Tuesday, December 01 200

Cyanobacteria live in the water, and can manufacture their own food through
"photosynthesis." Although the oldest known fossils, more than 3.5 billion years
old, are cyanobacteria, they are still around in large numbers; in fact, they one of
the largest and most important groups of bacteria. Individual cyanobacteria are
very small and usually just single cells, either round, ovoid, or stringlike in shape.
Some types grow in colonies that can be large. These colonies are built of many
layers, and are called stromatolites (if more or less dome-shaped) or oncolites (if
round).
The characteristic layered structure of fossilized stromatolites advertises their
presence, helping scientists locate them and identify their age through radioactive
dating of the surrounding rocks. To the upper left is a drawing of such a layered
colony, then of the layered surface, and to the lower left of a fossil cynaobacterium
such as would live near the surface.
As with all life, water is central to the life cycle of cyanobacteria. Because they are
small in size and so simple that they cannot work cooperatively, each living cell
needs to have access to a continuous supply. At the same time, they make food by
photosynthesis, using chlorophyll. That is, as with complex, advanced plants, they
absorb carbon dioxide from the atmosphere, and use the energy from the sun to
build it into complex energy-containing sugars, while releasing oxygen. To carry out
this process, each cell needs to be bathed in bright sunlight. To satisfy these two
critical requirements - access to water and sunlight - the cyanobacteria grow at the
shore of the ocean or the edges of ponds or pools of water. Single cells would be
easily dislodged from this location and would float to regions where they could not
survive, so the bacteria live in sheet-like films (thin enough so each cell has access
both to water and sunlight) that are held together by a slime that they secrete. The
stringlike ones can glide toward light at a speed of a few millimeters per hour, and
become tangled in their competition to get positioned to receive sunlight. Thus,
they develop into a felt-like structure that makes a particularly robust living mat.
Another complication for stromatolites on the early Earth is that the gases in the
atmosphere did not absorb ultraviolet photons from the sun well. Although
stromatolites are somewhat UV-resistant, it is likely that they tended to grow just
under the surface of the water to gain some additional protection. For colonies
exposed to direct sunlight, a top level of bacteria killed by the UV light may have
served to protect lower-lying layers.

The cyanobacteria form a veneer over a complex, layered colony.

Underneath them is another layer of photosynthetic bacteria that absorb sunlight at
wavelengths where the cyanobacteria are transparent. These bacteria are poisoned
by oxygen, so they are termed "anaerobic"; the mat of cyanobacteria acts as a
protective shield. Additional underlying layers, which can be millimeters or
centimeters deep, contain other forms of anaerobic bacteria. Because they do not
receive sunlight and do not conduct photosynthesis, these bacteria feed on dead
photosynthetic bacteria that have been left behind by the gliding of the live ones
toward the sun.

This structure by itself would be a stable colony. However, from time to time
some external event - say a rainstorm - causes the top layer of the mat to be buried
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in a layer of mud. The cyanobacteria then stop secreting slime and, freed from the
mat, glide upward toward the faint light filtering through until they are once again
in the sun. The underlying anaerobic bacteria follow along, since their survival
depends on the colony structure. In this way, a new layer grows on top of the silt,
and after many repetitions the process yields a multi-layered structure that can
grow to a foot or more in height. As the old, underlying layers dry out and are
compressed, the silt in them solidifies into rock.

To the far left is a fossil colony, and to the left a living one (from Schopf),
showing how this pattern has remained stable for billions of years! Detailed
comparisons of individual fossils of ancient bacteria also show them to be virtually
identical to those found in living colonies. Much of what we describe about ancient
stromatolites is based on our observations of the behavior of living ones, but it
appears that this line of evidence should be quite reliable. Stromatolites appear to
be the ultimate "living fossils", life forms that have survived for 3.5 billion years with
virtually no modifications of their form or mode of surviving. Stromatolites ruled the
earth for billions of years. One line of evidence is the huge deposits of fossils they
have left. Indeed, because of their striking appearance, they have been used as
ornamental stonework. To the left is a children's playground slide in China, made of
1.5-billion-year old stromatolite fossils (picture from Schopf). To the right is the
Chinese parliamentary building; the columns are stromatolite fossils of the same
age; note the people and van for scale (picture by G. Rieke).

Even more dramatically, the photosynthesis carried out by the stromatolites

was on such a large scale that they consumed most of the carbon dioxide in the
atmosphere and released enough oxygen to make permanent changes. The release
of enough greenhouse gases by humans to change the climate through global
warming makes us only the second species to have a major effect on the climate of
the earth, and at least so far our influence is dwarfed by that of these bacteria!

How did they do it? The simple cells of cyanobacteria can reproduce quickly,
in only about 30 minutes. When conditions change, the members of a colony that
are best able to cope are the ones that tend to survive, and since the reproduction
time is so short, their offspring can be manifold and continue the growth of the
colony. In this process of "natural selection," the colony soon consists only of
individual bacteria adapted to the new conditions. It is a different type of survival
than we usually associate with more complex forms of life. For example, if there is a
harsh winter, we are interested in whether enough individuals of lived through the
winter and can establish a breeding stock to produce a next generation that can
continue the species. With cyanobacteria in stromatolite, there may be as many as
10,000 generations in a single long, harsh winter. Thus, the survival depends on the
most cold-tolerant members of the colony producing offspring, and the nature of the
members of the colony will be subtly modified at the end of the ordeal. If a hot
summer follows, then the process of natural selection will favor those that survived
the harsh winter but still have some heat tolerance.

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Modern stromatolites have used this process to search out ecological niches
that are too harsh for other species. Thus, the snails and other animals that might
feed on them cannot tolerate extremely salty water, so to avoid predators the
colonies grow in bays where evaporation results in extremely saline pools of
seawater. Cyanobacteria have survived in

other ways. One of the most interesting is through symbiosis, where a cell
merges with another cell in a way that helps both survive. The most dramatic
example is that the chloroplast with which plants make food for themselves is
actually a cyanobacterium living within the plant's cell. The photosynthesis is
centered in the chloroplast, while the other parts of the cell provide a protective
environment for the chloroplast and integrate it into the plant.

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