The circadian clock modulates immune responses in plants and animals; however, it is unclear how host-pathogen interactions affect the clock. Here we analyzed clock function in Arabidopsis thaliana mutants with defective immune responses and found that enhanced disease susceptibility 4 (eds4) displays alterations in several circadian rhythms. Mapping by sequencing revealed that EDS4 encodes the ortholog of NUCLEOPORIN 205, a core component of the inner ring of the nuclear pore complex (NPC). Consistent with the idea that the NPC specifically modulates clock function, we found a strong enrichment in core clock genes, as well as an increased nuclear to total mRNA accumulation, among genes that were differentially expressed in eds4 mutants. Interestingly, infection with Pseudomonas syringae in wild-type (WT) plants downregulated the expression of several morning core clock genes as early as 1 h post-infection, including all members of the NIGHT LIGHT-INDUCIBLE AND CLOCK-REGULATED (LNK) gene family, and this effect was attenuated in eds4. Furthermore, lnk mutants were more susceptible than the WT to P. syringae infection. These results indicate that bacterial infection, acting in part through the NPC, alters core clock gene expression and/or mRNA accumulation in a way that favors bacterial growth and disease susceptibility.

中文翻译：

---

细菌感染破坏时钟基因表达以减弱免疫反应。

生物钟调节植物和动物的免疫反应；然而，尚不清楚宿主-病原体相互作用如何影响时钟。在这里，我们分析了具有缺陷免疫反应的拟南芥突变体的时钟功能，发现增强的疾病易感性 4 (eds4) 显示出几种昼夜节律的改变。测序图谱显示 EDS4 编码 NUCLEOPORIN 205 的直向同源物，NUCLEOPORIN 205 是核孔复合物 (NPC) 内环的核心成分。与 NPC 特异性调节时钟功能的想法一致，我们发现在 eds4 突变体中差异表达的基因中，核心时钟基因有很强的富集，以及核与总 mRNA 积累的增加。有趣的是，早在感染后 1 小时，野生型 (WT) 植物中丁香假单胞菌的感染就下调了几个早晨核心时钟基因的表达，包括夜光诱导和时钟调节 (LNK) 基因家族的所有成员，和这种影响在 eds4 中减弱了。此外，lnk 突变体比 WT 更容易受到丁香假单胞菌感染。这些结果表明，部分通过 NPC 起作用的细菌感染以有利于细菌生长和疾病易感性的方式改变了核心时钟基因表达和/或 mRNA 的积累。