KAI2-and MAX2-Mediated Responses to Karrikins and Strigolactones Are Largely Independent of HY5 in Arabidopsis Seedlings

Karrikins are butenolide compounds released from burning vegetation that stimulate seed germination and enhance seedling photomorphogenesis. Strigolactones are structurally similar plant hormones that regulate shoot and root development, and promote the germination of parasitic weed seeds. In Arabidopsis, the F-box protein MAX2 is required for responses to karrikins and strigolactones, and the a/~ hydrolase KAI2 is necessary for responses to karrikins. Both MAX2 and KAI2 are essential for normal light-dependent seedling development. The bZIP transcription factor HY5 acts downstream of multiple photoreceptors and promotes photomorphogenesis, but its relationship with MAX2 and KAI2 in terms of seedling development and responses to karrikins and strigolactones is poorly defined. Here, we dem- onstrate that HY5 action is genetically separable from that of MAX2 and KAI2. While by5 mutants have weak hypoco- tyl elongation responses to karrikins and the artificial strigolactone GR24, they have normal transcriptional responses, suggesting that HY5 is not involved in perception or action of karrikins or strigolactones. Furthermore, we show that overexpression of KAI2 is sufficient to enhance responses to both karrikins and GR24 in wild-type seedlings, and that KAI2 overexpression partially suppresses the hy5 long hypocotyl phenotype. These results suggest that KAI2 and MAX2 define a regulatory pathway that largely operates independently of HY5 to mediate seedling responses to abiotic signals such as smoke and light.

Karrikin and Cyanohydrin Smoke Signals Provide Clues to New Endogenous Plant Signaling Compounds

Two new types of signaling compounds have been discovered in wildfire smoke due to their ability to stimulate seed germination. The first discovered were karrikins, which share some structural similarity with the strigol- actone class of plant hormones, and both signal through a common F-box protein. However, karrikins and strigolactones operate through otherwise distinct signaling pathways, each distinguished by a specific a/13 hydrolase protein. Genetic analysis suggests that plants contain endogenous compounds that signal specifically through the karrikin pathway. The other active compounds discovered in smoke are cyanohydrins that release germination-stimulating cyanide upon hydrolysis. Cyanohydrins occur widely in plants and have a role in defense against other organisms, but an additional role in endogenous cyanide signaling should also now be considered.