B. J. Finlay

Respiration rates in heterotrophic, free-living protozoa

Published estimates of protozoan respiratory rates are reviewed with the object of clarifying their value in ecological studies. The data show a surprisingly large variance when similarly sized cells or individual species are compared. This is attributed to the range of physiological states in the cells concerned. The concept of basal metabolism has little meaning in protozoa. During balanced growth, energy metabolism is nearly linearly proportional to the growth rate constant; at the initiation of starvation, metabolic rate rapidly declines. Motility requires an insignificant fraction of the energy budget of protozoans. For growing cells, metabolic rate is approximately proportional to weight0.75 and the data fall nearly exactly on a curve extrapolated from that describing the respiration rates of poikilotherm metazoans as a function of body weight. It is conceivable that protozoan species exist with lower maximum potential growth and metabolic rates than those predicted from cell volume and the equations derived from the available data. However, the lack of information concerning the state of the cells studied prevents verification of this idea. Laboratory measurements of protozoan respiratory rates have no predictive value for protozoa in nature other than delimiting a potential range. For small protozoans, this range may, on an individual basis, represent a factor of 50.

Freshwater protozoa: biodiversity and ecological function

The purpose of this article is to pull together various elements from current knowledge regarding the natural history of free-living protozoa in fresh waters. We define their functional role, set the likely limits of ‘biodiversity’, and explore how the two may be related. Protozoa are unicellular, phagotrophic organisms, and 16 phyla of protists contain free-living freshwater protozoan species. They are the most important grazers of microbes in aquatic environments and the only grazers of any importance in anoxic habitats. In sediments, ciliates are usually the dominant protozoans. Benthic ciliate biomass accounts for slightly less than 10% of total benthic invertebrate biomass, but ciliate production may equal or even exceed invertebrate production. Freshwater protozoan species are probably ubiquitous, although many may persist locally for long periods in a cryptic state – as ‘potential’ rather than ‘active’ biodiversity. As protozoa are among the largest and most complex of micro-organisms, it follows that bacteria and all other smaller, more numerous microbes are also ubiquitous. The number of protozoan species recorded in local surveys (232) is about 10% of the estimated global species richness (2390). The seedbank’ of protozoan (and microbial) species ensures that local microbial diversity is never so impoverished that it cannot play its full part in ecosystem functions such as carbon fixation and nutrient cycling.

On the abundance and distribution of protozoa and their food in a productive freshwater pond

We have examined and quantified the protozoa living in a productive freshwater pond during a 2-day period in June 1987. Over 90 species were recognised. The planktonic and benthic communities were dominated by ciliates and heterotrophic flagellates although the large amoeba Pelomyxa palustris was abundant (102 ml(-1)) in anaerobic sediments. Picoplankton averaged 1.4 × 10(7) ml(-1), phototrophic nanoplankton 0.8 × 10(5) ml(-1), heterotrophic nanoplankton 0.9 × 10(5) ml(-1) and planktonic ciliates 1.3 × 10(2) ml(-1). Numbers were about two orders of magnitude higher in the sediment. Protozoan biomass ranged from 3% to 61% of the total plankton biomass. Heterotrophic flagellates were the principal grazers of the picoplankton. Planktonic ciliates fed mainly on phototrophic nanoplankton but they probably also ingested heterotrophic flagellates. Benthic ciliates were predominantly bactivorous. Competition between ciliate species was minimised by both spatial and food niche separation. Ten species of planktonic ciliates appeared to contain algal symbionts: one species (Strombidium viride) contained structures resembling sequestered chloroplasts. These findings concerning the diversity and abundance of protozoa in a freshwater pond are consistent with the consensus opinion expressed in the marine literature that protozoa play a fundamental role in microbial food webs within aquatic ecosystems.

Biodiversity at the microbial level: The number of free-living ciliates in the biosphere

For more than 200 years, ciliated protozoa have been identified and allocated Species names largely on the basis of the rich morphological variety they present. We have examined the species richness of all free-living ciliate genera, described historical trends in the descriptions of new species, and estimated the number of species currently known. We have quantified the value of taxonomic revisions, and conclude that the number of known, extant free-living species is close to 3000. We have investigated the concept of species and the meaning of biodiversity in relation to ciliates, and conclude that the bilogical species concept is neither appropriate nor practicable. Insofar as ciliate morphology is closely correlated with the function of the organism in nature, the morphospecies concept is as valid as any, and probably more pragmatic than any other. Thus, when speaking of species diversity, or the biodiversity of ciliates, we refer to diversity of form and function. The majority of ciliate species in the more frequently studied habitats have probably already been discovered, but an accurate picture of ciliate diversity on a global scale will require substantial taxonomic revision of many long-established and crowded genera, together with the investigation and description of new forms from previously unexplored habitats.

Cyclidium porcatum n. sp.: a Free-living anaerobic scuticociliate containing a stable complex of hydrogenosomes, eubacteria and archaeobacteria

A new ciliate species (Cyclidium porcatum) is the first freshwater anaerobic scuticociliate to be cultured and described. It contains a unique tripartite structure consisting of hydrogenosomes (confirmed by cytochemical staining for hydrogenase), interspersed with methanogens (confirmed by auto fluorescence and in situ hybridisation with an archaeobacterial 16S rRNA-specific probe) and unidentified eubacteria (confirmed with a eubacterial 16S rRNA-specific probe). This complex structure is stable and persistent, indicating that it is an anaerobic symbiotic consortium incorporating three functional partners.

Ciliated protozoa from a volcanic crater-lake in Victoria, Australia

We have investigated the ciliated protozoa living in the crater-lake of an extinct volcano in Australia. Our principal objective was to discover if such a habitat— geographically distant and isolated from Europe (the latter having provided most of the diversity on which ciliate taxonomy is based)— could yield species that were unusual, and perhaps new to science. Numerous samples were taken from the superficial layer of lake sediment, and examined fresh in the laboratory. Thereafter, the samples were manipulated to encourage growth of rare and cryptic ciliate species. Eighty-five species of ciliated protozoa were identified. None of these was new, all having been found previously in fresh-or brackish water, sea water, or soil. All, apart from one (Oxytricha salmastra), are already known from Europe. In order to test our ability to discover new ciliate species, we examined a variety of water samples from other lakes, including those known to harbour endemic algae. One new ciliate species (Lembadion curvatum) was discovered in a lake in Western Tasmania. We conclude that the ciliate fauna of Australia is remarkably similar to that in Europe and other parts of the world. This is supported by the example of those ‘endemic’ ciliate species described in the literature which have, in response to additional sampling, been found elsewhere in the world.