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Structurally,
prokaryotic organisms are far simpler than eukaryotic life forms.
Yet, prokaryotes are functionally or metabolically diverse with
regards to the reactions they mediate or stresses they can endure.
Prokaryotes do not possess membrane-bound organelles such as a nucleus,
endoplasmic reticulum, mitochondria, or chloroplasts. Therefore,
prokaryotes can not separate metabolically incompatible biochemical
processes into discrete compartments. The inability to compartmentalize
cellular processes poses a bottleneck for the biochemical and structural
evolution of prokaryotes. As a result, prokaryotes have diversified
with regards to biochemical abilities while remaining structurally
simple.
The prokaryotic solution to the packaging problem is the formation
of associations with other organisms the promote protection from
potentially inhibitory environmental factors. These include exposure
to adverse oxygen concentrations, ltraviolet radiation, desiccation,
adverse pH, and Eh. The steep biogeochemical gradients that exist
in mats allow and select for functional diversification such that
fully organisms possessing aerobic, microaerophilic, and anaerobic
requirements may co-exist and contemporaneously function along a
gradient. Such dramatic environmental changes occurring in a small
spatial scale set up association that facilitate mutualistic nutrient,
gas, and metabolite exchange. Associations reflect synergistic or
syntrophic lifestyles where growth and biogeochemical cycling are
conducted more effectively and efficiently than on an individual
population basis. Such associations may be termed microbial consortia.
Functionally, a consortium exceeds the sum of its parts. Members
of the consortium maintain metabolic and ecological compatibility,
as long as biogeochemcal and environmental gradients allow for individual
niches to exist in close proximity. Microbial mats typify these
conditions and, accordingly, are the focus of research on consortial
growth strategies in extreme environments.
The close spatial coupling and metabolic interdependence of microorganisms
in extreme environments begs the question of whether or not there
exist pairs or groups of organisms that are specifically dependent
on one another because of the traits each has evolved (i.e. have
consortial members co-evolved?). Alternatively, do consortia represent
functional relationships of opportunity in which participants are
not particular from whom they derive their needs? There are many
examples of obligate symbiotic relationships between prokaryotic
and eukaryotic organisms, e.g. rhizobia and leguminous plants, the
Anabaena-Azolla sp. (cyanobacterial-fern) association, methanogens
and sulphur bacteria with anaerobic ciliates, and sulfur bacteria
and Rivularia worms near deep-sea hydrothermal vents. In each of
these examples, a strong case may be made for co-evolution, whereby
strong selective pressures for highly specific traits allow the
pair to exploit a particular niche and survive. We do not know whether
similar relationships, as opposed to opportunistic encounters, characterize
microbial consortia. There are cases in which specific associations
between prokaryotes have been documented. Most widely known is the
relationship between heterotrophic bacteria (Pseudomonas sp. and
Zoogloea sp.) with the heterocysts of an Anabaena sp. Additionally,
there are instances of 'metabolic coupling between specific bacterial
strains and the cyanobacteria, Oscillatoria redekii. Anecdotal evidence
includes the fact that many cultured cyanobacterial isolates are
lost and no longer viable once their bacterial associations are
completely removed. The evolutionary basis for the development of
microbial consortia is a an under explored area of research!
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