Substrates

The existence of substrates and their physical and chemical nature is importance
for sessile organisms, as well as for those which need a surface for locomotion.
From the physical point of view, a low stream velocity is important and can be
observed close to surfaces in running waters. The peripheral region and dead
zones in streams and rivers are important habitats for protozoa.

In nearly all biotopes, one part of the community consists of sessile protozoa. In
the vicinity of sediment particles and aggregats, especially high numbers of
species can be observed due to the high abudance of microbial food particles
and potential nutrients.

Sessile filter feeders use mechanical fixation to ensure that the ciliary moion
results in the collection of food particles instead of locomotion. Simultaneously,
the cell needs to be elevated from its substrates to allow the creation of
adequate water flow. This is ensured by stalks or elongated body forms. E.g in
choanoflagellatas and ciliates such as Vorticella and Stentor. The effectiveness of
the filtering activity can be heightened by the formation of colonies, which can
become suspended in the water e. G. Spherical colonies of certain
chaonoflageliates and small colonies of ciliates such as Ophrydium versatile.

Crwaling amoeba overwhelm their prey by creating a trap formed by pseudopods

pressed againts the substrate. The plasmodia of Reticulomyxa
(Granuloreticulosa) form with their reticulopodia large networks to catch large
prey organisms such as Volvox colonies or rotifers.

Certain sessile ciliates attach, exclussively or at least preferentially to organic or

living subtrates. This is particularly true for the ciliate genus Spirochona,
members of which have been found, so far exclusively, on the gill plates of
gammarids or for certain Apostomatia, which are attached to the surfaces of
crutaceans. Certain suctorians live on animal or plants in a species-specific way
e. G. Tokophrya lemnae living on rhizomes of species of Lemna. The evolutionary
significance of this high degree of specialization might be seen in the avoidance
of competition. The mechanisms of recognition of the substrate, however, are
unknown.

Biotic factors

As mentioned above, the most important biotic factors are food resources,
competition with other organisms, and also predator-prey relationships.

Species with generation times of several weeks or months, such as foraminifers,

survive only in very stable habitats. Normally, they represent one link in a rather
complex biocoenose which is predetermined by the limitations of a given
habibat. In contrast to these, protozoa with reproductive times of a few hours
make rapid use of resources. Such populations, be they amoebae, flagellates, or
ciliates, tend to collapse as rapidly as they developed.
The first step in establishing a new biocoenose is the settlement of a new
habitat. Experiements with artificial substrates in fresh water ponds have shown
that within three weeks, about 60 different species settled on these substrates.
On the other hand, under extreme situations, a new settlemen might be
established very slowly, if, for instance a new habitat is distant from a source of
inoculating species.

The pioneer species are normally small flagellates such as choanoflagellates and
bodonids, which are followed by small amoebae and cilliates. Their food is
bacteria, cyanobacteria, and unicellular algae e. G. Diatoms. The composition of
the community of food organisms is greatly influenced (quantitavely, as well as
qualitatively), by the feeding activity of the protozoa. In the course of the
establishment of these biocoenoses are unstable, and that they can change very
rapidly and dramatically, often within one day.

In natural biotopes, the situation is more complex due to the influence of

additional factors on population densities of various species. One of these factors
is, for instance, the appearance of metazoan predators in protozoan
biocoenoses. Some of the protozoa will be ingested, for instance, by larval fish,
water snails, platyhelminths, or oligochaetes; others will die due to competition
for food with rotifers.

Protozoan populations can live in habitats in which few if any metazoa exist. As
shown for the marine interstitial, the microhabitats might be to small for
metazoa to enter. Other habitats with high numbers of protozoa may have
extreme physical or chemical characteristics, such as anaerobic systems where
few if any metazoa can survive. Finally, very dense populations of protozoa are
found in artificial habitats e.g. the activated sludge of biological sewage
treatment plants. In these plants, the sewage is pumped into large tanks in
which the organic material is degraded by bacteria that develop very dense
populations. This food resource is used by protozoa, especially sessile, filter-feed-
ing ciliates. Since the flow of water is relatively high in these plants, metazoan
predators cannot develop dense populations.

In cases of food resource depletion, protozoa have developed characteristic

survival strategies. One, as already mentioned, is the formation of resting cysts.
Sessile species may become temporarily motile by active swimming or by
passive flotation to other habitats. Examples of these strategies are the
formation of the telotroch by Vorticella and the transformation into flagellated
stages by schizopyrenids or into floating stages (radiosa form) by naked
amoebae. Others become completely immotile and switch to a minimal
metabolic rate which provides only 2-4% of energy normally needed for survival.
Another possibility is the secretion into the surrounding medium of metabolites
which inhibit growth. This phenomenon is called eg bacteriostasis, fungistasis,
and ciliostasis. Conspecific consumption is also known through the development
of macrostomes capable of cannibalism.

Aquatic Biocoenoses and Habitats

On a global scale, the aquatic biocoenoses of the oceans are the source of most
biological productivity. The role that unicellular organisms play has been explored
only during the last two decades.

Firstly, it is striking that the species composition of the plankton, neuston, and
benthos is very similar in marine and limnic habitats. Exceptions are the
radiolarians and foraminifers, as well as the large, heterotrophic dinoflagellates
which are found exclusively in the marine environment, especially in the marine
environment, especially in the open oceans, additionally, a typical interstitial
fauna, composed mainly of karyorelictid ciliates, is known only for marine
habitats. Consequently, the similarities between marine and freshwater habitats
concentrate mainly on autotrophic and heterotrophic flagellates, ciliates, and
amoebae.

A major habitat type is the pelagic water column which is characterized by a

certain uniformity of enviromental parameters. not just exclusively planktonic,
but also sessile forms attached to suspended particles. These biotopes are
inhabited by prokaryotes, eukaryotic unicells, and also by metazoa.

Compared to the pelagial, benthic habitats exhibit a versatile and heterogenous

structure. Local differences in chemical parameters, surface properties, and food
resources lead to the formation of microhabits with specific faunas. Some
microhabitats exist exclusively due to their small size. An example is the marine
interstitial, with grain sizes of 100 to 250 m. Normally, food competitors and
predators cannot invade the habitat due to their body size. It is therefore mainly
colonized by slender or verniform protozoa.

These and similar benthic microhabitats might be additionally subdivided

vertically. The basis for stratification horizons is mainly the gradient oxygen,
which is dictated by the degree of underlying bacterial degradation, and the flux
of O2 into the sediment. At a certain depth, the oxygen concentration is too low
for aerobic microorganisms, and the redox potential becomes negative. Under
these reducing, anoxic condotions, degradation in exclusively anaerobic. Many
ciliates which live in such habitats, prefer a certain redox potential and inhabit
specific horizons in the sediment.

Inside the different sediment horizons, protozoa live in specific ecological niches.
This is based on specialization on specific diets. For instance, some feed on
anaerobic, others an aerobic bacteria, some feed on algae, protozoa and small
metazoa, others on detritus. The specialization can reach such a point as
detected by different species of the cilliate genus Remanella, which live in the
same habitat but feed on differently sized food particles.

Small organisms are characterized by relatively high metabolic rates. The

contribution of the protozoa to the cycle of organic material is normally very
high. Experiments show that protozoa speed up the rate of degradation of
sedimented plant material. The experimentally obtained result that protozoa, in
cooperation with bacteria have the fastest rate of degradation, seems at first
glance inconsistent because bacteria are also phagocytosed by the ciliates.
However, the explanation for this apparent paradox is that due to the digestion
of bacteria, the surrounding medium becomes enriched in essential nutrients
which positively influence the growth and reproduction of the remaining bacteria.
Furthermore, ciliates positively influence the density of bacterial populations by
their ciliary activity, which causes a better circulation of dissolved nutrients and
gases in the fluid microhabitat. Additionally, larger protozoa are involved in the
destruction of animal and plant material as carnivorous and herbivorous
organisms. Of special importance are ciliates belonging to the genus
Ophryoglena, as well as the macrostome forms of Tetrahymena. The filopodial
organized stages of vampyreliids and the labyrinthulids also belong to these
protozoan groups.