What Drives Symbiotic Calcium Signalling in Legumes? Insights and Challenges of Imaging.

We review the contribution of bioimaging in building a coherent understanding of Ca 2 + signalling during legume-bacteria symbiosis. Currently, two different calcium signals are believed to control key steps of the symbiosis: a Ca 2 + gradient at the tip of the legume root hair is involved in the development of an infection thread, while nuclear Ca 2 + oscillations, the hallmark signal of this symbiosis, control the formation of the root nodule, where bacteria fix nitrogen. Additionally, different Ca 2 + spiking signatures have been associated with specific infection stages. Bioimaging is intrinsically a cross-disciplinary area that requires integration of image recording, processing and analysis. We used experimental examples to critically evaluate previously-established conclusions and draw attention to challenges caused by the varying nature of the signal-to-noise ratio in live imaging. We hypothesise that nuclear Ca 2 + spiking is a wide-range signal involving the entire root hair and that the Ca 2 + signature may be related to cytoplasmic streaming.

Nuclear-localized cyclic nucleotide-gated channels mediate symbiotic calcium oscillations.

Nuclear-associated Ca(2+) oscillations mediate plant responses to beneficial microbial partners--namely, nitrogen-fixing rhizobial bacteria that colonize roots of legumes and arbuscular mycorrhizal fungi that colonize roots of the majority of plant species. A potassium-permeable channel is known to be required for symbiotic Ca(2+) oscillations, but the calcium channels themselves have been unknown until now. We show that three cyclic nucleotide-gated channels in Medicago truncatula are required for nuclear Ca(2+) oscillations and subsequent symbiotic responses. These cyclic nucleotide-gated channels are located at the nuclear envelope and are permeable to Ca(2+) We demonstrate that the cyclic nucleotide-gated channels form a complex with the postassium-permeable channel, which modulates nuclear Ca(2+) release. These channels, like their counterparts in animal cells, might regulate multiple nuclear Ca(2+) responses to developmental and environmental conditions.

The role of DMI1 in establishing Ca (2+) oscillations in legume symbioses.

Calcium (Ca (2+)) is a key secondary messenger in many plant signaling pathways. One such pathway is the SYM pathway, required in the establishment of both arbuscular mycorrhizal and rhizobial root symbioses with legume host plants. (1) When the host plant has perceived the diffusible signals from the microbial symbionts, one of the earliest physiological responses are Ca (2+) oscillations in and around the nucleus. (2) These oscillations are essential for activating downstream gene expression, but the precise mechanisms of encoding and decoding the Ca (2+) signals are unclear and still under intense investigation. Here we put forward a hypothesis for the mechanism of the cation channel DMI1.