SummaryThe Arabidopsis trichome is a model system for studying cell development, cell differentiation and the cell cycle in plants. Our previous studies have shown that the ZINC FINGER PRO-TEIN5 (ZFP5) controls shoot maturation and epidermal cell fate through GA signaling in Arabidopsis.We have identified a novel C2H2 zinc finger protein ZINC FINGER PROTEIN 6 (ZFP6) which plays a key role in regulating trichome development in Arabidopsis.Overexpression of ZFP6 results in ectopic trichomes on carpels and other inflorescence organs. Gain-and loss-of-function analyses have shown that the zfp6 mutant exhibits a reduced number of trichomes in sepals of flowers, cauline leaves, lateral branch and main inflorescence stems in comparison to wild-type plants.Molecular and genetic analyses suggest that ZFP6 functions upstream of GIS, GIS2, ZFP8, ZFP5 and key trichome initiation regulators GL1 and GL3.We reveal that ZFP6 and ZFP5 mediate the regulation of trichome initiation by integrating GA and cytokinin signaling in Arabidopsis. These findings provide new insights into the molecular mechanism of plant hormone control of epidermal trichome patterning through C2H2 transcriptional factors.

Arabidopsis (Arabidopsis thaliana) trichome development is a model system for studying cell development, cell differentiation, and the cell cycle. Our previous studies have shown that the GLABROUS INFLORESCENCE STEMS (GIS) family genes, GIS, GIS2, and ZINC FINGER PROTEIN8 (ZFP8), control shoot maturation and epidermal cell fate by integrating gibberellins (GAs) and cytokinin signaling in Arabidopsis. Here, we show that a new C2H2 zinc finger protein, ZFP5, plays an important role in controlling trichome cell development through GA signaling. Overexpression of ZFP5 results in the formation of ectopic trichomes on carpels and other inflorescence organs. zfp5 loss-of-function mutants exhibit a reduced number of trichomes on sepals, cauline leaves, paraclades, and main inflorescence stems in comparison with wild-type plants. More importantly, it is found that ZFP5 mediates the regulation of trichome initiation by GAs. These results are consistent with ZFP5 expression patterns and the regional influence of GA on trichome initiation. The molecular analyses suggest that ZFP5 functions upstream of GIS, GIS2, ZFP8, and the key trichome initiation regulators GLABROUS1 (GL1) and GL3. Using a steroid-inducible activation of ZFP5 and chromatin immunoprecipitation experiments, we further demonstrate that ZFP8 is the direct target of ZFP5 in controlling epidermal cell differentiation.Cell differentiation and morphogenesis at appropriate times and places are critical for the normal development of multicellular organisms (Ishida et al., 2008).

Phosphoenolpyruvate carboxylase (PEPC) is a crucial enzyme that catalyzes an irreversible primary metabolic reaction in plants. Previous studies have used transgenic plants expressing ectopic PEPC forms with diminished feedback inhibition to examine the role of PEPC in carbon and nitrogen metabolism. To date, the in vivo role of PEPC in carbon and nitrogen metabolism has not been analyzed in plants. In this study, we examined the role of PEPC in plants, demonstrating that PPC1 and PPC2 were highly expressed genes encoding PEPC in Arabidopsis (Arabidopsis thaliana) leaves and that PPC1 and PPC2 accounted for approximately 93% of total PEPC activity in the leaves. A double mutant, ppc1/ppc2, was constructed that exhibited a severe growth-arrest phenotype. The ppc1/ppc2 mutant accumulated more starch and sucrose than wild-type plants when seedlings were grown under normal conditions. Physiological and metabolic analysis revealed that decreased PEPC activity in the ppc1/ppc2 mutant greatly reduced the synthesis of malate and citrate and severely suppressed ammonium assimilation. Furthermore, nitrate levels in the ppc1/ppc2 mutant were significantly lower than those in wild-type plants due to the suppression of ammonium assimilation. Interestingly, starch and sucrose accumulation could be prevented and nitrate levels could be maintained by supplying the ppc1/ppc2 mutant with exogenous malate and glutamate, suggesting that low nitrogen status resulted in the alteration of carbon metabolism and prompted the accumulation of starch and sucrose in the ppc1/ppc2 mutant. Our results demonstrate that PEPC in leaves plays a crucial role in modulating the balance of carbon and nitrogen metabolism in Arabidopsis.