SBT 321/BIO 310: PLANT PHYSIOLOGY
Lecture 5 Teaching Notes
18th March 2026 from 9:00a.m. to 11:00a.m.
ANABOLISM: THE USE OF
ENERGY IN BIOSYNTHESIS
Anabolism is the synthesis of complex molecules from simpler molecules with the
input of energy and reducing power.
During anabolism, an organism begins with simple inorganic molecules and a
carbon source and constructs ever more complex molecules until new organelles and
cells arise (figure 6.1). Anabolism is the creation of order. Because a cell is highly
ordered and immensely complex, much energy is required for biosynthesis. One of Comment [BNAM1]: Definition same as
anabolism.
the anabolic processes where energy is expended in plants is Photosynthesis.
PHOTOSYNTHESIS
All forms of life in the universe require energy for growth and maintenance. Plants
and some forms of bacteria capture light energy directly from solar radiation and
utilize it for synthesis of food materials besides producing basic raw materials from
which other cellular biomolecules are produced. The term photosynthesis describes
the process by which green plants synthesize organic compounds from inorganic raw
materials using light. Photosynthesis is the source of all biological energy, viz., food,
biological fuels, and biomass, and is also most important for availing free oxygen.
Whatever free oxygen is there in the atmosphere is the result of photosynthesis. Since
heterotrophic organisms including animals cannot use sunlight as direct source of
energy, they consume plants as the source of energy. Photosynthesis is the means
through which solar energy enters the global ecosystem, and it is the lone essential
biological process by which solar energy is transformed into metabolic form of energy
for all forms of life on earth.
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Session: 26th January to 24th Apilr 2026
Primitive life is believed to have existed in anaerobic conditions which utilized
energy stored in chemical compounds for the biosynthesis of biomolecules required
for their growth. However, hundreds of millions of years ago, with the depletion of
these compounds, photoautotrophs might have originated which utilized solar energy
to produce reduced organic compounds, which either oxidized water and released
oxygen (oxygenic photosynthesis), ferrous ions (Fe2+) to ferric ions (Fe3+) (e.g.,
purple photosynthetic bacteria), or used H2S as source of electrons (e.g., green sulphur
bacteria). In the latter case, sulphur got deposited by the organisms (anoxygenic
photosynthesis) rather than release oxygen. However, as time went by, almost 3.5 ×
109 years ago, water which was freely available is believed to have been utilized by
cyanobacteria in place of H2S. It was almost 2.7 × 109 years ago when oxygen is
presumed to have been released as a waste product which started to accumulate on
the earth surface resulting in an oxygenic environment. Accumulation of oxygen
shielded living organisms from damaging effects of UV irradiations.
The site of photosynthesis in eukaryotes (algae and higher plants) are the cells that
contain few to numerous (about 1–1000) chloroplasts which vary in size and shape.
Chloroplasts (Figures 5.1(a) and 5.1(b)) are unique double-membrane-bound
organelles that originated through an endosymbiotic association between free-living
oxygen-evolving photosynthetic bacteria which might have been incorporated into the
growing eukaryotic cells as chloroplast. Outer chloroplast membrane is relatively
freely permeable, while the inner membrane exhibits more selective permeability. The
sites of light reactions in the chloroplast are the sac-like structures, known as
chloroplast lamellae or thylakoids. The space within the chloroplasts is divided into
two compartments, viz., one enclosed within the thylakoids called lumen and the
other outside the thylakoids, which is called stroma. Stroma, the matrix around the
thylakoid, is the site where CO2 is assimilated, leading to the synthesis of sugars.
Thylakoids exist either as stacks called grana or are unstacked and are interconnected
to form stroma lamellae. Each chloroplast contains 10–100 grana. Light is captured
by various pigments which includes chlorophyll molecules as the photoreceptors for
photosynthesis. These exist as the chlorophyll-protein complexes which are involved
in harvesting light energy and transporting electrons, resulting in generation of
reductant and synthesis of ATP. In cyanobacteria, photosynthetic machinery required
for light reactions exists in plasma membrane which forms invaginations or folded
structures resembling grana of chloroplasts in eukaryotic cells.
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Session: 26th January to 24th Apilr 2026
Figure 5.1(a): Chloroplast structure. Note that the chloroplast is surrounded by a double membrane, but also
contains a third set of membranes, which enclose the thylakoids.
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Session: 26th January to 24th Apilr 2026
Figure 5.1(b): Illustration of general leaf structure and a chloroplast, which is a photosynthetic organelle, or
plastid, found in green plant cells. Image © copyright by Dave Carlson.
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Session: 26th January to 24th Apilr 2026
Photosynthesis is one of the most significant metabolic processes on Earth
because almost all our energy is ultimately derived from solar energy. Photosynthetic
organisms serve as the base of most food chains in the biosphere. One type of
photosynthesis is also responsible for replenishing our supply of O2. Although most
people associate photosynthesis with plants, over half the photosynthesis on Earth is
carried out by microorganisms (Table 5.1).
Table 5.1: Diversity of Phototrophic Organisms
Eukaryotic Organisms Prokaryotic Organisms
Plants Cyanobacteria
Multicellular green, brown, and red Green sulfur bacteria
algae Green nonsulfur bacteria
Unicellular protists (e.g., euglenoids, Halobacterium (archaeon), Purple sulfur
dinoflagellates, diatoms) bacteria, Purple nonsulfur bacteria
Photosynthesis is an oxidation-reduction process and is divided into two parts. In
the light reactions (Stage I), light energy is trapped and converted to chemical
energy and reducing power. These are then used to fix CO2 and synthesize cell
constituents in the dark reactions (Stage II).
Light
2H2O O2 + 4[H+] + 4e– (Stage I)
4e–+ 4[H+] +CO 2 → (CH2O) + H2O (Stage II)
In this section, three types of phototrophy are discussed: oxygenic photosynthesis,
anoxygenic photosynthesis, and rhodopsin-based phototrophy (Figure 5.2).
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Figure 5.2: Phototrophic Fueling Reactions. Phototrophs use light to generate a proton motive force
(PMF), which is then used to synthesize ATP by a process called photophosphorylation (photo phos). The
process requires light-absorbing pigments. When the pigments are chlorophyll or bacteriochlorophyll, the
absorption of light triggers electron flow through an electron transport chain, accompanied by the pumping
of protons across a membrane. The electron flow can be either cyclic (dashed line) or noncyclic (solid line),
depending on the organism and its needs. Rhodopsin-based phototrophy differs in that the PMF is formed
directly by the light-absorbing pigment, which is a light-driven proton pump. Many phototrophs are
autotrophs and must use much of the ATP and reducing power they make to fix CO2.
Phototrophic eukaryotes and cyanobacteria carry out oxygenic photosynthesis,
so named because oxygen is generated and released into the environment when light
energy is converted to chemical energy. Central to this process, and to all other
phototrophic processes, are light-absorbing pigments (Table 5.2).
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Session: 26th January to 24th Apilr 2026
Table 5.2: Properties of Chlorophyll-Based Photosynthetic Systems
Green Bacteria, Purple
Property Eukaryotes Cyanobacteria
Bacteria, and Heliobacteria
Photosynthetic pigment Chlorophyll a Chlorophyll a Bacteriochlorophyll
Number of
photosystems 2 2 1
Photosynthetic electron
donors H2O H2O H2, H2S, S, organic matter
O2 production pattern Oxygenic Oxygenica Anoxygenic Comment [Dr. M.BNA2]:
Primary products of ATP +
energy conversion ATP + NADPH ATP
NADPH
Carbon source CO2 CO2 Organic or CO2
aSome cyanobacteria can function anoxygenically under certain conditions. For example, Oscillatoria can use
H2S as an electron donor instead of H2O.
In oxygenic phototrophs, the most important pigments are the chlorophylls.
Chlorophylls are large planar molecules composed of four substituted pyrrole rings
with a magnesium atom coordinated to the four central nitrogen atoms (Figure 5.4).
Several chlorophylls are found in eukaryotes; the two most important are chlorophyll Figure 5.3a: The Electromagnetic Spectrum. A
a and chlorophyll b. These two molecules differ slightly in their structure and spectral portion of the spectrum is expanded at the bottom of
the figure. (Source: Willey et al., 2009).
properties. When dissolved in acetone, chlorophyll a has a light absorption peak at
665 nm; the corresponding peak for chlorophyll b is at 645 nm. In addition to
absorbing red light, chlorophylls also absorb blue light strongly (the second
absorption peak for chlorophyll a is at 430 nm). Because chlorophylls absorb
primarily in the red and blue ranges, green light is transmitted, and these organisms
appear green. A long hydrophobic tail attached to the chlorophyll ring aids in its
attachment to membranes, the site of the light reactions.
Figure 5.3b: Optimal absorption of light occurring at
different wavelengths for different pigments. "The
light-dependent reactions of photosynthesis: Figure 4,"
by Openstax College, Biology (CC BY 3.0)
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Figure 5.4: Chlorophyll Structure. The structures of chlorophyll a, chlorophyll b, and bacteriochlorophyll a.
The complete structure of chlorophyll a is given. Only one group is altered to produce chlorophyll b, and two
modifications in the ring system are required to change chlorophyll a to bacteriochlorophyll a. The side chain
(R) of bacteriochlorophyll a may be either phytyl (a 20-carbon chain also found in chlorophylls a and b) or
geranylgeranyl (a 20-carbon side chain similar to phytyl but with three more double bonds).
Other photosynthetic pigments also trap light energy. The most widespread of
these are the carotenoids, long molecules, usually yellowish in color, that possess an
extensive conjugated double bond system (Figure 5.5). β-Carotene is present in
Cyanobacteria belonging to the genus Prochloron and most photosynthetic protists;
fucoxanthin is found in algae such as diatoms, brown algae and dinoflagellates. Red
algae and cyanobacteria have photosynthetic pigments called phycobiliproteins,
consisting of a protein with a linear tetrapyrrole attached (Figure 5.4). Phycoerythrin
is a red pigment with a maximum absorption around 550 nm, and phycocyanin is
blue (maximum absorption at 620 to 640 nm).
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Session: 26th January to 24th Apilr 2026
Figure 5.5: Representative Accessory Pigments. Beta-carotene is a carotenoid found in photosynthetic
protists and plants. Note that it has a long chain of alternating double and single bonds called conjugated
double bonds. Fucoxanthin is a carotenoid accessory pigment in several divisions of algae (the dot in the
structure represents a carbon atom). Phycocyanobilin is an example of a linear tetrapyrrole that is attached to
a protein to form a phycobiliprotein. Comment [Dr. M.BNA3]: A photon is a particle
representing a quantum of light or other
electromagnetic radiation. A photon carries energy
Carotenoids and phycobiliproteins are often called accessory pigments because proportional to the radiation frequency and has zero
mass.
of their role in photosynthesis. They are important because they absorb light in the A photon is usually denoted by the symbol γ. Photons
are also symbolized as hν where h is the Planck
range not absorbed by chlorophylls (the blue-green through yellow range; about 470– constant (6.62607015×10−34 J⋅Hz−1)and the Greek
letter ν (nu) is the photon's frequency.
630 nm). This light is transferred to chlorophyll. In this way, accessory pigments make γ = hν
photosynthesis more efficient over a broader range of wavelengths. In addition, this Photons have a photoelectric effect: thus they emit
electrons when they hit materials.
allows organisms to use light not used by other phototrophs in their habitat.
Accessory pigments also protect microorganisms from intense sunlight, which could
oxidize and damage the photosynthetic apparatus.
Chlorophylls and accessory pigments are assembled in highly organized arrays
called antennas, whose purpose is to create a large surface area to trap as many
photons as possible. An antenna has about 300 chlorophyll molecules. Light energy is
captured in an antenna and transferred from chlorophyll to chlorophyll until it reaches
a reaction-center chlorophyll pair that is directly involved in photosynthetic
electron transport. In oxygenic phototrophs, there are two kinds of antennas
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associated with two different photosystems (Figure 5.6). Photosystem I absorbs
longer wavelength light (≥680 nm) and funnels the energy to a reaction center
chlorophyll a pair called P700. The term P700 signifies that this molecule most
effectively absorbs light at a wavelength of 700 nm. Photosystem II traps light at
shorter wavelengths (≤680 nm) and transfers its energy to the reaction center
chlorophyll pair P680.
Figure 5.6: The Z-Scheme of Photosynthesis. Electron flow during photosynthesis in higher plants.
Cyanobacteria and eucaryotic algae are similar in having two photosystems, although they may differ in some
details. The carriers involved in electron transport are ferredoxin (Fd) and other FeS proteins; cytochromes
b6, b563, and f; plastoquinone (PQ); copper containing plastocyanin (PC); pheophytin a (Pheo. a); possibly
chlorophyll a (A); and the unknown quinone Q, which is probably a plastoquinone. Both photosystem I (PS
I) and photosystem II (PS II) are involved in noncyclic photophosphorylation; only PS I participates in cyclic
photophosphorylation. The oxygen evolving complex (OEC) that extracts electrons from water contains
manganese ions and the substance Z, which transfers electrons to the PS II reaction center.
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When the photosystem I antenna transfers light energy to P700, P700 absorbs the
energy and is excited, and its reduction potential becomes very negative. This allows
P700 to donate its excited, high-energy electron to a specific acceptor, probably a
special chlorophyll a molecule or an iron-sulfur protein. The electron is eventually
transferred to ferredoxin and then travels in either of two directions. In the cyclic
pathway (the dashed lines in figure 5.6), the electron moves in a cyclic route through a
series of electron carriers and back to the oxidized P700. The pathway is termed cyclic
because the electron from P700 returns to P700 after traveling through the
photosynthetic ETC. PMF is formed during cyclic electron transport and used to
synthesize ATP. This process is called cyclic photophosphorylation because
electrons travel in a circle and ATP is formed. Only photosystem I participates.
Electrons also can travel in a noncyclic pathway involving both photosystems.
P700 is excited and donates electrons to ferredoxin as before. In the non-cyclic route,
however, reduced ferredoxin reduces NADP+ to NADPH (Figure 5.6). Because the
electrons contributed to NADP+ cannot be used to reduce oxidized P700,
photosystem II participation is required. It donates electrons to oxidized P700 and
generates ATP in the process. The photosystem II antenna absorbs light energy and
excites P680, which then reduces pheophytin a. Pheophytin a is chlorophyll a in which
two hydrogen atoms have replaced the central magnesium. Electrons subsequently
travel to the plastoquinone pool and down an ETC to reduce P700. Now P680 must
also be reduced if it is to accept more light energy. Figure 5.6 indicates that the
standard reduction potential of P680 is more positive than that of the ½O2/H2O
redox couple. Thus H2O can be used to donate electrons to P680 resulting in the
release of oxygen. Because electrons flow from water to NADP+ with the aid of
energy from two photosystems, ATP is synthesized by non-cyclic
photophosphorylation. It appears that one ATP and one NADPH are formed when
two electrons travel through the noncyclic pathway.
It is worth reemphasizing that although light is the source of energy for
chlorophyll-based phototrophy, the process used to make ATP is virtually the same as
seen for chemotrophs: oxidation-reduction reactions occurring in ETCs generate a
PMF that is used by ATP synthase to make ATP. Furthermore, just as is true of
mitochondrial electron transport, photosynthetic electron transport takes place within
a membrane. Chloroplast granal membranes contain both photosystems and their
antennas. Figure 5.7 shows a thylakoid membrane carrying out noncyclic
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photophosphorylation by the chemiosmotic mechanism. Protons move to the
thylakoid interior during photosynthetic electron transport and return to the stroma
when ATP is formed.
Figure 5.7: The Mechanism of Photosynthesis. An illustration of the chloroplast thylakoid membrane
showing photosynthetic ETC function and noncyclic photophosphorylation. The chain is composed of three
complexes: PS I, the cytochrome bf complex, and PS II. Two diffusible electron carriers connect the three
complexes. Plastoquinone (PQ) connects PS I with the cytochrome bf complex, and plastocyanin (PC)
connects the cytochrome bf complex with PS II. The light-driven electron flow pumps protons across the
thylakoid membrane and generates a PMF, which can then be used to make ATP. Water is the source of
electrons and the oxygen-evolving complex (OEC) produces oxygen.
It is believed that stromal lamellae possess only photosystem I and are involved in
cyclic photophosphorylation alone. In cyanobacteria, photosynthetic light reactions
are located in thylakoid membranes within the cell.
The dark reactions of oxygenic phototrophs use three ATPs and two NADPHs to
reduce one CO2 to carbohydrate (CH2O).
CO2 + 3ATP + 2NADPH + 2H+ + H2O → (CH2O) + 3ADP + 3Pi + 2NADP+
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The noncyclic system generates one NADPH and one ATP per pair of electrons;
therefore four electrons passing through the system produce two NADPHs and two
ATPs. A total of 8 quanta of light energy (4 quanta for each photosystem) is needed
to propel the four electrons from water to NADP+. Because the ratio of ATP to
NADPH required for CO2 fixation is 3:2, at least one more ATP must be supplied.
Cyclic photophosphorylation probably operates independently to generate the extra
ATP. This requires absorption of another 2 to 4 quanta. It follows that around 10 to
12 quanta of light energy are needed to reduce and incorporate one molecule of CO2
during photosynthesis.
Certain bacteria carry out a second type of photosynthesis called anoxygenic
photosynthesis. This phototrophic process derives its name from the fact that
molecules other than water are used as an electron source and therefore O2 is not
produced. The process also differs in terms of the photosynthetic pigments used, the
participation of just one photosystem, and the mechanisms used to generate reducing
power. Three groups of bacteria carry out anoxygenic photosynthesis: phototrophic
green bacteria, phototrophic purple bacteria, and heliobacteria.
Anoxygenic phototrophs have photosynthetic pigments called
bacteriochlorophylls (Figure 5.4). In some bacteria, these are located in
membranous vesicles called chlorosomes. The absorption maxima of
bacteriochlorophylls (Bchl) are at longer wavelengths than those of chlorophylls.
Bacteriochlorophylls a and b have maxima in ether at 775 and 790 nm, respectively. In
vivo maxima are about 830 to 890 nm (Bchl a) and 1,020 to 1,040 nm (Bchl b). This
shift of absorption maxima into the infrared region better adapts these bacteria to
their ecological niches.
Many differences found in anoxygenic phototrophs are because they have a single
photosystem. Because of this, they are restricted to cyclic photophosphorylation and
are unable to produce O2 from H2O. Indeed, almost all anoxygenic phototrophs are
strict anaerobes. A tentative scheme for the photosynthetic ETC of a purple
nonsulfur bacterium is given in figure 5.8. When the reaction-center
bacteriochlorophyll P870 is excited, it donates an electron to bacteriopheophytin.
Electrons then flow to quinones and through an ETC back to P870 while generating
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sufficient PMF to drive ATP synthesis by ATP synthase. Note that although both
green and purple bacteria lack two photosystems, the purple bacteria have a
photosynthetic apparatus similar to photosystem II of oxygenic phototrophs, whereas
the green sulfur bacteria have a system similar to photosystem I.
Figure 5.8: Purple Nonsulfur Bacterial Photosynthesis. The photosynthetic ETC in the purple nonsulfur
bacterium Rhodobacter sphaeroides. This scheme is incomplete and tentative. Ubiquinone (Q) is very similar to
coenzyme Q. BPh stands for bacteriopheophytin. The electron source succinate is in blue.
Anoxygenic photoautotrophs face a further problem because they also require
reducing power (NAD[P]H or reduced ferredoxin) for CO2 fixation and other
biosynthetic processes. They are able to generate reducing power in at least three
ways, depending on the bacterium. Some have hydrogenases that are used to produce
NAD(P)H directly from the oxidation of hydrogen gas. This is possible because
hydrogen gas has a more negative reduction potential than NAD+. Others, such as the
photosynthetic purple bacteria, use reverse electron flow to generate NAD(P)H
(Figure 5.8). In this mechanism, electrons are drawn off the photosynthetic ETC and
“pushed” to NAD(P)+ using PMF. Electrons from electron donors such as hydrogen
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sulfide, elemental sulfur, and organic compounds replace the electrons removed from
the ETC in this way. Phototrophic green bacteria and heliobacteria also draw off
electrons from their ETCs. However, because the reduction potential of the
component of the chain where this occurs is more negative than NAD+ and oxidized
ferredoxin, the electrons flow spontaneously to these electron acceptors. Thus these
bacteria exhibit a simple form of non-cyclic photosynthetic electron flow (Figure 5.9).
Figure 5.9: Green Sulfur Bacterial Photosynthesis. The photosynthetic ETC in the green sulfur bacterium
Chlorobium limicola. Light energy is used to make ATP by cyclic photophosphorylation and to move electrons
from thiosulfate (S2O32−) and H2S (green and blue) to NAD+. The ETC has a quinone called menaquinone
(MK).
Oxygenic and anoxygenic photosynthesis are chlorophyll-based types of
phototrophy—that is, chlorophyll or bacteriochlorophyll is the major pigment used to
absorb light and initiate the conversion of light energy to chemical energy. This type
of phototrophy is observed only in eukaryotes and bacteria; it has not been observed
in any archaea to date. However, some archaea are able to use light as a source of
energy. Instead of using chlorophyll, these microbes use a membrane protein called
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bacteriorhodopsin (more correctly called archaeorhodopsin). One such archaeon is Comment [BNAM4]: Singular of Archaea.
Archaea are single-celled prokaryotic microorganisms
the halophile Halobacterium salinarum. that includes methanogens and those of harsh
environments (such as acidic hot springs, hypersaline
lakes, and deep-sea hydrothermal vents) which obtain
H. salinarum normally depends on aerobic respiration for the release of energy energy from a variety of sources (such as carbon
dioxide, acetate, ammonia, sulphur or sunlight).
from an organic energy source. It cannot grow anaerobically by anaerobic respiration
or fermentation. However, under conditions of low oxygen and high light intensity, it
synthesizes bacteriorhodopsin, a deep-purple pigment that closely resembles the
rhodopsin found in the rods and cones of vertebrate eyes. Bacteriorhodopsin’s
chromophore is retinal, a type of carotenoid. The chromophore is covalently attached
to the pigment protein, which is embedded in the plasma membrane in such a way
that the retinal is in the center of the membrane.
Bacteriorhodopsin functions as a light-driven proton pump. When retinal absorbs
light, a proton is released and the bacteriorhodopsin undergoes a sequence of
conformation changes that translocate the proton into the periplasmic space. The
light-driven proton pumping generates a pH gradient that can be used to power the
synthesis of ATP by chemiosmosis. This phototrophic capacity is particularly useful
to Halobacterium because oxygen is not very soluble in concentrated salt solutions and
may decrease to an extremely low level in Halobacterium’s habitat. When the
surroundings become temporarily anoxic, the archaeon uses light energy to synthesize
sufficient ATP to survive until oxygen levels rise again. Note that this type of
phototrophy does not involve electron transport. It had been thought that rhodopsin-
based phototrophy is unique to Archaea. However, proton-pumping rhodopsins have
recently been discovered in some proteobacteria (proteorhodopsin) and a fungus.
Autotrophs use CO2 as their sole or principal carbon source, and the reduction
and incorporation of CO2 requires much energy. Many autotrophs obtain energy by
trapping light during the light reactions of photosynthesis, but some derive energy
from the oxidation of inorganic electron donors. Autotrophic CO2 fixation is crucial
to life on Earth because it provides the organic matter on which heterotrophs depend
on.
The most common CO2-fixation pathway used by autotrophs is the Calvin cycle,
which is also called the Calvin-Benson cycle or the Reductive Pentose Phosphate
cycle. The Calvin cycle is found in photosynthetic eukaryotes and most
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photosynthetic bacteria. It is absent in some obligatory anaerobic and microaerophilic
bacteria. Autotrophic archaea also use an alternative pathway for CO2 fixation.
The Calvin cycle pathway was elucidated by Melvin Calvin, James Bassham, and
Andrew Benson between 1946–1953 for which Melvin Calvin was awarded the Nobel
Prize for Chemistry in 1961. The cycle is also called the Reductive Pentose
Phosphate Cycle because it is essentially the reverse of the pentose phosphate
pathway. Thus many of the reactions are similar, in particular the sugar
transformations. The reactions of the Calvin cycle occur in the chloroplast stroma of
eukaryotic autotrophs. In cyanobacteria, some nitrifying bacteria, and thiobacilli
(sulfuroxidizing chemolithotrophs), the Calvin cycle is associated with inclusion
bodies called carboxysomes. These polyhedral structures contain the enzyme critical
to the Calvin cycle and may be the site of CO2 fixation.
The Calvin cycle is divided into three phases: carboxylation phase, reduction
phase, and regeneration phase (Figure 5.10). During the carboxylation phase, the
enzyme ribulose 1,5-bisphosphate carboxylase, also called ribulose bisphosphate
carboxylase/oxygenase (RuBisCO), catalyzes the addition of CO2 to the five-carbon
molecule ribulose 1,5-bisphosphate (RuBP), forming a six-carbon intermediate that
rapidly and spontaneously splits into two molecules of 3-phosphoglycerate (PGA).
PGA is then phosphorylated in a process called substrate level phosphorylation to
yield 1,3-bisphosphoglycerate. The phosphorylation is facilitated by the enzyme
Phosphoglycerate kinase and phosphate groups originate from ATP. 1,3-
bisphosphoglycerate is then reduced by NADPH to glyceraldehyde 3-phosphate
under the influence of Glyceraldehyde-3-phosphate dehydrogenase. Finally, in the
regeneration phase, RuBP is regenerated in a series of reactions involving
Transketolases, Aldolases and kinases to give ribulose 1,5-bisphosphate, so that the cycle
can repeat. This portion of the cycle is similar to the pentose phosphate pathway and
involves the transketolase and transaldolase reactions.
In addition, this final phase produces carbohydrates such as glyceraldehyde 3-
phosphate, fructose 6-phosphate, and glucose 6-phosphate, all of which are precursor
metabolites. Fructose 6-phosphate, and glucose 6-phosphate can liberate their
phosphate groups in yet another process of substrate level phosphorylation under
the influence of kinases to give fructose and glucose respectively.
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Comment [BNAM5]:
Figure 5.10: The Calvin Cycle. This overview of the cycle shows only the carboxylation and reduction
phases in detail. Three ribulose 1,5-bisphosphates are carboxylated to give six 3-phosphoglycerates in the
carboxylation phase. These are converted to six glyceraldehyde 3-phosphates, which can be converted to
dihydroxyacetone phosphate (DHAP). Five of the six trioses (glyceraldehyde phosphate and
dihydroxyacetone phosphate) are used to reform three ribulose 1,5-bisphosphates in the regeneration phase.
The remaining triose is used in biosynthesis. The numbers in parentheses at the lower right indicate this
carbon flow.
To synthesize fructose 6-phosphate or glucose 6-phosphate from CO2, the cycle
must operate six times to yield the desired hexose and reform the six RuBP molecules.
6RuBP + 6CO2 → 12PGA → 6RuBP + fructose-6-P
The incorporation of one CO2 into organic material requires three ATPs and two
NADPHs. The formation of glucose from CO2 may be summarized by the following
equation.
6CO2 + 18ATP + 12NADPH + 12H+ + 12H2O → Glucose + 18ADP + 18Pi +
12NADP+
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