Ethylene, an important hormone in plant growth, development and response to environmental stimuli, is rapidly induced by mechanical injury or wounding. Although induction of ACS (1-aminocyclopropane-1-carboxylic acid synthase) gene expression has been associated with this process, the detailed regulatory mechanism is unclear. Here, we report that the wounding-induced ethylene production is modulated by both mitogen-activated protein kinase (MAPK) pathway and calcium-dependent protein kinase (CPK) pathway. Study using acs mutants demonstrated that four ACS isoforms, including ACS2, ACS6, ACS7 and ACS8, contribute to ethylene production in response to wounding. Loss-of-function analysis defines the role of MPK3 and MPK6, and their upstream MKK4 and MKK5, in wounding-induced ethylene production. They play an important role in the wounding-induced upregulation of all four ACS genes expression. Independent of MAPK pathway, CPK5 and CPK6 are also involved in the wounding-induced ethylene production by regulating the expression of ACS2, ACS6 and ACS8 genes. Taken together, we demonstrate that two independent kinase signalling pathways, MPK3/MPK6 cascade and CPK5/CPK6, are involved in the wounding-induced ethylene biosynthesis via differential regulation of ACS genes at transcriptional level.

Signaling pathways that control the activities in non-photosynthetic plastids, important sites of plant metabolism, are largely unknown. Previously, we demonstrated that WRKY2 and WRKY34 transcription factors play an essential role in pollen development downstream of mitogen-activated protein kinase 3 (MPK3) and MPK6 in Arabidopsis. Here, we report that GLUCOSE-6-PHOSPHATE/PHOSPHATE TRANSLOCATOR 1 (GPT1) is a key target gene of WRKY2/WRKY34. GPT1 transports glucose-6-phosphate (Glc6P) into plastids for starch and/or fatty acid biosynthesis depending on the plant species. Loss of function of WRKY2/WRKY34 results in reduced GPT1 expression, and concomitantly, reduced accumulation of lipid bodies in mature pollen, which leads to compromised pollen viability, germination, pollen tube growth, and male transmission in Arabidopsis. Pollen-specific overexpression of GPT1 rescues the pollen defects of wrky2 wrky34 double mutant. Furthermore, gain-of-function activation of MPK3/MPK6 enhances GPT1 expression; whereas GPT1 expression is reduced in mkk4 mkk5 double mutant. Together, this study revealed a cytoplasmic/nuclear signaling pathway capable of coordinating the metabolic activities in plastids. High-level expression of GPT1 at late stages of pollen development drives Glc6P from cytosol into plastids, where Glc6P is used for fatty acid biosynthesis, an important step of lipid body biogenesis. The accumulation of lipid bodies during pollen maturation is essential to pollen fitness and successful reproduction.

Gamma-aminobutyric acid (GABA) is an important metabolite which functions in plant growth, development, and stress responses. However, its role in plant defense and how it is regulated are largely unknown. Here, we report a detailed analysis of GABA induction during the resistance response to Pseudomonas syringae in Arabidopsis thaliana. While searching for the mechanism underlying the pathogen-responsive mitogenactivated protein kinase (MPK)3/MPK6 signaling cascade in plant immunity, we found that activation of MPK3/ MPK6 greatly induced GABA biosynthesis, which is dependent on the glutamate decarboxylase genes GAD1 and GAD4. Inoculation with Pseudomonas syringae pv tomato DC3000 (Pst) and Pst-avrRpt2 expressing the avrRpt2 effector gene induced GAD1 and GAD4 gene expression and increased the levels of GABA. Genetic evidence revealed that GAD1, GAD2, and GAD4 play important roles in both GABA biosynthesis and plant resistance in response to Pst-avrRpt2 infection. The gad1/2/4 triple and gad1/2/4/5 quadruple mutants, in which the GABA levels were extremely low, were more susceptible to both Pst and Pst-avrRpt2. Functional loss of MPK3/MPK6, or their upstream MKK4/ MKK5, or their downstream substrate WRKY33 suppressed the induction of GAD1 and GAD4 expression after Pst-avrRpt2 treatment. Our findings shed light on both the regulation and role of GABA in the plant immunity to a bacterial pathogen.

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