Morphological evolution from a unicellular to multicellular state provides greater opportunities for organisms to attain larger and more complex living forms. As the most common freshwater cyanobacterial genus, Microcystis is a unicellular microorganism, with high phenotypic plasticity, which forms colonies and blooms in lakes and reservoirs worldwide. We conducted a systematic review of field studies from the 1990s to 2017 where Microcystis was dominant. Microcystis was detected as the dominant genus in waterbodies from temperate to subtropical and tropical zones. Unicellular Microcystis spp. can be induced to form colonies by adjusting biotic and abiotic factors in laboratory. Colony formation by cell division has been induced by zooplankton filtrate, high Pb concentration, the presence of another cyanobacterium (Cylindrospermopsis raciborskii), heterotrophic bacteria, and by low temperature and light intensity. Colony formation by cell adhesion can be induced by zooplankton grazing, high Ca concentration, and microcystins. We hypothesise that single cells of all Microcystis morphospecies initially form colonies with a similar morphology to those found in the early spring. These colonies gradually change their morphology to that of M. ichthyoblabe, M. wesenbergii and M. aeruginosa with changing environmental conditions. Colony formation provides Microcystis with many ecological advantages, including adaption to varying light, sustained growth under poor nutrient supply, protection from chemical stressors and protection from grazing. These benefits represent passive tactics responding to environmental stress. Microcystis colonies form at the cost of decreased specific growth rates compared with a unicellular habit. Large colony size allows Microcystis to attain rapid floating velocities (maximum recorded for a single colony, ∼ 10.08 m h ) that enable them to develop and maintain a large biomass near the surface of eutrophic lakes, where they may shade and inhibit the growth of less-buoyant species in deeper layers. Over time, accompanying species may fail to maintain viable populations, allowing Microcystis to dominate. Microcystis blooms can be controlled by artificial mixing. Microcystis colonies and non-buoyant phytoplankton will be exposed to identical light conditions if they are evenly distributed over the water column. In that case, green algae and diatoms, which generally have a higher growth rate than Microcystis, will be more successful. Under such mixing conditions, other phytoplankton taxa could recover and the dominance of Microcystis would be reduced. This review advances our understanding of the factors and mechanisms affecting Microcystis colony formation and size in the field and laboratory through synthesis of current knowledge. The main transition pathways of morphological changes in Microcystis provide an example of the phenotypic plasticity of organisms during morphological evolution from a unicellular to multicellular state. We emphasise that the mechanisms and factors influencing competiti...

Microcystis aeruginosa and Cylindrospermopsis raciborskii are two cyanobacterial species that dominate freshwaters globally. Multiple strains of each species with different physiology occur, however, many studies have focused only on one or two strains, limiting our understanding of both strain variation and characterisation of the species. Therefore, in this study we examined the variation in growth and morphology of multiple isolates of both species, isolated from two adjacent Australian reservoirs. Four M. aeruginosa strains (=isolates) (one colony-forming, three single-celled morphology) and eight C. raciborskii isolates (five with straight trichomes, three with coiled trichomes) were cultured individually in a factorial designed experiment with four light intensities (L: 10, 30, 50 and 100μmol photons ms) and two temperatures (T: 20 and 28°C). The specific growth rate (μ), cell volume, and final cell concentration was measured. The light attenuation coefficient (k), a measure of self-shading, was calculated. The results showed that the intraspecific variation was greater than the interspecific variation. The μ of all isolates of M. aeruginosa and C. raciborskii ranged from 0.16 to 0.55d and 0.15 to 0.70d, respectively. However, at a specific light and temperature the mean μ of all M. aeruginosa isolates and C. raciborskii isolates were similar. At the species level, M. aeruginosa had higher growth rates at higher light intensity but lower temperature (L100T20), while straight C. raciborskii had higher growth rates at lower light intensity but higher temperature (L50T28), and coiled C. raciborskii had higher growth rates at higher light intensity and higher temperature (L100T28). The final cell concentrations of M. aeruginosa were higher than C. raciborskii. However, C. raciborskii isolates had greater variation in μ, k and cell volume than M. aeruginosa. k varied with light and temperature, and decreased with surface-to-volume ratio within each species. k was lower for M. aeruginosa compared to C. raciborskii as expected based on cell size, but interestingly, C. raciborskii coiled isolates had lower k than the straight isolates suggesting lower effect of self-shading. This study highlights the extent of strain variation to environmental conditions and to species variability.

The freshwater cyanobacterium Microcystis is a nuisance species. It forms large blooms on the water surface and overwhelmingly dominates the ecosystem through the formation of colonies from single cells surrounded by mucilage; however, the mechanism of colony formation is poorly understood. Two mechanisms of Microcystis colony formation have been proposed: cell-division, where cells remain attached after binary fission; and cell-adhesion, where single cells stick together. This paper examined the published literature on Microcystis colony formation to clarify the mechanism of colony formation and its relationship to environmental drivers. This meta-analysis showed that in laboratory experiments, colony formation by cell-division was mainly induced by zooplankton filtrate, high Pb concentrations, the presence of the cyanobacterium Cylindrospermopsis raciborskii, heterotrophic bacteria, and low temperature and low light intensities. Alternatively, colony formation by cell-adhesion was mainly induced by zooplankton grazing, high Ca concentrations, and microcystins. Therefore, colony formation by cell-division appears to be a slower process and to occur under an environmental stress factor, while cell-adhesion occurs more quickly to an environmental threat. Applying the criteria to the different morphospecies of Microcystis, it was found that under natural conditions M. ichthyoblabe colonies formed predominantly through cell-division, whereas M. wesenbergii colonies formed predominantly through cell-adhesion. This study provides new insights into the mechanisms and environmental drivers of colony formation by Microcystis.