PHYSIOLOGICAL ECOLOGY OF CYANOBACTERIA IN MICROBIAL MATS AND OTHER COMMUNITIES

In this review some aspects of the physiological ecology of cyanobacteria are discussed by taking a microbial mat as an example. The majority of microbial mats are built and dominated by cyanobacteria which are primary producers at the basis of the microbial foodweb in microbial mats. These micro-scale ecosystems are characterized by steep and fluctuating physico-chemical gradients of which those of light, oxygen and sulphide are the most conspicuous. Light is strongly attenuated in the sediment, and owing to constant sedimentation, the mat-forming cyanobacteria have to move upwards towards the light. However, at the sediment surface, light intensity, particularly in the u.v. part of the spectrum, is often deleterious. The gliding movement of the cyanobacteria, with photo- and chemotaxis, allows the organism to position itself in a thin layer at optimal conditions. The organic matter produced by cyanobacterial photosynthesis is decomposed by the microbial community. Sulphate-reducing bacteria are important in the end-oxidation of the organic matter. These organisms are obligate anaerobes and produce sulphide. Gradients of sulphide and oxygen move up and down in the sediment as a response to diurnal variations of light intensity. Cyanobacteria, therefore, are sometimes exposed to large concentrations of the extremely toxic sulphide. Some species are capable of sulphide-dependent anoxygenic photosynthesis. Other cyanobacteria show increased rates of oxygenic photosynthesis in the presence of sulphide and have mechanisms to oxidize sulphide while avoiding sulphide toxicity. Iron might play an important role in this process. Under anoxic conditions in the dark, mat-forming cyanobacteria switch to fermentative metabolism. Many species are also capable of fermentative reduction of elemental sulphur to sulphide. The gradients of sulphide and oxygen are of particular importance for nitrogen fixation. Very fe iv microbial mars are formed by heterocystous cyanobacteria, which are best adapted to diazotrophic growth. However, these organisms probably cannot tolerate greater concentrations of sulphide or anoxic conditions or both. Under such conditions non-heterocystous cyanobacteria become dominant as diazotrophs. These organisms avoid conditions of oxygen supersaturation. In the ecosystem, nitrogen fixation and photosynthesis might be separated temporally as well as spatially. In addition, non-heterocystous diazotrophic cyanobacteria have mechanisms at the subcellular level to protect the oxygen-sensitive nitrogenase from inactivation.