Microbial nutrition and basic metabolism

E.C.S. Chan, in Handbook of Water and Wastewater Microbiology, 2003

7.2.2 Anoxygenic photosynthesis

Green and purple sulphur bacteria differ from cyanobacteria because most of them are strict anaerobes
and do not use water as an electron source. The green and purple sulphur bacteria use H2, H2S and
elemental sulphur as electron donors and possess different light-harvesting pigments called
bacteriochlorophylls. They absorb light at longer wavelengths. Bacteriochlorophyll a has an absorption
maximum at 775 nm, while bacteriochlorophyll b has a maximum at 790 nm. The Photosystem (PS) or
Reaction Centre in purple bacteria is called P870, while that in green bacteria is called P840 (denoting
the absorption wavelength maxima associated with them). Both bacterial groups exhibit cyclic electron
transport which can be used to generate ATP. They are unable to synthesize NADPH + H+ directly by
photosynthetic electron movement. The generalized photosynthetic electron flow in anoxygenic
photosynthesis is shown in Fig. 1.4. Briefly, an electron is removed from the PS by bacteriopheophytin
and passed along to ubiquinone. Electrons pass from ubiquinone through an electron transport system
(creating a proton motive force, Δp) and return to the PS restoring it to the reduced state.

Purple bacteria generate NADPH + H+ by reversed electron flow to drive electrons from organic
compounds or inorganic compounds to NADP+ and energized by the proton motive force. Alternatively,
in the presence of H2, NADPH + H+ can be produced directly as H2 has a reduction potential more
negative than NAD+. In the green sulphur bacteria, the primary acceptor of electrons from the PS is not
bacteriopheophytin but an isomer called bacteriochlorophyll 663. The subsequent electron acceptor is
not a quinone but ferredoxin. Green sulphur bacteria also exhibit a form of non-cyclic photosynthetic
electron flow in order to reduce NAD+ (not shown in Fig. 1.4). They oxidize sulphide to sulphur, with
donation of electrons to the PS, to bacteriochlorophyll 663 and then to an iron sulphurcytochrome b
complex with ferredoxin serving as the immediate donor of electrons to NAD+. The elemental sulphur
accumulates as refractile granules outside of the bacterial cells.
What Is a Phototroph?
The word phototroph gives the first clue revealing what makes these organisms
important. It means “light nourishment” in Greek. Put simply, phototrophs are
organisms that get their energy from photons, or particles of light. You probably
already know that green plants use light to make energy through photosynthesis.

However, this process isn’t restricted to plants. Many prokaryotic and eukaryotic
organisms carry out photosynthesis to make their own food, including photosynthetic
bacteria and some algae.

While photosynthesis is similar among all organisms that do it, the process of bacterial
photosynthesis is less complicated than plant photosynthesis.

What Is Bacterial Chlorophyll?

Just like green plants, phototrophic bacteria use pigments to capture photons as
energy sources for photosynthesis. For bacteria, these are bacteriochlorophylls found
in the plasma membrane (rather than in chloroplasts like plant chlorophyll pigments).

Bacteriochlorophylls exist in seven known varieties, labeled a, b, c, d, e, c s or g. Each

variant is structurally different and therefore able to absorb a specific type of light from
the spectrum, ranging from infrared radiation to red light to far red light. The type of
bacteriochlorophyll a phototrophic bacterium contains depends on its species.

Steps in Bacterial Photosynthesis

Just like plant photosynthesis, bacterial photosynthesis occurs in two stages: light
reactions and dark reactions.

In the light stage, the bacteriochlorophylls capture photons. The process of absorbing

this light energy excites the bacteriochlorophyll, triggering an avalanche of electron
transfers and ultimately producing adenosine triphosphate (ATP) and nicotinamide
adenine dinucleotide phosphate (NADPH).

In the dark stage, those ATP and NADPH molecules are used in chemical reactions
that transform carbon dioxide into organic carbon through a process called carbon
fixation.

Different types of bacteria make fuel by fixing carbon in different ways using a carbon
source such as carbon dioxide. For example, cyanobacteria use the Calvin cycle.
This mechanism uses a compound with five carbons called RuBP to catch one
molecule of carbon dioxide and form a molecule with six carbons. This splits into two
equal pieces, and one half exits the cycle as a sugar molecule.

The other half transforms into a molecule with five carbons, thanks to reactions
involving ATP and NADPH. Then, the cycle begins again. Other bacteria rely on the
reverse Krebs cycle, which is a series of chemical reactions that use electron donors
(such as hydrogen, sulfide or thiosulfate) to produce organic carbon from the inorganic
compounds carbon dioxide and water.

Why Are Phototrophs Important?

Phototrophs that use photosynthesis (called photoautotrophs) form the base of the
food chain. Other organisms that can’t perform photosynthesis get their fuel by using
photoautotrophic organisms as a food source.

Because they can’t convert light to fuel on their own, these organisms simply eat the
organisms that do and use their bodies as a source of energy. Since carbon fixing
uses carbon dioxide to produce fuel in the form of sugar molecules, phototrophs help
reduce excess carbon dioxide in the atmosphere.

Phototrophs may even be responsible for the free oxygen in the atmosphere that
enables you to breathe and thrive on Earth. This possibility – called the Great
Oxygenation Event – proposes that cyanobacteria performing photosynthesis and
releasing oxygen as a byproduct eventually produced too much oxygen to be
absorbed by iron in the environment.

This excess became part of the atmosphere and shaped evolution on the planet from
that point forward, making it possible for humans to eventually emerge.

Chemosynthesis

Photosynthesis occurs in plants and some bacteria, wherever there is sufficient sunlight – on land,
in shallow water, even inside and below clear ice. All photosynthetic organisms use solar energy to
turn carbon dioxide and water into sugar (food) and oxygen: CO 2 + 6H2O -> C6H12O6 + 6O2.

Chemosynthesis occurs in bacteria and other organisms and involves the use of energy released
by inorganic chemical reactions to produce food. All chemosynthetic organisms use energy released
by chemical reactions to make a sugar, but different species use different pathways. For example, at
hydrothermal vents, vent bacteria oxidize hydrogen sulfide, add carbon dioxide and oxygen, and
produce sugar, sulfur, and water: CO2 + 4H2S + O2 -> CH20 + 4S + 3H2O. Other bacteria make
organic matter by reducing sulfide or oxidizing methane.

Our knowledge of chemosynthetic communities is relatively new, brought to light by ocean

exploration when humans first observed a vent on the deep ocean floor in 1977 and found a thriving
community where there was no light. Since then, chemosynthetic bacterial communities have been
found in hot springs on land and on the seafloor around hydrothermal vents, cold seeps, whale
carcasses, and sunken ships. No one had ever thought to look for them, but these communities
were there all along.

New article

All living things require a way to produce energy in order to power the metabolic,
synthetic and reproductive machinery inside their cells. Ultimately, every living thing
uses the molecule ATP (adenosine triphosphate)  for this purpose.

In turn, to derive energy from molecules, those molecules, called nutrients, must be
easy to find and simple to break down. Glucose fits this description for most life on
Earth. Some organisms get glucose by digesting what they eat; others have to make it
or make other carbohydrates.

Far under the ocean's surface, where pressures are extreme and nutrients scarce,
certain communities of organisms are able to not merely survive but thrive. Not by
accident, in fact, they do so while clustering around hydrothermal vents, openings in
the sea floor that emit extreme heat and chemicals that many species cannot tolerate
(like miniature volcanoes). These chemosynthetic organisms represent both a
curiosity and a triumph of evolution in terms of how they make food.

How Organisms Get Food

Organisms can be classified as prokaryotes, the cells of which lack membrane-bound
organelles and reproduce asexually, or eukaryotes, whose cells have their DNA
enclosed in nuclei and feature a host of membrane-bound organelles in the cytoplasm.
Among those membrane-bound organelles are mitochondria and, in plants,
chloroplasts.

Mitochondria allow all eukaryotes to break down glucose aerobically to carbon dioxide,
water and energy; chloroplasts allow plants to build glucose from carbon dioxide since
they cannot ingest it.
Chemosynthesis is the derivation of carbon from carbon dioxide plus energy from
other agents, described below. Chemosynthesis is thus closely related to
photosynthesis. In fact, together, chemosynthetic organisms and photosynthetic
organisms make up the autotrophs, or the class of living things that make, rather than
ingest, their own food. These can be either prokaryotes or eukaryotes, as you'll see.

What Are Autotrophs?

Autotrophs are organisms that can produce, or synthesize, their own food as long as
a source of carbon and a source of energy is present. This minimal source of carbon
is usually in the form of carbon dioxide (CO2), a molecule that is virtually everywhere
on and above the planet.

Humans and other animals excrete it as waste. Plants and other autotrophs use it as
fuel, maintaining one of nature's more grand and definitive biochemical cycles.

Plants are the most familiar type of autotroph, but various others dot the global
biosphere, often far from human eyes. Algae, phytoplankton and certain bacteria are
autotrophs. In particular, the bacteria that can survive deep in the sea are of special
interest because of their chemosynthetic metabolism.

Chemosynthesis: Definition
Chemosynthesis is a process by which energy is derived via the microbial mediation
of certain chemical reactions. The source of energy for chemosynthesis is energy
liberated from a chemical reaction (the oxidation of an inorganic substance) rather
than energy harvested from sunlight or other light.

The carbon source remains CO 2, and oxygen (as O2) must be present to operate on
the inorganic molecule, but that inorganic molecule may be hydrogen gas (H 2),
hydrogen sulfide (H2S) or ammonia (NH3), depending on the environment in question.
Whatever carbohydrate is formed for the cell's use will have the form (CH 2O)N, as this
is true of all carbohydrates by definition.

One chemosynthesis equation depicts the conversion of carbon dioxide to

carbohydrate as hydrogen sulfide is oxidized to water and sulfur:

CO2+ O2 + 4 H2S → CH2O + 4 S + 3 H2O

Chemosynthetic Bacteria and Life Examples

Some organisms can survive in the vicinity of sea floor vents, because these emit
water with a temperature of around 5 to 100 °C (41 to 212 °F). This is not precisely
warm and welcoming, but inconsistent and sometimes violent heat is better than no
heat at all if you have the right enzymatic equipment.
Some "bacteria" in these so-called hydrothermal vent communities are
actually Archaea, prokaryotic organisms closely related to bacteria (and formerly
called archaebacteria). One example is Methanopyrus kandleri, which tolerates very
salty and very warm environments with unusual ease. This species gets energy from
hydrogen gas and releases methane (CH 4).

Script from the MIT https://ocw.mit.edu/high-school/biology/exam-prep/cellular-

energetics/photosynthesis/chemosynthesis/

So for carbon the choices are inorganic or organic.

So this would be CO2 and this might be glucose or sugars, any sugars. And then on the energy axis they can use
solar energy, as in photosynthesis, or they can use chemical energy.
And within the chemical energy sources they can be inorganic or organic like sugars, etc. And often here you have
reduced compounds such as hydrogen sulfide, ammonia, and we'll talk about these.
So these are the ways we divide up the possibilities for carbon and energy sources to be alive. All organisms also
need to have an energy currency in the cell. And you've talked about this a lot already in the biochemistry lectures so
I'm, again, just giving you the impressionist view of this. You know the details.
This is just to get you organized. And so all life uses redox reactions. And in your handouts for today there's a primer
on redox reactions just in case you want to review that.
And one of the key reactions we'll talk about today is the conversion of NADP. If you put energy in you can reduce it
to NADPH.
So that's a reduction. And the reverse you get energy out when it's oxidized. Now, we're going to be talking about
oxidation and reduction today. And then they all use ATP whichyou've talked a lot about here. And the couple here is
ADP. Put energy in.
You make ATP which is a high energy intermediate. And in converting it back to ADP that energy can be released.
And this is used in the biochemistry of the cell. So all cells have these two energy conversion processes in common.
OK, so let's look at just summarizing what we're going to go over today. This is a summary of options for life. See
also Freeman,Chapter 25. There is some discussion of this.
And we can divide life here between what we call autotrophs.
These are organisms that can make their own organic carbon.
In other words, they can convert carbon dioxide to organic carbon.
Heterotrophs are organisms that can only use organic carbon.
They rely on the guts of other organisms in order to get through life. And so now we're going to systematically go
through these processes that fall under each one of these. Oxygenic photosynthesis is the one we've been talking
about last time and in my abbreviated version of life on earth.
And this is carried out by eukaryotic organisms, plants, trees, etc., and also by prokaryotic organisms.
Those are the cyanobacteria, microscopic photosynthetic plants.
They use CO2 and sunlight. So our first variant on this theme we'll get into is a group of bacteria that do
anoxygenic photosynthesis. Oxygenic means they evolve oxygen.
These guys use solar energy but they don't evolve oxygen.
And we'll get into how that works. And then there's a group of organisms that still use CO2. And in the very similar
pathway the Calvin Cycle is photosynthesis. But they use chemical energy in order to make these intermediates to fix
CO2. OK, so let's talk about those first. And so we're going to talk about the autotrophs.
And all of them share this pathway, CO2 to C6H12. This would be glucose.
And it takes ATP to run this reaction and it also takes reduced NADPH -- -- to run this reaction. It also takes this
enzyme ribisco which you've talked about I'm sure, ribulose bisphosphate carboxylase.
And this is the enzyme that initially takes the CO2 from the atmosphere and binds it to an organic carbon.
Now, in a detailed version of this is what's called the Calvin Cycle or the Calvin/Benson Cycle. I don't know which one
your book calls it. Calvin got the Nobel Prize but Benson was the graduate student that did all the work, so you
should recognize that.
Anyway, you studied this in great deal. But an interesting factoid is that ribisco is the most abundant protein on earth.
That tells you how important this reaction is for sustaining life on earth. So notice that in order to drive this reaction,
which is the Calvin Cycle, it requires energy and reducing power. So where do they get it?
Well, there are three ways that autotrophs can get energy and reducing power to drive thisreaction. And the first is
oxygenic photosynthesis. And the second is anoxygenic. And the third is chemosynthesis.
OK, those first three there. So now we're going to go through each of these and look at how they work remembering
that all of them are generating ATP and NADPH in order to drive that. So all of the autotrophs have that in common.
Well, oxygenic photosynthesis is the one that you know well already. You've studied it in great detail in biochemistry.
So we're going to,again, give you the abbreviated version here just so you have a template to map these other ones
onto.
These are what are known as the light reactions of photosynthesis, the Z scheme taking solar energy, splitting water,
evolving oxygen and synthesizing ATP and NADPH. This is all familiar, right? Very familiar. I'm just writing it in a
cartoon version. OK, so this is the NADPH and ADPthat goes to fuel that process.
OK, so now, well, at least I can do it on that board.
Let me do it on this board. Anoxygenic -- -- is almost exactly like this process, but instead of splitting water these guys
oxidize hydrogen sulfide. So here's our ATP and NADPH.
And they use sunlight to do this.
So these are called photosynthetic bacteria. And they were around very early on the earth. Long before the earth's
atmosphere was oxygenated these were the guys that were able to use solar energy and make organic carbon but
without evolving oxygen.
Then somewhere along the line some cell evolved, had some mutations and somehow figured out that water, this
abundant source of water was a much better electron donor than hydrogen sulfide.
And once the biochemistry figured this out, you can see the simple substitution here, the whole earth started going in
a different direction. So this is an interesting example of how a small biochemical innovation can dramatically
change the whole nature of the planet. Now, these guys are still around on earth. In fact, I'm going to show you some.
I'll explain this at the end, but I have some captured in here.
See that little purple band? Those are those guys.
I've got other little tricks in here but I'll save those.
Well, you cannot really see the purple band. But you can come up later and look at it. Those are photosynthetic
bacteria.
So they're still around on the earth but they're stuck in places where there's no oxygen. So they have a rather
restricted niche on the planet now, but they're still extremely important. What did I do? Oh, here it is.
So one of the places that they can be found, and if you're interested in them a great place to go find some is out at
the Mystic Lakes in Arlington which is a permanently stratified lake so the bottom of the lake is always anaerobic.
There's never oxygen there.
In a typical lake like that you have a lot of mud on the bottom and you have a lot of hydrogen sulfide coming out of the
mud from bacterial processes that we'll talk about. And you have light here.
And so you have a gradient here of this is oxygen and this is H2S.
And these photosynthetic bacteria have to life somewhere where there's enough light to photosynthesize and enough
hydrogen sulfide to use in this part of the reaction. But they're very sensitive to oxygen so they cannot be in the
oxygenated part of the lake.
So you find them in a layer. It's called the squeeze. They have to have light so they have to be up, but they cannot
have oxygen so they have to be down. And they need hydrogen sulfide so they have to be down. So they're layered
in lakes.
OK. So what about these guys, chemosynthesis?
They don't rely on solar energy. Again, they're still driving the Calvin Cycle reducing CO2 from the air into organic
carbon, but they're not using sunlight. So what do they do? They get their energy -- -- from redox reactions. And let's
just show you an example.
Redox reactions couple to the conversion of oxygen to H2O. So oxygen is involved in thesereactions. And one
organism, for example, can take ammonia and convert it to nitrite. Another type of organism can take nitrite and
convert it to nitrate. And there are other organisms thatcan take hydrogen sulfide and convert it to sulfate.
And some can take hydrogen sulfide, oh, no, take iron, ferrous iron, Fe2+ and convert it to Fe3+. So in all of these
cases what is happening to these compounds? Are they being oxidized or reduced?
I heard an oxidized. Yes, they're being oxidized.
So these reduced compounds, relatively reduced compounds can be utilized by oxidizing them. The organism can
release the energy that's needed. ATP is generated here.
And NADPH is generated by any of these redox couples. So using this energy then the celltakes the reduced NADPH
and the ATP and it runs the Calvin Cycle, chemosynthesis. OK. Now, you may think that these are kind of
strange, weird bacteria that life in strange pockets of the earth where there's no oxygen. And who cares anyway?
They're outdated.
They dominated the earth way back in the early stages of the earth but they're not so important now. Well, that's not
true. They're incredibly important. In some ecosystems they're the total base of the entire ecosystem. But also on a
global scale, as you'll learn, you should have a feeling for this by the end of this lecture, but also when we talk about
global biogeochemical cycles you will learn that these microbes are really messengers for electrons in the
environment. Without them the redox balance of the earth would not be maintained, OK? You cannot have nothing
but oxidizing reactions or nothing but reduction reactions and have a system sustain itself. So it's these microbes that
are playing a really important role in maintaining the redox balance of the earth. OK. Now, one system that I'm going
to show you in that DVD, that will do much better justice to it than my drawings here, that's a deep-sea volcano in
case you didn't recognize it. And this is 2500 meters at the bottom of the ocean, very, very deep. And there is intense
heat. I mean just think of a volcano on the surface of the earth. Intense heat and reduced compounds are found in
the earth's mantle that are ready to erupt through this deep-sea volcano. And you have sulfate in the sea water that
percolates through here. And as it percolates in and gets draw into the volcanic stuff that's coming out of here it's
reduced to hydrogen sulfide coming out of the volcano.
But you have oxygen in the water in the deep-sea. And we'll be talking about this when we talk about ocean
circulation. But the oceans have a global ocean circulation where the surface water that's in equilibrium with the
atmosphere actually sinks and travels along the bottom of the ocean. So there is oxygen in the bottom of the
ocean, unlike many lakes where you don't have oxygen.
And we'll talk about that difference. And in the hot vents the water coming out of here can be very, very hot, but
there's a gradient right as it comes out meeting the colder sea water.
And so what you have here is a perfect incubator for chemosynthetic bacteria -- -- that use the hydrogen sulfide in
chemosynthesis to fix carbon dioxide using the oxygen here. And that forms the base of the entire food web in the
deep ocean because there's no light down there.
There's no photosynthesis. There's only chemosynthesis.

At the heart of these deep-sea communities is a process called chemosynthesis. Chemosynthesis is the use of
energy released by inorganic chemical reactions to produce food. It is analogous to the more familiar process of
photosynthesis. In photosynthesis, plants grow in sunlight, capturing solar energy to make organic matter. In
chemosynthesis, bacteria grow in mineral-rich water, harnessing chemical energy to make organic material.
Chemosynthesis can sustain life in absolute darkness.

The most extensive ecosystem based on chemosynthesis lives around undersea hot springs. At these hydrothermal
vents, a chemical-rich soup bubbles out of the crust and into the bottom of the sea. Boiling hot, saturated with toxic
chemicals and heavy metals, and more acidic than vinegar, vent waters are deadly to most marine animals.

This noxious brew is paradise to the bacteria that coats the rocks around the vent in thick orange and white mats.
The bacteria absorb hydrogen sulfide streaming from the vents, and oxidize it to sulfur. They use the chemical energy
released during oxidation to combine carbon, hydrogen, and oxygen into sugar molecules.

From this simple reaction, an entire ecosystem grows. Snails, clams, mussels, and a host of other grazing animals
feed on the bacterial mats. Crabs and shrimp eat the grazers, and then are hunted by larger crabs, fish, and octopi.

The largest and most abundant vent creatures are tube worms and giant white clams—animals that thrive because
they have developed a symbiotic, or mutually beneficial, relationship with the bacteria. Bacteria live within the hard-
shelled animals where they are protected from predators. The tube worms and clams receive a built-in food supply
because they absorb nutrients directly from the bacteria.

Tube worms, the signature inhabitants of hydrothermal vents, are absolutely dependant on their internal bacteria—as
adults they have no mouth or digestive system, no means of getting food apart from their symbionts. Their blood rich
tissues, colored red by hemoglobin, absorb dissolved gases from the vent water and from the seawater, and then
carry them to the bacteria. The bacteria convert the chemicals to organic matter and share the excess with the tube
worms.

This extraordinary relationship is highly satisfactory to both species. Millions of bacteria live safely within each tube
worm. The tube worms in return are so well nourished that they are the fastest growing invertebrates on Earth,
stretching up to 2 meters long in a single year.

Despite the total darkness, crushing water pressure, and temperatures that swing from above boiling to near freezing,
life is good at hydrothermal vents thanks to chemosynthetic bacteria. Vent faunas have both large biomass and high
diversity—over 300 species of animals have been found at vents, most living nowhere else on the planet.

But life based on chemosynthesis is also precarious. The hydrothermal vents—the source of life-sustaining chemicals
—can be extinguished at any time by earthquakes, lava flows, or rock falls. Many vents close after a few months or
years, and few seem to survive more than a couple of decades. Once the supply of chemicals stops, the bacteria die
and the rest of the fauna either migrates or perishes.

Chemosynthetic communities are also found in marine settings other than hydrothermal vents. At so-called cold-
seeps, where tectonic activity squeezes mineral water out of the ground and around sea bottom petroleum deposits,
methane, ammonia, and hydrogen sulfide are released. Bacteria use these compounds to make organic molecules,
which support a web of symbionts, carnivores, and scavengers.

Chemosynthesis and Hydrothermal Vent Life

http://noaacontent.nroc.org/lesson05/l5text.htm
At the heart of these deep-sea communities is a process called chemosynthesis. Chemosynthesis is the use of
energy released by inorganic chemical reactions to produce food. It is analogous to the more familiar process of
photosynthesis. In photosynthesis, plants grow in sunlight, capturing solar energy to make organic matter. In
chemosynthesis, bacteria grow in mineral-rich water, harnessing chemical energy to make organic material.
Chemosynthesis can sustain life in absolute darkness. Potential for intro

The most extensive ecosystem based on chemosynthesis lives around undersea hot springs. At these hydrothermal
vents, a chemical-rich soup bubbles out of the crust and into the bottom of the sea. Boiling hot, saturated with toxic
chemicals and heavy metals, and more acidic than vinegar, vent waters are deadly to most marine animals.

This noxious brew is paradise to the bacteria that coats the rocks around the vent in thick orange and white mats.
The bacteria absorb hydrogen sulfide streaming from the vents, and oxidize it to sulfur. They use the chemical energy
released during oxidation to combine carbon, hydrogen, and oxygen into sugar molecules.

From this simple reaction, an entire ecosystem grows. Snails, clams, mussels, and a host of other grazing animals
feed on the bacterial mats. Crabs and shrimp eat the grazers, and then are hunted by larger crabs, fish, and octopi.

The largest and most abundant vent creatures are tube worms and giant white clams—animals that thrive because
they have developed a symbiotic, or mutually beneficial, relationship with the bacteria. Bacteria live within the hard-
shelled animals where they are protected from predators. The tube worms and clams receive a built-in food supply
because they absorb nutrients directly from the bacteria.

Tube worms, the signature inhabitants of hydrothermal vents, are absolutely dependant on their internal bacteria—as
adults they have no mouth or digestive system, no means of getting food apart from their symbionts. Their blood rich
tissues, colored red by hemoglobin, absorb dissolved gases from the vent water and from the seawater, and then
carry them to the bacteria. The bacteria convert the chemicals to organic matter and share the excess with the tube
worms.

This extraordinary relationship is highly satisfactory to both species. Millions of bacteria live safely within each tube
worm. The tube worms in return are so well nourished that they are the fastest growing invertebrates on Earth,
stretching up to 2 meters long in a single year.

Despite the total darkness, crushing water pressure, and temperatures that swing from above boiling to near freezing,
life is good at hydrothermal vents thanks to chemosynthetic bacteria. Vent faunas have both large biomass and high
diversity—over 300 species of animals have been found at vents, most living nowhere else on the planet.

But life based on chemosynthesis is also precarious. The hydrothermal vents—the source of life-sustaining chemicals
—can be extinguished at any time by earthquakes, lava flows, or rock falls. Many vents close after a few months or
years, and few seem to survive more than a couple of decades. Once the supply of chemicals stops, the bacteria die
and the rest of the fauna either migrates or perishes.

Chemosynthetic communities are also found in marine settings other than hydrothermal vents. At so-called cold-
seeps, where tectonic activity squeezes mineral water out of the ground and around sea bottom petroleum deposits,
methane, ammonia, and hydrogen sulfide are released. Bacteria use these compounds to make organic molecules,
which support a web of symbionts, carnivores, and scavengers.

this reaction - CO2+ 4H2S + O2→ CH2O + 4S + 3H2O - is one of the most important chemosynthetic
pathways, supporting diverse deep-sea communities and helping to regulate the chemistry of ocean
water.
Hydrogen sulfide chemosynthesis: 18H2S + 6CO2 + 3O2 → C6H12O6 (carbohydrate) + 12H2O + 18S.
Instead of releasing oxygen gas while fixing carbon dioxide as in photosynthesis, hydrogen sulfide
chemosynthesis produces solid globules of sulfur in the process.
Chemosynthesis

Why do bacteria that live deep below the ocean’s surface rely on chemical compounds instead of
sunlight for energy to make food?

Most autotrophs make food by photosynthesis, but this isn’t the only way that autotrophs produce
food. Some bacteria make food by another process, which uses chemical energy instead of light energy.
This process is called chemosynthesis. In chemosynthesis, one or more carbon molecules (usually carbon
dioxide or methane, CH4) and nutrients is converted into organic matter, using the oxidation of
inorganic molecules (such as hydrogen gas, hydrogen sulfide (H2S) or ammonia (NH3)) or methane as a
source of energy, rather than sunlight. In hydrogen sulfide chemosynthesis, in the presence of carbon
dioxide and oxygen, carbohydrates (CH2O) can be produced:

CO2 + O2 + 4H2S → CH2O + 4S + 3H2O

Many organisms that use chemosynthesis are extremophiles, living in harsh conditions, such as in the
absence of sunlight and a wide range of water temperatures, some approaching the boiling point. Some
chemosynthetic bacteria live around deep-ocean vents known as “black smokers.” Compounds such as
hydrogen sulfide, which flow out of the vents from Earth’s interior, are used by the bacteria for energy
to make food. Consumers that depend on these bacteria to produce food for them include giant
tubeworms, like those pictured in Figure below. These organisms are known as chemoautotrophs. Many
chemosynthetic microorganisms are consumed by other organisms in the ocean, and symbiotic
associations between these organisms and respiring heterotrophs are quite common.