The smallest organisms may have played the largest role in influencing the Earth's biogeochemistry, and a special section in this week's issue of Science called Environmental Microbiology ( ) indicates that microorganisms have affected terrestrial, aquatic, and atmospheric environments through their sheer abundance, diversity, and metabolic potential. Understanding how various microbial communities assemble, function, and survive requires elucidating how their physical, chemical, and biological components interconnect. How does a particular consortium of bacteria form or catabolize particular metals and minerals in the ocean or in the deep layers of the Earth's mantle? How do certain microorganisms thrive in aerobic, anaerobic, or extreme conditions of temperature, pH, or salinity? The dynamic relations of species with one another and with their environment help define a particular microbial ecosystem. Such relations are regulated in part by biochemical processes that control how microorganisms respond to, and affect, their local environment as well as each other. We are at the frontier of understanding how particular networks of genes, proteins, and signaling pathways define microbial communities and consequently, the microbiology that underlies earth history. The general ubiquity and diversity of microbial eukaryotes and prokaryotes, described in Viewpoints by Finlay ( ) and by Torsvik et al . ( ) , indicates that microorganisms can make a living just about anywhere, and as a result, microbial communities are heterogenous and complex. Clues to the microbial strategies employed for survival and growth in a given environment have been gleaned from studying isolated species or mixed populations under defined conditions. For example, bacteria may detect changes in osmolarity and temperature through physical changes in their membranes and associated alterations in membrane-associated signaling proteins such as receptors and ion channels [see Perspective by Los and Murata ( ) in the STKE Archive]. Reviews by Reysenbach and Shock ( ) and by Fenchel ( ) point out that because motility is the primary mechanism by which microorganisms seek out optimal environments, nearly all microorganisms are motile during some part of their life cycle. Gliding, amoeboid-like motion, or flagella-driven movement can propel a microorganism in the right direction. A Perspective by Dahlquist ( ) describes our current understanding of how a bacterium can respond to temporal changes in the intensity or concentration of a stimulus by modulating its flagellar rotary engine. Alterations in bacterial orientation and movement (chemotaxis) involve transducing a perceived environmental change to the flagella motor. A phosphorelay signaling system is in place that couples transmembrane chemoreceptors with a sensor kinase and a response regulator, a cascade that ultimately affects the rotational bias of the flagella. Signal amplification can occur at multiple points of this pathway and receptor methylation further marks a of a bacterium's exposure to a stimulus, ensuring appropriate responses to the same stimulus later on. Santos and Shiozaki ( ) describe a similar phosphorelay system in yeast, indicating that growth responses and morphological transitions in such lower eukaryotes utilize a conserved signaling mechanism (see Review in the STKE Archive). Stock et al . ( ) points out that although each bacterium in a population should move toward optimal conditions for survival and growth, microbial dispersal requires a diversity of responses that permits exploration in other directions. A molecular understanding of how bacteria efficiently interpret multiple stimuli and choose the best direction to move in is not yet clear. In contrast to a free-swimming state, three-dimensional biofilms represent stable, surface-adherent communities of bacteria that are largely cooperative and communicate through chemical signals. They are not only geologically important, but their link to persistent bacterial infections and antibiotic resistance have made them a force to be reckoned with in medicine. A Review by Schembri et al . ( ) describes the signaling mechanisms that Gram-negative bacteria use to control adhesion during biofilm development. Managing adhesion in response to environmental conditions requires the expression of adhesin molecules and cell surface extensions called fimbrae in coordination with a process called autoaggregation. A cell-to-cell signaling mechanism called quorum sensing is also in place to monitor biofilm population density. If the community structure is such that expansion into nutrient-rich areas is needed, bacterial signaling molecules called N -acylhomoserine lactones are synthesized that stimulate biosurfactant production and induce cellular flagellation to move the biofilm. As Newman and Banfield ( ) point out in their Review, our present knowledge of microbial life in geological systems and the evolution of Earth history has involved integration of geochemical and biological data. Genomic sequence information, DNA microtechnology, and development of good model systems should bring further insight into the relationships between gene expression, inter- and intracellular signaling, and survival strategies that underlie the diversity and complexity of microbial life. Featured in This Focus Issue on Environmental Microbiology Related Resources at STKE
Read full abstract