chlorophylls (Fig. 11.4) indicates that absorption is
maximum in blue and red parts of the spectrum, two absorption peaks being at around
430 nm and 670 nm respectively. Absorption peaks of carotenoids are different from
those of chlorophylls.
Action spectrum
Different wavelengths are not only differently absorbed by photosynthetic
pigments but are also differently effective in photosynthesis.
Graph showing relative effectiveness
of different wavelengths (colours) of light in
driving photosynthesis is called action
spectrum of photosynthesis.
First action spectrum:
The first action spectrum was during photosynthesis.
a. Absorption spectrum of chlorophyll and carotenoids.
b. Action spectrum for photosynthesis.
obtained by German biologist, T.W.Engelmann in 1883. He worked on Spirogyra.
Action spectrum can be obtained by illuminating plant with light of different wavelengths (or colours) and then estimating relative CO2 consumption or oxygen release.
during photosynthesis.
a. Absorption spectrum of chlorophyll and carotenoids.
b. Action spectrum for photosynthesis.
As is evident from above figure 11.4, action spectrum of photosynthesis
corresponds to absorption spectrum of chlorophyll. The same two peaks and the valleyare obtained for absorption of light as well as for CO2 consumption. This also shows that
chlorophyll is the photosynthetic pigment.
Difference Between Absorption and Action Spectrum
However, the action spectrum of photosynthesis does not parallel the
absorption spectrum of chlorophyll exactly. Compared to the peaks in absorption
spectrum, the peaks in action spectrum are broader, and the valley is narrower and not
as deep.
(Photosynthesis in the most absorbed range is more than the absorption itself.
Likewise, photosynthesis in 500-600 nm (including green light) is more than the
absorption of green light by the chlorophyll).
Accessory Pigments
This difference occurs because the accessory pigments, the carotenoids, absorb
light in this zone and pass on some of the absorbed light to chlorophylls which then
convert light energy to chemical energy.
When equal intensities of light are given there is more photosynthesis in red
than in blue part of spectrum.
Role of Carbon Dioxide:
A Photosynthetic Reactant:
Sugar is formed during light − independent reactions of photosynthesis by the
reduction of CO2, using ATP and NADPH, the products of light − dependent reactions.
Obviously photosynthesis does not occur in the absence of CO2.
CO2 from water or air
About 10 percent of total photosynthesis is carried out by terrestrial plants, the
rest occurs in oceans, lakes and ponds. Aquatic photosynthetic organisms use dissolved
CO2, bicarbonates and soluble carbonates that are present in water as carbon source.
Air contains about 0.03 − 0.04 percent CO2. Photosynthesis occurring on land utilizes
this atmospheric CO2.
Stomata
Carbon dioxide enters the leaves through stomata and gets dissolved in the
water absorbed by the cell walls of mesophyll cells. Stomata are found in a large number in a leaf; their number being proportional to the amount of gas diffusing into the leaf. Stomata cover only 1 − 2 percent of the leaf surface but they allow proportionality much more gas to diffuse.
Role of Carbon Dioxide:
A Photosynthetic Reactant:
Sugar is formed during light − independent reactions of photosynthesis by the
reduction of CO2, using ATP and NADPH, the products of light − dependent reactions.
Obviously photosynthesis does not occur in the absence of CO2.
CO2 from water or air
About 10 percent of total photosynthesis is carried out by terrestrial plants, the
rest occurs in oceans, lakes and ponds. Aquatic photosynthetic organisms use dissolved
CO2, bicarbonates and soluble carbonates that are present in water as carbon source.
Air contains about 0.03 − 0.04 percent CO2. Photosynthesis occurring on land utilizes
this atmospheric CO2.
Stomata
Carbon dioxide enters the leaves through stomata and gets dissolved in the
water absorbed by the cell walls of mesophyll cells. Stomata are found in a large number in a leaf; their number being proportional to the amount of gas diffusing into the leaf. Stomata cover only 1 − 2 percent of the leaf surface but they allow proportionality much more gas to diffuse.
Reactions of Photosynthesis
Redox Process
Photosynthesis is a ‘redox proc
Redox Process
Photosynthesis is a ‘redox process’ that can be represented by the following
simplified summary equation:
Water Glucose
Oxidation
Carbon Oxygen
Dioxide
Reduction
+ + +
6 CO2 12 H O2 + light energy C H O 6 12 6 6O2 6H O2
Chlorophyll
However, it is not a simple, single step process, but is a complex one that is
completed by a series of simple steps or reactions. These reactions of photosynthesis
consist of two parts:
The light-dependent reactions (light reactions) which use light directly and
The light-independent reactions (dark reactions) which do not use light directly.Light reactions
Light dependent reactions constitute that phase of photosynthesis during which
light energy is absorbed by chlorophyll and other photosynthetic pigment molecules and
converted into chemical energy. As a result of this energy conversion, reducing and
assimilating power in the form of NADPH2 (NADPH + H+
) and ATP, are formed, both
temporarily storing energy to be carried alongwith H to the light independent reactions.
Dark reactions
NADPH2 provides energized
electron (and H+
), while ATP provides chemical energy for the synthesis of sugar by
reducing CO2, using reducing power and chemical energy of NADPH2 and ATP
respectively, produced by light reactions. The energy is thus stored in the molecules of
sugar. This phase of photosynthesis is also called dark reactions because these reactions
do not use light directly an can take place equally well both in light and dark provided
NADPH2 and ATP of light reactions are available.
An overview of photosynthesis: Light − Depend Reactions (Energy − conversion) and Light −
Independence Reactions (Energy − conservation)
Light Dependent Reactions
(Energy-Conversion Phase; Formation of ATP and NADPH)
Photo System
As has been described previously, sunlight energy which is absorbed by
photosynthetic pigments drives the process of photosynthesis. Photosynthetic pigmentsare organized into clusters, called photosystems, for efficient absorption and utilization
of solar energy thylakoid membranes.
are organized into clusters, called photosystems, for efficient absorption and utilization
of solar energy thylakoid membranes.
Light harvesting photosystem. Energy of light (photon) absorbed by photosynthetic pigment molecules is
transferred from molecule to molecule, and finally reaches the reaction center where actual energy
conversion begins.
Each photosystem consists of a light-gathering ‘antenna complex’ and a ‘reaction
centre’.
Antenna Complex
The antenna complex has many molecules of chlorophyll a, chlorophyll b and
carotenoids, most of them channeling the energy to reaction center.
Reaction Center
Reaction center has one or more molecules of chlorophyll a along with a primary
electron acceptor, and associated electron carriers of ‘electron transport system’.
Chlorophyll a molecules of reaction center and associated proteins are closely linked to
the nearby electron transport system. Electron transport system plays role in generation of ATP by chemiosmosis. Light energy absorbed by the pigment molecules of antenna complex is transferred ultimately to the reaction center. There the light energy isconverted into chemical energy.
Types of Photo System
There are two photosystems, photosystem I (PS I) and photosystem II (PS II).
These are named so in order of their discovery.
Photo system I
Photo-system I has chlorophyll a molecule which absorbs maximum light of 700
nm and is called P700, Photo-system II whereas reaction center of photo-system II has
P680, the form of chlorophyll a which absorbs best the light of 680 nm. A specialized
molecule called, primary electron acceptor is also associated nearby each reaction
centre. This acceptor traps the high-energy electrons from the reaction center and then
passes them on to the series of electron carriers. During this energy is used to generate
ATP by chemiosmosis.
Non-Cyclic Electron Flow
In predominant type of electron transport called non-cyclic electron flow, the
electrons pass through the two photo systems. In less common type of path called cyclic
electron flow only photo system I is involved. Formation of ATP during non-cyclic
electron flow is called non-cyclic phosphorylation while that during cyclic electron flow
is called cyclic phosphorylation.