Phototrophs convert the energy of the sun into chemical energy, which feeds

chemoorganotrophs. Chemoorganotrophs recycle the wastes of other organisms and play

important roles in industry; chemolithotrophs oxidize inorganic molecules and in the process
contribute to biogeochemical cycles such as the iron and sulfur cycles.

Chemoorganotrophs oxidize an organic energy source and conserve the energy released in
the form of ATP. The electrons released are accepted by a variety of electron acceptors, and
whether the acceptor is exogenous (that is, externally supplied) or endogenous (internally
supplied) defines the energy conserving process used by the organism. When the electron
acceptor is exogenous, the metabolic process is called respiration and may be divided into
two different types. In aerobic respiration, the final electron acceptor is oxygen, whereas the
terminal acceptor in anaerobic respiration is a different exogenous acceptor such as NO3-,
SO42-, CO2, Fe3-, and SeO42-. Organic acceptors such as fumarate and humic acids also may be
used. Respiration involves the activity of an electron transport chain. As electrons pass
through the chain to the final electron acceptor, a type of potential energy called the proton
motive force (PMF) is generated and used to synthesize ATP from ADP and Pi. In contrast,
fermentation uses an endogenous electron acceptor and does not involve an electron
transport chain or the generation of PMF. The endogenous electron acceptor is usually an
intermediate (e.g., pyruvate) of the catabolic pathway used to degrade and oxidize the
organic energy source. During fermentation, ATP is synthesized only by substrate level
phosphorylation, a process in which a phosphate group is transferred to ADP from a highenergy molecule (e.g., phosphoenolpyruvate) generated by catabolism of the energy source.

I. AEROBIC RESPIRATION:
The process of aerobic respiration is represented in the following summary form:
C6H12O6 + 6O2 + 38 ADP + 38 Pi = 6CO2 + 6H2O + 38 ATP
A. The breakdown of glucose to pyruvate
Microorganisms employ several metabolic pathways to catabolize glucose and other sugars. The
way in which microorganisms breakdown sugar to pyruvate are classified into 3 pathways: (1)

the Embden-Meyerhof pathway, (2) the pentose phosphate pathway, and (3) the
Entner-Doudoroff pathway. Collectively these pathways are called glycolytic pathways or
glycolysis.

1. The Embden-Meyerhof Pathway

The Embden-Meyerhof pathway is undoubtedly the most common pathway for glucose
degradation to pyruvate in stage two of aerobic respiration. It is found in all major groups of
microorganisms and functions in the presence or absence of O2. As noted earlier, it is also an
important amphibolic pathway and provides several precursor metabolites. The EmbdenMeyerhof pathway occurs in the cytoplasm of prokaryotes and eukaryotes.

The pathway as a whole may be divided into two parts. In the initial six-carbon
phase, energy is consumed as glucose is phosphorylated twice, and is converted to fructose
1,6-bisphosphate. This preliminary phase consumes two ATP molecules for each glucose

molecule and adds phosphates to each end of the sugar. In essence, the organism invests
some of its ATP so that more can be made later in the pathway. The three-carbon, energyconserving phase begins when the enzyme fructose 1,6-bisphosphate aldolase catalyzes the
cleavage of fructose 1,6-bisphosphate into two halves, each with a phosphate group. One of
the products, dihydroxyacetone phosphate, is immediately converted to
glyceraldehyde 3-phosphate. This yields two molecules of glyceraldhyde 3-phosphate,
which are then converted to pyruvate in a five-step process. Because dihydroxyacetone
phosphate can be easily changed to glyceraldehyde 3-phosphate, both halves of fructose 1,6bisphosphate are used in the three-carbon phase. First, glyceraldehyde 3-phosphate is oxidized
with NAD+ as the electron acceptor (to form NADH), and a phosphate (Pi) is simultaneously
incorporated to give a high energy molecule called 1,3-bisphosphoglycerate. The high-energy
phosphate on carbon one is subsequently donated to ADP to produce ATP. This synthesis of
ATP is called substrate-level phosphorylation because ADP phosphorylation is coupled with
the exergonic breakdown of a high-energy bond.
A somewhat similar process generates a second ATP by substrate-level phosphorylation.
The phosphate group on 3- phosphoglycerate shifts to carbon two, and 2-phosphoglycerate is
dehydrated to form a second high-energy molecule, phosphoenolpyruvate. This molecule
donates its phosphate to ADP forming a second ATP and pyruvate, the final product of the
pathway.
The Embden-Meyerhof pathway degrades one glucose molecule to two pyruvate
molecules. ATP and NADH are also produced. The yields of ATP and NADH may be calculated
by considering the two phases separately. In the six-carbon phase, two ATPs are used to form
fructose 1,6-bisphosphate. For each glyceraldehyde 3-phosphate transformed into pyruvate,
one NADH and two ATP are formed. Because two glyceraldehyde 3-phosphates arise from a
single glucose (one by way of dihydroxyacetone phosphate), the three carbon phase generates
four ATPs and two NADHs per glucose. Subtraction of the ATP used in the six-carbon phase
from that produced by substrate-level phosphorylation in the three-carbon phase gives a net
yield of two ATPs per glucose. Thus the catabolism of glucose to pyruvate can be represented
by this simple equation.
Glucose + 2 ADP + 2 Pi + 2 NAD+ = 2 pyruvate + 2 ATP + 2 NADH + 2 H+

2. The Pentose Phosphate Pathway

A second pathway, the pentose phosphate or hexose monophosphate pathway, may be
used at the same time as either the Embden-Meyerhof or the Entner-Doudoroff pathways. It
can operate either aerobically or anaerobically and is important in both biosynthesis and
catabolism. The pentose phosphate pathway begins with the oxidation of glucose 6phosphate to 6-phosphogluconate followed by the oxidation of 6-phosphogluconate to the
pentose sugar ribulose 5-phosphate and CO2. NADPH is produced during these oxidations.
Ribulose 5-phosphate is then converted to a mixture of three- through seven-carbon sugar
phosphates. Two enzymes play a central role in these transformations: (1) transketolase
catalyzes the transfer of two-carbon ketol groups, and (2) transaldolase transfers a threecarbon group from sedoheptulose 7-phosphate to glyceraldehyde 3-phosphate. The overall
result is that three glucose 6-phosphates are converted to two fructose 6-phosphates,
glyceraldehyde 3-phosphate, and three CO2 molecules, as shown in this equation.

3 glucose 6-phosphate + 6NADP+ + 3H2O 2 fructose 6-phosphate + glyceraldehyde 3phosphate + 3CO2 + 6NADPH + 6H+

These intermediates are used in two ways. The fructose 6-phosphate can be changed
back to glucose 6-phosphate while glyceraldehyde 3-phosphate is converted to pyruvate by
enzymes of the Embden-Meyerhof pathway. Alternatively two glyceraldehyde 3- phosphates
may combine to form fructose 1,6-bisphosphate, which is eventually converted back into
glucose 6-phosphate. This results in the complete degradation of glucose 6-phosphate to CO2
and the production of a great deal of NADPH.
Glucose 6-phosphate + 12NADP++ 7H2O 6CO2 + 12NADPH + 12H+ + Pi
The pentose phosphate pathway is a good example of an amphibolic pathway as it has
several catabolic and anabolic functions that are summarized as follows:

1. NADPH from the pentose phosphate pathway serves as a source of electrons for the
reduction of molecules during biosynthesis.
2. The pathway produces two important precursor metabolites: erythrose 4-phosphate, which is
used to synthesize aromatic amino acids and vitamin B6 (pyridoxal) and ribose 5-phosphate,
which is a major component of nucleic acids. Note that when a microorganism is growing on a
pentose carbon source, the pathway can function biosynthetically to supply hexose sugars (e.g.,
glucose needed for peptidoglycan synthesis).
3. Intermediates in the pentose phosphate pathway may be used to produce ATP.
Glyceraldehyde 3-phosphate from the pathway can enter the three-carbon phase of the
Embden-Meyerhof pathway and be converted to pyruvate, as ATP is produced by substratelevel phosphorylation. Pyruvate may be oxidized in the tricarboxylic acid cycle to provide more
energy.
Although the pentose phosphate pathway may be a source of energy in many
microorganisms, it is more often of greater importance in biosynthesis.
3. The Entner-Doudoroff Pathway

The Entner-Doudoroff pathway is used by soil microbes, such as Pseudomonas,

Rhizobium, Azotobacter, and Agrobacterium, and a few other gram-negative bacteria. Very few
gram-positive bacteria have this pathway, with the intestinal bacterium Enterococcus faecalis

being a rare exception. The Entner-Doudoroff pathway begins with the same reactions as the
pentose phosphate pathway: the formation of glucose 6-phosphate, which is then converted to
6-phosphogluconate. Instead of being further oxidized, 6-phosphogluconate is dehydrated to
form 2-keto-3-deoxy-6-phosphogluconate or KDPG, the key intermediate in this pathway. KDPG
is then cleaved by KDPG aldolase to pyruvate and glyceraldehyde 3-phosphate. The
glyceraldehyde 3-phosphate is converted to pyruvate in the Embden-Meyerhof pathway. If
the Entner-Doudoroff pathway degrades glucose to pyruvate in this way, it yields one ATP, one
NADPH, and one NADH per glucose metabolized.
B. THE TRICARBOXYLIC ACID CYCLE

In the glycolytic pathways, the energy captured by the oxidation of glucose to pyruvate
is limited to no more than two ATP generated by substrate-level phosphorylation. During
aerobic respiration, the catabolic process continues by oxidizing pyruvate to three CO2. The
first step of this process employs a multi-enzyme system called the pyruvate dehydrogenase
complex. It oxidizes and cleaves pyruvate to form one CO2 and the two-carbon molecule
acetyl-coenzyme A (acetyl-CoA). Acetyl-CoA is energy-rich because a high-energy thiol links
acetic acid to coenzyme A. Acetyl-CoA then enters the tricarboxylic acid (TCA) cycle, which
is also called the citric acid cycle or the Krebs cycle.
In the first reaction acetyl-CoA is condensed with (i.e. added to) a four-carbon
intermediate, oxaloacetate, to form citrate, a molecule with six carbons. Citrate (a tertiary
alcohol) is rearranged to give isocitrate, a more readily oxidized secondary alcohol. Isocitrate is
subsequently oxidized and decarboxylated twice to yield -ketoglutarate (five carbons), and
then succinyl-CoA (four carbons), a molecule with a high-energy bond. At this point two NADH
molecules have been formed and two carbons lost from the cycle as CO2. The cycle continues
when succinyl-CoA is converted to succinate. This involves breaking the high-energy bond in
succinyl-CoA and using the energy released to form one GTP by substrate-level
phosphorylation. GTP is also a high-energy molecule, and it is functionally equivalent to ATP.
Two oxidation steps follow, yielding one FADH2 and one NADH. The last oxidation step
regenerates oxaloacetate, and as long as there is a supply of acetyl-CoA the cycle can repeat
itself. TCA cycle generates two CO2 molecules, three NADH molecules, one FADH2, and one GTP
for each acetyl-CoA molecule oxidized.
TCA cycle enzymes are widely distributed among microorganisms. In procaryotes, they
are located in the cytoplasmic matrix. In eucaryotes, they are found in the mitochondrial
matrix. The complete cycle appears to be functional in many aerobic bacteria, free-living
protists, and fungi. Even those microorganisms that lack the complete TCA cycle usually have
most of the cycle enzymes, because the TCA cycle is also a key source of carbon skeletons for
use in biosynthesis.
C. ELECTRON TRANSPORT AND OXIDATIVE PHOSPHORYLATION
In oxidizing glucose, the cell has also generated numerous molecules of NADH and
FADH2. Both of these molecules can be used to conserve energy. In fact, most of the ATP
generated during aerobic respiration comes from the oxidation of these electron carriers in the
electron transport chain.
ETC of E.coli
The NADH generated by the oxidation of organic substrates (during glycolysis and the TCA
cycle) is donated to the electron transport chain, where it is oxidized to NAD+ by the
membrane-bound NADH dehydrogenase. The electrons are then transferred to carriers
with progressively more positive reduction potentials. As electrons move through the carriers,
protons are moved across the plasma membrane to the periplasmic space (i.e., outside the cell)
rather than to an intermembrane space as seen in the mitochondria. Another significant
difference between the E. coli chain and the mitochondrial chain is that the bacterial electron
transport chain contains a different array of cytochromes. Furthermore, E. coli has evolved two
branches of the electron transport chain that operate under different aeration conditions. When
oxygen is readily available, the cytochrome bo branch is used. When oxygen levels are reduced,
the cytochrome bd branch is used because it has a higher affinity for oxygen. However, it is less

efficient than the bo branch because the bd branch moves fewer protons into the periplasmic
space.

Oxidative Phosphorylation
Oxidative phosphorylation is the process by which ATP is synthesized as the result of
electron transport driven by the oxidation of a chemical energy source. The most widely
accepted hypothesis for the mechanism of oxidative phosphorylation is the chemiosmotic
hypothesis, which was formulated by British biochemist Peter Mitchell. Chemiosmosis
proposes that electron transport between cytochrome complexes provides the energy to
pump protons (H+) across the membrane; in the case of bacterial and archaeal cells, this is

from the cytosol to the environment. The result of proton expulsion during electron transport is
the formation of a concentration gradient of protons (pH; chemical potential energy) and a
charge gradient (; electrical potential energy). Thus, the cytoplasm is more alkaline and
more negative than the periplasmic space. The combined chemical and electrical potential
differences make up the proton motive force (PMF). The PMF is used to perform work when
protons flow back across the membrane, down the concentration and charge gradients, and
into the prokaryotic cytoplasm. This flow is exergonic and is often used to phosphorylate ADP to
ATP with the help of enzyme ATP synthase.

ATP Yield during Aerobic Respiration

II.

ANAEROBIC RESPIRATION:

As we have seen, during aerobic respiration sugars and other organic molecules are
oxidized and their electrons transferred to NAD+ and FAD to generate NADH and FADH2,
respectively. These electron carriers then donate the electrons to an electron transport chain
that uses O2 as the terminal electron acceptor. However, it is also possible for other terminal
electron acceptors to be used for electron transport. Anaerobic respiration, aprocess
whereby an exogenous terminal electron acceptor other than O2 is used for electron transport,
is carried out by many bacteria and archaea. The most common terminal electron acceptors
used during anaerobic respiration are nitrate, sulfate, and CO2, but metals and a few organic
molecules can also be reduced.
Although some bacteria and archaea grow using only anaerobic respiration, many can
perform both aerobic and anaerobic respiration, depending on the availability of oxygen. One
example is Paracoccus denitrificans, a gram-negative, facultative anaerobic soil bacterium
that is extremely versatile metabolically. It can degrade a wide variety of organic compounds
and can even grow chemolithotrophically. Under anoxic conditions, P. denitrificans uses NO3- as
its electron acceptor. Two different electron transport chains are used by this bacterium, one
for aerobic respiration and the second for anaerobic respiration. Notice that during
chemoorgantrophic growth, the source of electrons in both chains is NADH. The aerobic chain
has four complexes that correspond to the mitochondrial chain. When P. denitrificans grows
without oxygen, using NO3- as the terminal electron acceptor, the electron transport chain is
more complex. Also, not as many protons are pumped across the membrane during anaerobic
growth, but nonetheless a PMF is established. The anaerobic reduction of nitrate makes it
unavailable to the cell for assimilation or uptake. Therefore this process is called dissimilatory
nitrate reduction. Nitrate reductase replaces cytochrome oxidase to catalyze the reaction:
NO3- + 2e- + 2H+ NO2- + H2O
Nitrite is quite toxic. Bacteria such as P. denitrificans avoid the toxic effects of nitrite by
reducing it to nitrogen gas, a process known as denitrification. By donating five electrons to a
nitrate molecule, NO3- is converted into a nontoxic product.
2NO3- + 10e- + 12H+ N2 + 6H2O
Denitrification is a multistep process with four enzymes participating: nitrate reductase,
nitrite reductase, nitric oxide reductase, and nitrous oxide reductase.
NO3- NO2- NO N2O N2
In addition to P. denitrificans, some members of the genera Pseudomonas and Bacillus
carry out denitrification. All three genera use denitrification as an alternative to aerobic
respiration and may be considered facultative anaerobes.
Not all microbes employ anaerobic respiration facultatively. Some are obligate
anaerobes that can carry out only anaerobic respiration. The methanogens are an example.
These archaea use CO2 or carbonate as a terminal electron acceptor. They are called
methanogens because the electron acceptor is reduced to methane. Bacteria such as
Desulfovibrio are another example. They donate eight electrons to sulfate, reducing it to sulfide
(S2- or H2S).
SO42- + 8e- + 8H+ S2- + 4H2O

Also, both methanogens and Desulfovibrio are able to function as chemolithotrophs,

using H2 as an energy source.
In anaerobic respiration, the amount of ATP produced is less than in aerobic respiration.
There are several reasons for this. First, only a portion of the citric acid cycle functions in
anaerobic respiration, so fewer reduced coenzymes are available to the electron transport
chain. Also, not all of the cytochrome complexes function during anaerobic respiration, so the
ATP yield will be less. The exact amount of ATP produced therefore will depend on the
organism and where in the respiratory pathway intermediates enter.

III.

FERMENTATION

Despite the tremendous ATP yield obtained by oxidative phosphorylation, some

chemoorganotrophic microbes do not respire because either they lack electron transport
chains or they repress the synthesis of electron transport chain components under anoxic
conditions, making anaerobic respiration impossible. Yet NADH produced by the EmbdenMeyerhof pathway reactions during glycolysis must still be oxidized back to NAD+. If NAD+ is
not regenerated, the oxidation of glyceraldehyde 3-phosphate will cease and glycolysis will
stop. Many microorganisms solve this problem by slowing or stopping pyruvate
dehydrogenase activity and using pyruvate or one of its derivatives as an electron acceptor
for the reoxidation ofNADH in a fermentation process. There are many kinds of
fermentations, and they often are characteristic of particular microbial groups.
Three unifying themes should be kept in mind when microbial fermentations are
examined: (1) NADH is oxidized to NAD+, (2) the electron acceptor is often either pyruvate
or a pyruvate derivative, and (3) oxidative phosphorylation cannot operate, reducing theATP
yield per glucose significantly. In fermentation, the substrate is only partially oxidized,ATP is
formed exclusively by substrate-level phosphorylation, and oxygen is not needed.
Many fungi, protists, and some bacteria ferment sugars to ethanol and CO2 in a process
called alcoholic fermentation. Pyruvate is decarboxylated to acetaldehyde, which is then
reduced to ethanol by alcohol dehydrogenase with NADH as the electron donor. Lactic acid
fermentation, the reduction of pyruvate to lactate, is even more common. It is present in
bacteria (lactic acid bacteria, Bacillus), protists (Chlorella and some water molds), and even
in animal skeletal muscle. Alcoholic and lactic acid fermentations are quite useful. Alcoholic
fermentation by yeasts produces alcoholic beverages; CO2 from this fermentation causes
bread to rise. Lactic acid fermentation can spoil foods, but also is used to make yogurt,
cheese, and pickles. Also, many bacteria, especially members of the family
Enterobacteriaceae, can metabolize pyruvate to formic acid and other products in a process
sometimes called the formic acid fermentation.

Photosynthesis
Photosynthesis is a process by which light energy is converted to chemical energy that is then
stored as carbohydrate or other organic compounds. In the cyanobacteria, the process takes place in
special thylakoid membranes, which contain chlorophyll or chlorophyll-like pigments. Among
eukaryotes, photosynthesis occurs in the chloroplasts of such organisms as diatoms, dinoflagellates, and
green algae. In all cases, these microbes carry out oxygenic photosynthesis; that is, where oxygen gas
(O2) is a by-product of the process.
The Energy-Fixing Reactions:
In the first part of photosynthesis, light energy is converted into or fixed as chemical energy in
the form of ATP and NADPH. Thus, the process is referred to as energy-fixing reactions, or lightdependent reactions, because light energy is required for the process. Like the reactions of cellular
respiration, the energy-fixing reactions of photosynthesis are dependent on electrons and protons. The
source of these atomic particles is water. In the cyanobacteria and algae, the splitting of water not only
produces the needed atomic particles, it releases oxygen as a by-product:
2H2O

4H+ + 4e + O2

1. Light energy is absorbed by the green pigment chlorophyll a, a magnesium-containing, lipid soluble
compound. Chlorophylls and accessory pigments make up light-receiving complexes called
photosystems. The light excites pigment molecules in photosystem II, resulting in the loss of one
electron.
Photosystem I absorbs longer wavelength light (680 nm) and funnels the energy to a
special 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 special chlorophyll pair P680.

2. These electrons are replaced from the splitting of water. Excited electrons are immediately accepted
by the first of a series of electron carriers. The electrons are passed along the membrane carriers and
cytochrome complexes, and eventually the electrons are taken up by other chlorophyll pigments that
form photosystem I. As the electrons move between cytochromes, energy is made available for
proton pumping across the thylakoid membrane of the cyanobacterium, followed by
chemiosmosis. As described for oxidative phosphorylation, ATP is formed when protons pass back
across the membrane and release their energy. Because light was involved in the formation of ATP,
this process is called photophosporylation.
3. The electrons in photosystem I again are excited by light energy and are boosted out of the pigment
molecules to the first of another set of membrane carriers;
4. And finally to a coenzyme called nicotinamide adenine dinucleotide phosphate (NADP+). The
coenzyme functions much like NAD+ in that NADP+ receives pairs of electrons and protons from
water molecules to form NADPH.
The Carbon-Fixing Reactions:
5. In the second stage of photosynthesis, another cyclic metabolic pathway forms carbohydrates. The
process is known as the carbon-fixing reactions because the carbon in carbon dioxide is trapped (or
fixed) into carbohydrates and other organic compounds. It also is called the Calvin cycle, named
after Melvin Calvin who worked out the sequence of reactions. An enzyme, ribulose bisphosphate
carboxylase, bonds carbon dioxide to a 5-carbon organic substance called ribulose 1, 5-bisphosphate
(RuBP).

6. The resulting 6-carbon molecule then splits to form two molecules of 3-phosphoglycerate (3PG). In
the next step, the products of the energyfixing reactions, ATP and NADPH, drive the conversion of
3PG to glyceraldehyde-3-phosphate (G3P).
7. Two molecules of G3P then condense with each other to form a molecule of glucose.
Thus, the overall formula for photosynthesis may be expressed as:
6 CO2 + 6 H2O + ATP

light

C6H12O6 + 6 O2 + ADP + Pi

Notice that this reaction is the reverse of the equation for aerobic respiration. The fundamental
difference is that aerobic respiration is a catabolic, energy-yielding process, while photosynthesis is an
anabolic, energy-trapping process.
8. To finish off the cycle, most G3P molecules undergo a complex series of enzyme-catalyzed reactions
that require ATP to reform RuBP. However, some G3P exits the cycle and combines in pairs to form
glucose. The sugar then can be used for cell respiration, stored as glycogen, or used for other cellular
purposes.

In addition to the cyanobacteria, a few other groups of bacteria trap energy by photosynthesis. Two
such groups are the green bacteria and purple bacteria, so named because of the colors imparted by their
pigments. These bacterial organisms have chlorophyll-like pigments known as bacteriochlorophylls to
distinguish them from other chlorophylls. In the energy-fixing reactions, the organisms do not use water
as a source of hydrogen ions and electrons. Consequently, no oxygen is liberated and the process is
therefore called anoxygenic photosynthesis. Instead of water, a series of inorganic or organic substances,
such as hydrogen sulfide gas (H2S) and fatty acids are used as a source of electrons and hydrogen ions.
2H2S + CO2

light

Carbohydrates + H2O + 2S

Thus, the green and purple bacteria commonly live under anaerobic conditions in environments
such as sulfur springs and stagnant ponds.

Rhodopsin-Based Phototrophy
Oxygenic and anoxygenic photosynthesis are chlorophyll-based types of phototrophythat 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 eucaryotes 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

bacteriorhodopsin (more correctly called archaeorhodopsin). One such archaeon is the halophile
Halobacterium salinarum.
H. salinarum normally depends on aerobic respiration for the release of energy from an organic
energy source. However, under conditions of low oxygen and high light intensity, it synthesizes
bacteriorhodopsin, a deep-purple pigment that closely resembles the rhodopsin from the rods and cones of
vertebrate eyes. 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. However, that this type of
phototrophy does not involve electron transport.

The overall electron flow resulting in ATP generation in photosynthesis can be of

2 main types cyclic & Non cyclic.
CYCLIC PHOTO PHOSPHORYLATION IN BACTERIA:In the cyclic type, the high energy electrons ejected by the reaction center pigment flow through
a series of electron acceptors from a higher energy level to a gradually lower energy level &
return to the reaction center, forming there by a close circuit. The loss of energy of electrons in
this cyclic path is utilized for phosphorylation of ADP to ATP. The only product of cyclic path is
ATP. No NADH2 is produced.

CYCLIC
AND
NON
CYCLIC
PHOTO PHOSPHORYLATION
IN
CYANOBACTERIA:In non-cyclic photo phosphorylation electrons ejected by the reaction center pigment
complex & accepted by ferredoxin are used for reduction of NAD+/NADP+.

Chemoautotrophy
Chemoautotrophs can grow in a mineral medium, during carbon from CO2 & energy
from the oxidation of inorganic compounds. Some of these bacteria are capable of
growing both

Chemoorganotrophically

&

chemoautotrophically

i.e.,

they

are

facultative autotrophs, e.g. Alcaligenes eutrophus. Other chemoautotrophic bacteria are

obligate in nature e.g. Thiobacillus, Nitrosomonas.
Reaction which yield energy in chemoautotrophs are the oxidation of H2, NH4+,
NO3-, S & reduced

sulphur compounds and Fe++. All these oxidations, except H2

oxidation, couple electron transport to the cytochrome system & NAD+ reduction occurs
by energy dependent reverse electron flow.
The assimilation of CO2 in these organisms occurs through the reaction of
the calvin cycle. When grown chemoautotrophically, cells contain high levels of
the 2 enzymes of this pathway namely carboxy dismutase, phosphoribulokinase.
Depending on the oxidisable in organic substrate, the chemoautotrophic bacteria
can be distinguished into following groups: Nitrifying bacteria, sulfur oxidizing bacteria,
H2 oxidizing bacteria, Iron oxidizing bacteria & carbon monoxide bacteria.
Nitrifying bacteria:
Nitrification is a natural process carried out by the nitrifying bacteria occurring in soil &
aquatic bodies. It involves oxidation of ammonia liberated by decomposition of
nitrogenous organic matter like proteins, nucleic acids, urea etc. The oxidation takes place
in 2 steps ammonia to nitrous acid & nitrous acid to nitric acid. The acids react with
metal ions to produce the corresponding salts, nitrite & nitrate. Nitrate acts as main Nsource of plants.
The 2 step nitrification carried out by two different groups of bacteria.
I.

First

step

involving oxidation of

NH3

to nitrous

acid

is

called

nitrosification. Eg. Nitrosomonas. The members of this genus are highly

aerobic & strictly autotrophic.
The energy-yielding oxidation reaction of these bacteria can be represented as:
NH3 + O2 NO2 + 3H+ + 2e
NH3 + O2 + 2H+ + 2e NH2OH + H2O
NH2OH + H2O NO2- + 5H+ + 4e

II.

The second step of nitrification involves oxidation of nitrous acid to nitric acid
& organisms are known as nitrite oxidizing bacteria., Eg. Nitrobacter

NO2- + H2O NO3- + 2H+ + 2e

The organisms of both groups are capable of generating ATP by oxidative

phosphorylation in course of electron transport through the cytochrome system of
the respiratory chain & final electron acceptor is O2.
ATP generated in this way is utilized for CO2 fixation by Calvin Benson cycle.
The part of the ATP generated by oxidative phosphorylation is spent for driving
electrons from nitrite to NAD through a reverse electron transport.
SULFUR OXIDIZING BACTERIA:
Oxidation of elemental sulfur (S0) & various reduced sulfur compounds, like
sulfide (S2-), thio sulfate (S2O32-) etc. takes place in soil & aquatic bodies mediated by
both Eubacteria & Archae bacteria.
The best known among sulfur oxidizing eubacteria are members of genus
Thiobacillus. Some species like T. thiooxidans, T. thioparus & T. denitrificans are
obligately chemoautotrophic while T. novellus or T. intermedins are facultative.
Some eubacteria, designated as filamentous sulfur oxidizing bacteria belonging
to the genera Beggiatoa, Thiothrix are able to oxidise sulfide (H2S) to elemental sulfur (S0)
The anoxygenic sulfur purple & green bacteria like Chromatium, Chlorobium etc.
are able to oxidize sulfide to sulfur.
Thiobacillus oxidize elemental sulfur or sulfur compounds to sulfuric acid. The
reactions are represented as:

Other chemoautotrophic bacteria include

Iron - oxidizing bacteria - Ferrobacillus ferro oxidans, Hydrogen oxidizing bacteria - Hydrogen
oxidizing bacteria are Alcaligenes eutrophus, All are facultative chemoautotrophs. A number of
extremophilic archaebactria can grow as hydrogen bacteria eg. Pyrodictium (Anaerobic, obligately
chemo autotrophic).

IMPORTANCE OF CHEMOAUTOTROPHS:
Chemoautotrophs play an important role in oxidation of reduced N and S
compounds like NH3 and H2S to NO3 and SO4 respectively. Nitrates, sulphates are
the utilized forms of nutrients for higher plants.