Occurrence of Raphidiopsis raciborskii blooms in cool waters: Synergistic effects of nitrogen availability and ecotypes with adaptation to low temperature.

Raphidiopsis raciborskii is a diazotrophic and potentially toxic cyanobacterium. To date, this species has successfully invaded many regions from the tropics to sub-tropical and temperate regions, typically forming blooms at temperatures greater than 25 °C. However, there have been a few cases in which R. raciborskii blooms have occurred at low temperatures (below 15 °C), but its cause and mechanisms remain unclear. In this study, field investigations revealed that R. raciborskii blooms occurred at 10-15 °C in Lake Xihu, Yunnan, China. The biomass of R. raciborskii was found to be positively related to nitrate concentrations in this lake. Three strains of R. raciborskii, two isolated from Lake Xihu (CHAB 6611 and CHAB 6612) and one from Lushui Reservoir in central China (CHAB 3409), were used for growth experiments at 15 °C. The three strains exhibited genotypic (16S rRNA and ITS-L genes) and physiological differences in response to nitrogen concentrations at low temperature. The growth rates of strains CHAB 6611 and CHAB 6612 increased with nitrogen concentration while CHAB 3409 could not grow at 15 °C. Furthermore, the growth and phenotypic responses of CHAB 6611 and CHAB 6612 to nitrogen concentrations were different, despite the closer genetic relationship shared by these two strains. Thus, increased nitrogen concentration in water may enhance the biological availability and utilization of nitrogen by R. raciborskii, which is the external promoter, leading to improving the resistance of R. raciborskii to low temperature. The internal cause is the presence of ecotypes in R. raciborskii populations with adaptation to low temperature. With increasing global eutrophication, the distribution range of R. raciborskii as well as the scale of its blooms will increase. As such, the risk of exposure of aquatic biota and humans to cylindrospermopsin is also expected to increase.