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Bioquímica e Fisiologia Microbianas Licenciatura em Engenharia Química e Biológica 2º Ano - 1º Semestre - 2021/2022

The document discusses various microbial metabolic processes, focusing on chemolithotrophy, nitrification, and photosynthesis. It explains how chemolithotrophs use inorganic substrates for energy, the Calvin-Benson cycle for CO2 fixation, and the differences between oxygenic and anoxygenic photosynthesis. Key microbial groups involved in these processes include nitrifying bacteria, hydrogen-oxidizing microorganisms, and sulfur-oxidizing bacteria.

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23 views26 pages

Bioquímica e Fisiologia Microbianas Licenciatura em Engenharia Química e Biológica 2º Ano - 1º Semestre - 2021/2022

The document discusses various microbial metabolic processes, focusing on chemolithotrophy, nitrification, and photosynthesis. It explains how chemolithotrophs use inorganic substrates for energy, the Calvin-Benson cycle for CO2 fixation, and the differences between oxygenic and anoxygenic photosynthesis. Key microbial groups involved in these processes include nitrifying bacteria, hydrogen-oxidizing microorganisms, and sulfur-oxidizing bacteria.

Uploaded by

Ana Branca
Copyright
© © All Rights Reserved
We take content rights seriously. If you suspect this is your content, claim it here.
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Bioquímica e Fisiologia Microbianas

Licenciatura em Engenharia Química e Biológica


2º ano | 1º semestre | 2021/2022

T9
Chemoorganotrophic

So far, we have considered microbes that synthesize


ATP with the energy liberated by the oxidation of
organic substrates.
Chemolithotrophy

Certain bacteria and archaea are


chemolithotrophs. These microbes
donate electrons to their ETCs by
oxidizing inorganic molecules rather
than organic nutrientes.
Chemolithotrophy

The most common energy sources (electron donors) are hydrogen, reduced nitrogen
compounds, reduced sulfur compounds, and ferrous iron (Fe2+).

The acceptor is usually O2, but nitrate may also be used.


Chemolithotrophy

Much less energy is available from the oxidation of


inorganic molecules than from the complete oxidation of
glucose.

Inorganic substrates used are much more positive than


the reduction potentials of organic molecules such as
glucose and the NADH and FADH2 electron carriers.

To have enough ATP to grow and reproduce,


chemolithotrophs must oxidize a large quantity of
inorganic material.
Chemolithotrophy

Because they must consume a large amount of inorganic


material, chemolithotrophs have significant ecological impact.

They make important contributions to several biogeochemical


cycles, including the nitrogen, sulfur, and iron cycles.

Nitrifying bacteria
Hydrogen-oxidizing microorganisms
Sulfur-oxidizing bacteria
Nitrifying bacteria

Some bacteria and archaea use


the oxidation of nitrogenous
compounds as a source of
electrons.

They are soil and aquatic bacteria


that carry out nitrification - the
oxidation of ammonia to nitrate.

Nitrification is a two-step process:


first ammonia is oxidized to nitrite
and in the second step, the nitrite
is oxidized to nitrate. When two bacteria are involved one performs the ammonia oxidation (e.g.,
Nitrosomonas spp.) while another oxidizes nitrite to yield nitrate (e.g., Nitrobacter
spp.).

Some members of the genus Nitrospira can perform both steps, a process called
comammox, for complete ammonia oxidation.

A unique metabolic process that combines aspects of both nitrification and


denitrification is carried out by anammox bacteria (phylum Planctomycetes).
Chemolithotrophy

The ATP yields for chemolithotrophs are low; this reflects


their short ETCs.

Molecules such as ammonia, nitrite, and H2S have more


positive reduction potentials than the NAD(P)+/NAD(P)H
conjugate redox pair, therefore, they cannot directly donate
their electrons to NAD(P)+.

But many of these microbes are also autotrophs, and


autotrophs need NAD(P)H (reducing power) as well as ATP
to reduce CO2 and other molecules.

How to make the NAD(P)H they need?

Using a process called reverse electron flow.


Chemolithotrophy

Electrons derived from


oxidation of inorganic
substrates (reduced
nitrogen or sulfur
compounds) are moved
up their ETCs to reduce
NAD(P)+ to NAD(P)H.
Calvin-Benson Cycle

CO2 Fixation: Reduction and Assimilation of CO2 Carbon

The reactions of the Calvin-Benson cycle occur in the chloroplast stroma of eukaryotic autotrophs.

In cyanobacteria, some nitrifying bacteria, and thiobacilli (sulfur-oxidizing chemolithotrophs), the cycle is
associated with inclusions called carboxysomes.

These polyhedral structures contain the enzyme critical to the Calvin-Benson cycle and are the site of CO2
fixation.
Calvin-Benson Cycle

Three phases:

Carboxylation phase: RuBisCO catalyzes the


addition of CO2 to the ribulose-1,5-bisfosfato
(RuBP), forming a 6-carbon intermediate that
rapidly and spontaneously splits into two PGA.

Reduction phase: PGA is reduced to


glyceraldehyde 3-phosphate by two reactions
(ATP and NADPH are used).

Regeneration phase: RuBP is reformed, so that


the cycle can repeat. This phase produces
carbohydrates such as fructose 6-phosphate and
glucose 6-phosphate.
Hydrogen-oxidizing microorganisms

Members of several bacterial and archaeal genera can oxidize hydrogen gas using
a hydrogenase enzyme: H2 → 2H+ + 2e− .

Hydrogen is oxidized by a membrane-


bound hydrogenase causing proton
pumping via electron transfer to various
quinones and cytochromes.

In many organisms, a second


cytoplasmic hydrogenase is used to
generate reducing power in the form of
NADH, which is subsequently used to
fix carbon dioxide via the Calvin cycle.
Sulfur-oxidizing bacteria

Thiobacillus and Acidithiobacillus


spp. use sulfur (S0), hydrogen sulfide
(H2S), thiosulfate (S2O32−), and
other reduced sulfur compounds as
electron donors to their ETCs.

Electrons released by the oxidation of


sulfur-containing compounds are
donated to an ETC, thereby
generating a PMF that can be used
to make ATP by oxidative
phosphorylation.
Phototrophy

Many microbes capture the energy in light and use it to


synthesize ATP and reducing power (e.g., NADPH). When
the ATP and reducing power are used to reduce and
incorporate CO2, the process is called photosynthesis.

Although most people associate photosynthesis with plants,


at least half the photosynthesis on Earth is carried out by
microorganisms.

Indeed, Prochlorococcus, a cyanobacterium, is thought to be the most


abundant photosynthetic organism on Earth and thus a major
contributor to the functioning of the biosphere.
Photosynthesis

In the light reactions, light energy is trapped


and converted to chemical energy (ATP) and
reducing power (e.g., NADPH).

ATP synthesis is accomplished by formation


of a PMF, initiated by the absorption of light
— a process called photophosphorylation.

The dark reaction uses this ATP and


reducing power to fix CO2 and synthesize
cell constituents (Calvin Cycle).
Light Reactions in Oxygenic Photosynthesis

Photosynthetic 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.
Chlorophylls

Several chlorophylls are found


in eukaryotes; the two most
important are chlorophyll a
and chlorophyll b.

Because chlorophylls absorb


primarily in the red and blue
ranges, green light is
transmitted, and these
organisms appear green.
Accessory pigments

Because chlorophylls absorb a


narrow range of light wavelengths,
other photosynthetic pigments
assist in trapping light energy:

Carotenoids and phycobiliproteins.

Absorb light in the range not


absorbed by chlorophylls.

The light energy is transferred to


chlorophyll.
Oxygenic Photosynthesis

Chlorophyll molecules and the 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.

Light energy captured in an antenna is transferred


from accessory pigment to chlorophyll and from
chlorophyll to chlorophyll until it reaches a
reaction-center chlorophyll pair that is directly
involved in photosynthetic electron transport.

Photosystem I absorbs longer wavelength light


and funnels the energy to a reaction center
chlorophyll a pair called P700.

Photosystem II traps light at shorter wavelengths


and transfers its energy to the reaction center
chlorophyll pair P680.
Two Photosystems Working Together Generate ATP
and NADPH; Oxygen Is Released
Oxygenic Photosynthesis

The absorption of light causes the reaction-center chlorophylls to become excellent electron
donors.

When it absorbs light, the energized P680 donates electrons to a series of electron carriers that
ultimately donate the electrons to P700 of photosystem I.

The flow of electrons through the P680-associated ETC generates a PMF, which can be used
by ATP synthase to make ATP from ADP and Pi.

In the meantime, photosystem I has also absorbed light and its energized reaction-center
chlorophylls donate electrons to a short ETC that ultimately transfers electrons to NADP+,
yielding NADPH.

As P680 continues to absorb light, its electrons must be replaced H2O can be used to donate
electrons to P680, resulting in the release of oxygen. This is accomplished by a metal-
containing enzyme called the oxygen-evolving complex (OEC) which is closely associated with
photosystem II.
Light Reactions in Anoxygenic Photosynthesis

Certain bacteria carry out a second type of


photosynthesis called anoxygenic photosynthesis.

Molecules other than water are used as an electron


source and therefore O2 is not produced.

The process also differs in terms of:

(1) the pigments used,


(2) the participation of just one photosystem, and
(3) the mechanisms used to generate reducing
power.
Light Reactions in Anoxygenic Photosynthesis

Anoxygenic phototrophs have light-


absorbing pigments called
bacteriochlorophylls.

They are very sensitive to oxygen so are


found only in bacteria that perform
anoxygenic phototrophy.

Almost all anoxygenic phototrophs are


strict anaerobes.

Anoxygenic phototrophs have only a single


photosystem.
Light Reactions in Anoxygenic Photosynthesis

When the reaction center bacteriochlorophyll P870 is excited, it donates an electron to an


ETC that transfers the electron back to P870 while generating sufficient PMF to drive ATP
synthesis by ATP synthase.
External electron donors: Sulphur and hydrogen sulphite
Calvin-Benson Cycle
Oxygenic vs Anoxygenic Photosynthesis

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