Types of Microbial Energy Conservation

What is Fermentation?
Fermentation is a metabolic process that converts sugar to acids, gases or alcohol. It
occurs in yeast, bacteria and molds. Fermentation is the process in which a substance breaks
down into a simpler substance. Microorganisms like yeast and bacteria usually play a role in the
fermentation process, creating beer, wine, bread, kimchi, yogurt and other foods.

History of Fermentation

• Louis Pasteur - "First Zymurgist" 1850's - 1860's.

• Yeast - "First solid evidence" Europe.
• 1860's – Pasteur concluded that Bacteria causes milk to be sour.
• 1870's - "General used of the term Bacteria".
• 1877 – Pasteur published a book entitled as "Estudes sur la Biere".
• 1879 – Translated in English as "Studies on fermentation".
• Pasteur states that fermentation is "Life without Air".

The process of lactic acid fermentation replaces

the process of aerobic respiration so that the cell can
have a continual source of energy, even in the absence
of oxygen. However this shift is only temporary and
cells need oxygen for sustained activity.
Lactic Acid Fermentation Alcohol Fermentation

If no oxygen is available, cells can obtain energy through the process of anaerobic
respiration. A common anaerobic process is fermentation. Fermentation is not an efficient
process and results in the formation of far fewer ATP molecules than aerobic respiration.
There are two primary fermentation processes:
1. Lactic Acid Fermentation

2. Alcohol Fermentation
Lactic acid fermentation
occurs when oxygen is not available.
For example, in muscle tissues during rapid and
vigorous exercise, muscle cells may be depleted of
oxygen. They then switch from respiration to
fermentation.

The pyruvic acid formed during glycolysis

is broken down to lactic acid and energy is
released (which is used to form ATP).

Glucose → Pyruvic acid → Lactic acid + energy

C6H12O6 → 2 CH3CHOHCOOH
Lactic acid that builds up in the tissue causes a
burning, painful sensation.
Alcohol fermentation
occurs in yeasts and some bacteria.
Pyruvic acid formed during glycolysis is broken
down to produce alcohol and carbon dioxide and is
released (which is used to form ATP).
 Glucose → Pyruvic acid → alcohol + carbon dioxide + energy

C6H12O6 → 2C2H5OH → 2CO2

All 3 fates of pyruvate from glycolysis provide for the regeneration of NAD+ from NADH.
RESPIRATION (Tricarboxylic Acid Cycle)

In order for ATP to be produced through oxidative phosphorylation, electrons are

required for ATP to pass down the electron transport chain. These electrons come from electron
carriers such as NADH and FADH₂, which are produced by the Tricarboxylic Acid Cycle (TCA)
also known as the Krebs/Citric Acid cycle.The Krebs cycle is responsible for cellular respiration.
When glucose is broken down to pyruvate and then oxidized, acetyl CoA forms. The citric acid
cycle in eukaryotes takes place in the mitochondria while in prokaryotes, it takes place in the
cytoplasm. The different enzymes involved in the citric acid cycle are located either in the inner
membrane or in the matrix space of the mitochondria.
Equation: The overall reaction/ equation of the citric acid cycle is:

Acetyl CoA + 3 NAD+ + 1 FAD + 1 ADP + 1 Pi → 2 CO2 + 3 NADH + 3 H+ + 1 FADH2 + 1 ATP

Krebs cycle Enzymes

1. Citrate synthase
2. Aconitase
3. Isocitrate dehydrogenase
4. α-ketoglutarate
5. Succinyl-CoA synthetase
6. Succinate dehydrogenase
7. Fumarase
8. Malate dehydrogenase
It was named after Hans Adolf Kreb who discover it in 1937.
The TCA Cycle

The citric acid cycle, like the conversion of pyruvate to acetyl CoA, takes place in the
mitochondrial matrix. With the exception of succinate dehydrogenase, which is lodged in the
inner membrane of the mitochondrion, almost all enzymes in the citric acid cycle are soluble.
Unlike glycolysis, the citric acid cycle is a closed loop in which the molecule utilized in the first
stage is recycled in the last step. A series of redox, dehydration, hydration, and decarboxylation
events create two carbon dioxide molecules, one GTP/ATP, and reduced forms of NADH and
FADH2 in the cycle's eight phases. Because the NADH and FADH2 generated must send their
electrons to the next pathway in the system, which will use oxygen, this is termed an aerobic
process. The oxidation steps of the citric acid cycle do not occur if this transfer does not take
place. It's important to note that the citric acid cycle produces relatively little ATP and does not
utilize oxygen directly.The TCA cycle is a central pathway that provides a unifying point for
many metabolites, which feed into it at various points. It takes place over eight different steps:

Step 1: Acetyl CoA joins with oxaloacetate to form citrate (6 carbon molecule).
 The first step is a condensation step, combining the two-carbon acetyl group (from acetyl
CoA) with a four-carbon oxaloacetate molecule to form a six-carbon molecule of citrate. CoA is
bound to a sulfhydryl group (-SH) and diffuses away to eventually combine with another acetyl
group. This step is irreversible because it is highly exergonic. The rate of this reaction is
controlled by negative feedback and the amount of ATP available. If ATP levels increase, the
rate of this reaction decreases. If ATP is in short supply, the rate increases.

Step 2: Citrate is converted to isocitrate

Citrate loses one water molecule and gains another as citrate is converted into its isomer,
isocitrate.
Step 3:
Isocitrate is oxidised to alpha ketoglutarate (a five carbon molecule) which results in the release
of carbon dioxide. One NADH molecule is formed.

The enzyme responsible for catalyzing this step is isocitrate dehydrogenase. This is a rate
limiting step, as isocitrate dehydrogenase is an allosterically controlled enzyme.

Step 4: Alpha-ketoglutarate is oxidized to form a 4 carbon molecule. This binds to coenzyme A,

forming succinyl CoA. A second molecule of NADH is produced, alongside a second molecule
of carbon dioxide.

Step 5: Succinyl
CoA is then converted
to succinate (4 carbon
molecule) and one
GTP molecule
is produced.
 A phosphate group is substituted for coenzyme A, and a high- energy bond is formed. This
energy is used in substrate-level phosphorylation (during the conversion of the succinyl group to
succinate) to form either guanine triphosphate (GTP) or ATP. There are two forms of the
enzyme, called isoenzymes, for this step, depending upon the type of animal tissue in which they
are found.

Step 6: Succinate is converted into fumarate (4 carbon molecule) and a molecule of FADH₂ is
produced.

Dehydration process that converts succinate into fumarate. Two hydrogen atoms are
transferred to fad, producing fadh2. The energy contained in the electrons of these atoms is
insufficient to reduce nad+ but adequate to reduce fad.

Step 7: Fumarate is converted to malate.

The fumarate is reversibly hydrated to form L-malate in the presence of the enzyme
fumarate hydratase. As it is a reversible reaction, the formation of L-malate involves hydration,
whereas the formation of fumarate involves dehydration.
Step 8: Malate is then converted into oxaloacetate. The third molecule of NADH is also
produced.

The last step of the citric acid cycle is also an oxidation-reduction reaction where L-
malate is dehydrogenated to oxaloacetate in the presence of L-malate dehydrogenase, which is
present in the mitochondrial matrix.

Steps Reactant Product Enzymes Details

1 Acetyl CoA+ Citrate Citrate Synthase
oxaloacetate

2 Citrate Isocitrate Aconitase One water lost, one

water gained

3 Isocitrate a-Ketoglutarate Isocitrate uses one NAD+

dehydrogenase produces NADH+ CO2

4 a-Ketoglutarate Succinyl Ketoglutarate uses one NAD+

dehydrogenase produces NADH+ CO2

5 Succinyl CoA succinate Succinyl CoA uses one phosphate

dehydrogenase produces GTP

6 succinate fumarate succinate uses one FAD produces

dehydrogenase
FADH2
7 fumarate malate fumarase one water gained
8 malate oxaloacetate malate uses one NAD+
dehydrogenase produces NADH
ANAEROBIC RESPIRATION
Under anoxic conditions, electron acceptors other than oxygen support respiration in
certain prokaryotes and this is called Anaerobic Respiration. It is a metabolic reactions and
processes that takes place in the cell of organisms that use electron acceptors other that oxygen.
Anaerobic Respiration uses inorganic and organic compounds as their electron acceptor because
of the absence of oxygen. It uses inorganic compounds such as nitrate, sulfate, carbonate, ferric
iron, and other compounds. These molecules have a lower reduction potential than oxygen; thus,
less energy is formed per molecule of glucose in anaerobic versus aerobic conditions. Anaerobic
Respiration is very important means of energy generation to microbial habitats that oxygen is
limited or totally absent and like the aerobic condition, it requires electron transport to make
ATP.
Nitrate as an electron acceptor in the process of Denitrification
Denitrification is the process that converts nitrate (NO3−) to nitrogen gas (N2) and then
return it to the atmosphere. It is a type of anaerobic respiration the uses nitrate as an electron
acceptor. Many facultative anaerobes use denitrification because like oxygen, nitrate has a high
reduction potential. Facultative anaerobes are organisms that survived in aerobic or anaerobic
conditions. Denitrification is a significant process in acid rain and wastewater treatment that it
reduces eutrophication, the process wherein nutrients such as nitrates and phosphates that
promotes growth of algae and other aquatic plants are abundant. As a result, oxygen depletion
occurs that causes the aquatic animal’s death. Denitrification usually occurs in soil, sediments,
and anoxic environment, where oxygen is depleted, oxygen consumption exceeds the oxygen
supply and where enough nitrate is present. Denitrification is performed by heterotrophic
bacteria called denitrifiers.

Figure 1. Escherichia coli Figure 2. Pseudomonas species

 Figure 3. Rhizobia
Escherichia coli and Pseudomonas species are some examples of denitrifiers that reduces
nitrate to nitrite. Another example is Rhizobia, a soil bacterium that establish N2-fixing symbiosis
on legume roots. It can switch from aerobic to anaerobic respiration when there is an oxygen
shortage and use nitrate to support the respiration.

Sulfate as an electron acceptor in the process of Sulfate Reduction

Sulfate Reduction is a type of anaerobic respiration that utilizes sulfate (SO 42−) as
terminal electron acceptor. Sulfate (SO42−) is reduced to hydrogen sulfide (H 2S) in this process
and compared to aerobic respiration, it is relatively energetically process, though it is a vital
mechanism for bacteria and archaea living in oxygen-depleted, sulfate-rich environment. Sulfate
reducers can be organotrophic that uses carbon compounds as electron donors or lithotrophic,
uses hydrogen gas (H2). Organotrophic are organisms that obtain electrons for respiration from
organic compounds while lithotrophic are from inorganic compounds. Sulfate-reducing bacteria
was tracked 3.5 billion years ago and considered among the oldest forms of microorganisms
contributed to the sulfur cycle soon after life emerged on Earth. Many bacteria reduce small
amounts of sulfates in order to synthesize sulfur-containing cell components and this is known as
assimilatory sulfate reduction. By contrast, sulfate-reducing bacteria reduce sulfate in large
amounts to obtain energy and expel the resulting sulfide as waste and known as “dissimilatory
sulfate reduction. Sulfate-reducing bacteria may be utilized for cleaning up contaminated soils
and be a way to deal with acid mine waters.
 Figure 4. Desulfovibrio species that reduces
sulfate to hydrogen sulfide

Carbonate as an electron acceptor in the process of Methanogenesis

Methanogenesis is a form of anaerobic respiration that uses carbon as electron acceptor
and results in the production of methane (CH₄). Microbes capable of producing methane are
called Methanogens, and they have been identified only from the domain Archaea. The methane
production is a significant and widespread form of microbial metabolism, wherein, it is the final
step in the decomposition of biomass. Electron acceptors including oxygen, ferric iron, sulfate,
and nitrate are depleted during the decay process, while hydrogen, carbon dioxide, and light
organics formed by fermentation accumulate. Except for carbon dioxide, which is a result of
most catabolic processes, all electron acceptors become depleted at advanced phases of organic
breakdown. It isn't depleted in the same way as other potential electron acceptors are. Methane
being one of the Earth’s most important greenhouse gases, methane produced by methanogenesis
is a considerable contributor to global warming

Figure 5. Methanogens Figure 6. Methanogens in ruminants

Methanogenesis also occurs in the guts of human and other animals, especially in the ruminant’s
 gut. Their rumen has anaerobic organisms, mainly methanogens that helped them to digest
cellulose from the plants. The useful products of methanogenesis are absorbed by the gut and the
methane as the by-product was released by the ruminants mainly by belching.

Ferric iron as an electron acceptor

Ferric iron (Fe3+) reduced to Ferrous ion, is a widespread anaerobic terminal electron
acceptor. It uses autotrophic and heterotrophic organisms and the ferric iron reducing- organisms’
final enzymes are ferric iron reductase. Since these bacteria can use toxic hydrocarbons as a
carbon source, there is a significant interest in using these organisms as bioremediation agents in
ferric iron contaminated aquifers.

Figure 7. Geobacter species that reduces

ferric iron to ferrous ion Chemolithotrophic
respiration

Chemolithotrophy is the oxidation of inorganic chemicals for the generation of

energy. The process can use oxidative phosphorylation, just like aerobic and anaerobic
respiration, but now the substance being oxidized (the electron donor) is an inorganic compound.
The electrons are passed off to carriers within the electron transport chain, generating a proton
motive force that is used to generate ATP with the help of ATP synthase.An organism that
obtains its energy from the oxidation of inorganic compounds is known as chemolithotroph.
 The concept of chemolithotrophic autotrophy was first conceived by Sergei
Winogradsky, a Russian Microbiologist who contributed so much in the development of
environmental microbiology. Chemolithotrophy was discovered by Winogradsky while studying
the microorganisms involved in the oxidation of sulfur compounds.

Chemolithotrophy is found only in prokaryotes and is widely distributed among Bacteria

and Archaea. Electrons donors in Chemolithotrophs use a variety of inorganic compounds as
electron donors, with the most common substances being hydrogen gas, sulfur compounds (such
as sulfide and sulfur), nitrogen compounds (such as ammonium and nitrite), and ferrous iron.
Examples of Electron Donors
- Ammonia (NH4+) – Nitrate (NO2-)
- Nitrate (NO2-) – Nitrite (NO32-) in Nitrobacter
- Hydrogen sulfide (H2S) – Sulfur (S0) in Thiobacillus and Beggiatoa
- Sulfur (S0) – Sulfate (S02-) in Thiobacillus
- Hydrogen (H2) – Water (H20) in Alcaligene
 o Hydrogen oxidizers – these organisms oxidize hydrogen gas (H2) with the use of a
hydrogenase enzyme. Both aerobic and anaerobic hydrogen oxidizers exist, with the
aerobic organisms eventually reducing oxygen to water.
o Sulfur oxidizers – as a group these organisms are capable of oxidizing a wide variety of
reduced and partially reduced sulfur compounds such as hydrogen sulfide (H2S),
elemental sulfur (S0), thiosulfate (S2O32-), and sulfite (SO32-). Sulfate (SO42-) is
frequently a byproduct of the oxidation. Often the oxidation occurs in a stepwise fashion
with the help of the sulfite oxidase enzyme.
o Nitrogen oxidizers – the oxidation of ammonia (NH3) is performed as a two-step process
by nitrifying microbes, where one group oxidizes ammonia to nitrite (NO2-) and the
second group oxidizes the nitrite to nitrate (NO3-). The entire process is known as
nitrification and is performed by small groups of aerobic bacteria and archaea, often
found living together in soil or in water systems.
o Iron oxidizers – these organisms oxidize ferrous iron (Fe2+) to ferric iron (Fe3+). Since
Fe2+ has such a positive standard reduction potential, the bioenergetics are not extremely
favorable, even using oxygen as a final electron acceptor. The situation is made more
difficult for these organisms by the fact that Fe2+ spontaneously oxidizes to Fe3+ in the
presence of oxygen; the organisms must use it for their own purposes before that
happens.

A classic example of a sulfur-oxidizing bacterium is Beggiatoa, a microbe originally

described by Sergei Winograd sky, one of the founders of environmental microbiology.
Beggiatoa can be found in marine or freshwater environments. They can usually be found in
habitats that have high levels of hydrogen sulfide. These environments include cold seeps, sulfur
springs, sewage contaminated water, mud layers of lakes, and near deep hydrothermal vents.
Beggiatoa can also be found in the rhizosphere of swamp plants. During his research in Anton de
Bary’s laboratory of botany in 1887, Russian botanist Winograd sky found that Beggiatoa
oxidized hydrogen sulfide (H2S) as an energy source, forming intracellular sulfur droplets.
Winogradsky referred to this form of metabolism as inorgoxidation (oxidation of inorganic
compounds). The finding represented the first discovery of lithotrophy.)
 Electron acceptors in Chemolithotrophy
can occur aerobically or anaerobically. Just
as with either type of respiration, the best
electron acceptor is oxygen, to create the
biggest distance between the electron donor
and the electron acceptor. Using a
nonoxygen acceptor allows
chemolithotrophs to have greater diversity
and the ability to live in a wider variety of
environments, although they sacrifice
energy production.

Examples of Electron Donors

- Ammonia (NH4+) – Nitrate (NO2-) in Nitrosomonas
- Nitrate (NO2-) – Nitrite (NO32-) in Nitrobacter
- Hydrogen sulfide (H2S) – Sulfur (S0) in Thiobacillus and Beggiatoa
- Sulfur (S0) – Sulfate (S02-) in Thiobacillus
- Hydrogen (H2) – Water (H20) in Alcaligenes
-
Nitrogen Metabolism
Nitrogen fixation- describes the conversion of the relatively inert dinitrogen gas (N2) into
ammonia (NH3), a much more useable form of nitrogen for most life forms. It is the process
performed by diazotrophs. Nitrogen fixation is an essential process for Earth’s organisms.
Nitrogen fixation is an extremely energy and electron intensive process. Assimilation- is a
reductive process by which an inorganic form of nitrogen is reduced to organic nitrogen.
Ammonia assimilation occurs when the ammonia (NH3)/ammonium ion (NH4+) formed
during nitrogen fixation is incorporated into cellular nitrogen. Assimilative nitrate reduction
is a reduction of nitrate to cellular nitrogen,

Nitrification- is a 2-step process performed by chemolithotrophs. One group of

chemolithotrophs can perform the first part of the nitrification process, ammonia oxidation,
while a different group of chemolithotrophs can perform the nitrite oxidation that occurs in
the second part of nitrification.

Denitrification- is the reduction of NO3- to gaseous nitrogen compounds, such as N2.a type
of dissimilatory nitrate reduction. This produces large amounts of excess byproducts,
resulting in the loss of nitrogen from the local environment to the atmosphere.
 Anammox or anaerobic ammonia oxidation- is performed by marine bacteria. Ammonia is
oxidized anaerobically as the electron donor while nitrite is utilized as the electron acceptor,
Photophosphorylation

For any organism, the general process of phototrophy is going to be the same. A
photosystem antennae absorbs light and funnels the energy to a reaction center, specifically to a
special pair of chlorophyll or bacteriochlorophyll molecules. The molecules become excited,
changing to a more negative reduction potential (i.e. jumping up the electron tower).
The electrons can then be passed through an electron transport chain of carriers, such as
ferredoxin and cytochromes, allowing for the development of a proton motive force. The protons
are brought back across the plasma membrane through ATPase, generating ATP in the process.
Since the original energy from the process came from sunlight, as opposed to a chemical, the
process is called photophosphorylation.

Fig. 1. Photophosphorylation

If the electrons are returned to the special pair of chlorophyll/bacteriochlorophyll molecules

(cyclic photophosphorylation), the process can be repeated over and over again. If the electrons
are diverted elsewhere, such as for the reduction of NAD(P) (non-cyclic
photophosphorylation), then an external electron source must be used to replenish the system.
Fig. 2. Cyclic Photophosphorylation Fig. 3. Non-Cyclic Photophosphorylation

Cyclic Photophosphorylation
Cyclic photophosphorylation is part of Photosystem I (PS I). It is perhaps the earliest way
cells formed ATP. In modern plants, it occurs when so much NADPH is made that no NADP+ is
available to pick up the energized electron & H+ (NADP+ is the final electron acceptor). This
potentially could shut down PS I. Instead, the excited electrons are shunted through the cyclic
electron transport chain allowing it to keep using the excited electrons while creating ATP.

Fig. 2.1. Cyclic Photophosphorylation

Non-cyclic photophosphorylation
Non-cyclic photophosphorylation produces ATP using the energy from excited electrons
provided by photosystem II.

As electrons pass through the non-cyclic pathway, they do not return to the original
photosystem. This does not create a cycle, hence the name non-cyclic.

1. Photolysis provides H+ ions to replace those lost in the photosystems. The excited
electrons provide energy for a proton pump to actively transport additional H+ into the thylakoid.
The high concentration of H+ diffuse past ATP synthase as they pass out of the membrane to the
lower H+ concentration. The energy created as H+ passes the ATP synthase forms ATP.

Diffusion of H+ ions from high to low concentration through ATP synthase to form ATP
is called chemiosmosis.

2. The electrons provided by non-cyclic photophosphorylation are boosted a 2nd

time to a higher energy level by photosystem I to then create 2 NADPH.

Fig. 3.1. Cyclic Photophosphorylation

Chemiosmosis
As for chemiosmosis, the movement of ions is driven by an electrochemical gradient,
such as a proton gradient. Not only is chemiosmosis similar to osmosis. It is also similar to other
forms of passive transport, such as facilitated diffusion. It employs a similar principle. The ions
move downhill. Also, the molecules are transferred to the other side of the membrane with the
help of membrane proteins. Membrane proteins help the ions to move across since the
membrane is not readily permeable to ions, basically because of its bilipid feature. These
proteins in the membrane facilitate their movement by acting as a temporary shuttle or by
serving as a channel or a passageway.

Chemiosmosis uses membrane proteins to transport specific ions. Furthermore, it does

not require chemical energy (e.g. ATP) as opposed to an active transport system that does. In
chemiosmosis, the formation of an ion gradient leads to the generation of potential energy that is
sufficient to drive the process. In eukaryotes, it occurs in the mitochondria during cellular
respiration and in the chloroplasts during photosynthesis. Prokaryotes lack these organelles and
therefore chemiosmosis will occur in their cell membrane.

Fig. 4. Chemiosmosis
Comparison and Contrast of Cyclic Photophosphorylation and Non-Cyclic
Photophosphorylation
Photosystem I is involved in the cyclic photophosphorylation process. In the cyclic
photophosphorylation, P700 is known to be the active reaction centre. Electrons tend to pass in a
cyclic manner as the electrons tend to pass in a cyclic manner. ATP molecules get generated in
this process. Water is not needed in the cyclic photophosphorylation process.
NADPH does not get produced. Oxygen does not get produced as a by-product. This process is
ideal only in the case of bacteria.

On the other hand, both Photosystem I and II are involved in the non-cyclic
photophosphorylation process. In the non-cyclic photophosphorylation, P680 is known to be the
active reaction centre. Electrons tend to pass in a non–cyclic manner as the electrons from
Photosystem I am accepted by NADP and it does not return back. Both ATP and NADPH
molecules get formed. Water is needed in the process and the process of photolysis takes place as
well. NADPH gets produced in the non-cyclic photophosphorylation. Oxygen gets produced as
a by-product. This process is ideal amongst all the green plants.

Fig. 5. Diffreneces between Cyclic and Non-Cyclic Photophosphorylation

Anoxygenic Phototrophy
Anoxygenic photosynthesis is the phototrophic process where light energy is captured
and converted to ATP, without the production of oxygen. Water is therefore not used as an
electron donor.
Purple Phototrophic Bacteria
Purple phototrophic bacteria engage in anoxygenic phototrophy, indicating that they do
not generate oxygen during the process. They have a single photosystem with
bacteriochlorophyll, allowing them to use cyclic photophosphorylation as described above for
the formation of ATP. But if a purple bacterium wants to grow as a photoautotroph, it will also
need reducing power in the form of NAD(P)H. The reaction center of purple bacteria (known as
P870) has an E0’ of +0.5V. After being hit by a photon of light, the potential changes to -1.0V,
which is insufficient to reduce
NAD(P) with its reduction potential (E0’) of -0.32V.

Thus, autotrophic purple bacteria must engage in a process known as reverse electron
flow, using energy from the proton motive force to drive electrons up the electron tower.
Additionally, they must find an external electron donor to replenish the electrons now diverted to
NAD(P). Typically the electrons come from H2S or elemental sulfur, with various sulfur
byproducts produced.

Fig. 6. Anoxygenic Photosynthesis and Reduction Potential in

Purple Phototrophic Bacteria

In the presence of organic compounds, the purple bacteria often exist as

photoheterotrophs, utilizing cyclic photophosphorylation to generate ATP and getting their
organic compounds from the environment. This eliminates the need for using reverse electron
flow, an energetically unfavorable process, as well as the need for external electron donors.
Green Phototrophic Bacteria
Green phototrophic bacteria also engage in anoxygenic phototrophy, utilizing a single
photosystem with bacteriochlorophyll for cyclic photophosphorylation in the production of ATP.
However, they also use this same photosystem for generation of reducing power, by periodically
drawing off electrons to NAD+. The use of reverse electron flow is unnecessary, however, since
the initial carrier, ferredoxin (Fd) has a reduction potential (E0’) with a more negative reduction
potential than NAD(P). An external electron donor is required, typically by using H2S or
thiosulfate. Thus, the green bacteria operate as photoautotrophs, by alternating the use of their
photosystem for ATP or NAD(P)H.

Fig. 7. Anoxygenic Photosynthesis and Reduction Potential in Green Phototrophic Bacteria

Oxygenic Phototrophy
Oxygenic photosynthesis is a non-cyclic photosynthetic electron chain where the initial
electron donor is water and, as a consequence, molecular oxygen is liberated as a byproduct. The
use of water as an electron donor requires a photosynthetic apparatus with two reaction centers.

Cyanobacteria
Oxygenic phototrophy is used by cyanobacteria containing chlorophyll a, with two
distinct photosystems, each containing separate reaction centers. This allows for the generation
of both ATP and reducing power in one process, facilitating photoautotrophic growth through
the fixation of CO2. This can appropriately be referred to as photosynthesis and it is the same
process used by plants, commonly referred to as the “Z pathway.”
 The process starts when light energy decreases the reduction potential of P680
chlorophyll a molecules contained in photosystem II (PSII). The electrons are then passed
through an electron transport chain, generating ATP via a proton motive force. Electrons are then
passed to photosystem I (PSI), where they get hit by another photon of light, decreasing their
reduction potential even more. The electrons are then passed through a different electron
transport chain, eventually being passed off to NADP+ for the formation of NADPH.

Fig.8. Oxygenic Photosynthesis in Cyanobacteria

Overview of Oxygenic Photosynthesis in Cyanobacteria

The process is an example of noncyclic photophosphorylation, since the electrons are not
returned to the original photosystem. Thus, an external electron donor is required in order to
allow the process to repeat. Water, found on the right side of the redox couple O2/H2O, is
normally a poor electron donor, due to its extremely positive reduction potential. But the
reduction potential of P680 chlorophyll a is even more positive when not excited, allowing for
water to serve as an electron donor. The hydrolysis of water leads to the evolution of oxygen, a
welcome byproduct for all organisms that use aerobic respiration. It is thought that cyanobacteria
are responsible for the oxygenation of Earth, allowing for the development of aerobic respiration
as a form of metabolism.

There are some conditions under which cyanobacteria only use PSI, essentially
performing a form of anoxygenic phototrophy, despite their possession of chlorophyll a. This
occurs within the heterocysts of cyanobacteria, where oxygen inactivates the nitrogenase
enzymes. Heterocysts degrade PSII, ensuring that oxygen will not be produced as a byproduct,
while still allowing for the production of ATP with the remaining photosystem.

Rhodopsin-Based Phototrophy
One unusual form of phototrophy is used by archaea, without the use of chlorophyll or
bacteriochlorophyll. Instead these organisms use a bacteriorhodopsin (more appropriately called
an archaearhodopsin), a retinal molecule related to the one found in vertebrate eyes. When the
rhodopsin absorbs light it undergoes a conformational change, pumping a proton across the cell
membrane and leading to the development of a proton motive force, without the participation of
an electron transport chain.

Fig.9. Simple scheme for phototrophy based on bacteriorhodopsin (BR) or proteorhodopsin

(PR) and ATP synthase.

REFERENCES:

Culpepper C.J. (2006). 1911 Encyclopedia Britannica. Retrieve November 11, 2021 from.

https://doi.org/10.1108/09504120610655222

Wakim, S., & Grewal, M. (2021). Human Biology (Wakim & Grewal). Butte College. Retrieved

November 11, 2021 from https://bio.libretexts.org/@go/page/16710

Fermentation of food (2013) Retrieved November 11, 2021, from

https://todosobrecarbohidratos.wordpress.com/2013/08/07/fermentacion-de-los-alimentos/

The Citric Acid (Krebs) Cycle | Boundless Microbiology. (n.d.) Retrieved November 11, 2021

from https://courses.lumenlearning.com/boundless-microbiology/chapter/the-citric-acid-

krebscycle/

Sapkota A. (2021). Krebs cycle / Citric acid cycle / TCA Cycle with steps and diagram.

Retrieved November 11, 2021 from https://microbenotes.com/krebs-cycle/

Anaerobic Respiration | Boundless Microbiology. (n.d.). Retrieved November 11, 2021 from

https://courses.lumenlearning.com/boundless-microbiology/chapter/anaerobic-respiration/

Chemolithotrophy | Boundless Microbiology. (n.d) .Retrieved November 11, 2021 from

https://courses.lumenlearning.com/boundless-microbiology/chapter/chemolithotrophy/

Chemolithotrophy & Nitrogen Metabolism. (n.d.). Retrieved November 11, 2021 from

https://open.oregonstate.education/generalmicrobiology/chapter/chemolithotrophy-

nitrogenmetabolism/

Anoxygenic Phototrophy | Microbial Physiology. (n.d.). Retrieved November 10, 2021, from

https://courses.lumenlearning.com/boundless microbiology/chapter/phototrophy/
Photophosphorylation: Anoxygenic and Oxygenic. (2019). Retrieved November 10, 2021

from https://bio.libretexts.org/@go/page/8525

Cyclic Photophosphorylation. (n.d.). Microbial Physiology. Retrieved November 10, 2021

from

http://science.halleyhosting.com/sci/ibbio/cellenergy/photosynnotes/photosyn/cyclic.htm

Non-Cyclic Photophosphorylation. (n.d.). Microbiology. Retrieved November 10, 2021 from

http://science.halleyhosting.com/sci/ibbio/cellenergy/photosynnotes/photosyn/noncyclic.htm