ABSTRACT
Numerous links have been reported between immune response and DNA damage repair pathways in both plants and animals but the precise nature of the relationship between these fundamental processes is not entirely clear. Here, we report that XAP5 CIRCADIAN TIMEKEEPER (XCT), a protein highly conserved across eukaryotes, acts as a negative regulator of immunity in Arabidopsis thaliana and plays a positive role in responses to DNA damaging radiation. We find xct mutants have enhanced resistance to infection by a virulent bacterial pathogen, Pseudomonas syringae pv. tomato DC3000, and are hyper-responsive to the defense-activating hormone salicylic acid (SA) when compared to wild-type. Unlike most mutants with constitutive effector-triggered immunity (ETI), xct plants do not have increased levels of SA and retain enhanced immunity at elevated temperatures. Genetic analysis indicates XCT acts independently of NONEXPRESSOR OF PATHOGENESIS RELATED GENES1 (NPR1), which encodes a known SA receptor. Since DNA damage has been reported to potentiate immune responses, we next investigated the DNA damage response in our mutants. We found xct seedlings to be hypersensitive to UV-C and γ radiation and deficient in phosphorylation of the histone variant H2A.X, one of the earliest known responses to DNA damage. These data demonstrate that loss of XCT causes a defect in an early step of the DNA damage response pathway. Together, our data suggest that alterations in DNA damage response pathways may underlie the enhanced immunity seen in xct mutants.
ABSTRACT
The phytohormone ethylene differentially regulates plant architecture and growth in both a light- and nutrient-dependent fashion. The modulation of plant development by ethylene in response to both external and internal signals can also generate tissue-specific differential responses. Here, we report that XAP5 CIRCADIAN TIMEKEEPER (XCT) is involved in blue light-dependent ethylene responses in the aerial tissues of Arabidopsis (Arabidopsis thaliana) seedlings. XCT was first identified as a circadian clock mutant with a short free-running period. The xct mutation also causes sugar-specific hypocotyl growth defects, in which mutants are short in blue light when grown on a sucrose-rich medium but tall when grown on sucrose-deficient medium. Our data suggest that the hypocotyl defects in blue light are not directly caused by defects in clock or light signaling but rather by enhanced ethylene responses. In blue light, xct mutants have a more active ethylene response pathway and exhibit growth phenotypes similar to the constitutive ethylene signaling mutant constitutive triple response1 (ctr1). xct mutants also have reduced ethylene emission, analogous to plants that have lost CTR1 function. Genetic analysis suggests that XCT negatively regulates ethylene responses downstream of ETHYLENE-INSENSITIVE3 in aerial tissues. However, XCT is not required for all ethylene-mediated processes, such as the inhibition of root growth. Thus, XCT acts downstream of a major transcriptional regulator in an organ-specific manner, playing an environment-dependent role in the regulation of plant growth.
