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Morphological and gene expression changes in Arabidopsis thaliana under clinorotated conditions;with special reference to WRKY46

Soh Hyuncheol 東北大学

2020.03.04

概要

Gravity is an important environmental factor that controls plant growth and development. However, the mechanistic aspect of plant response to gravity is far from being fully understood at present. Microgravity conditions are now available by utilizing space environment, but spaceflight opportunities for biological research are very limited. On the ground, clinorotation facilities have been used for simulating plant growth in microgravity. Clinorotation experiments could bring about novel results and hypothesis that should be ultimately verified under ‘real microgravity’ conditions in space. In this study, I examined morphology and gene expression changes of Arabidopsis thaliana seedlings under short-term (1 hour) and long-term (1 to 6 days) 3-D clinorotation.

To understand the 3-D clinorotation effect on the gene expression pattern of Arabidopsis roots, using 44k Arabidopsis microarray (Agilent) I analyzed transcript abundance of genes in seedlings 1 h and 6 days after the start of 3-D clinorotation. The microarray data analysis showed that Arabidopsis seedlings respond to 3-D clinorotation by dynamic up- and down-regulations of genes. Gene ontology and qRT- PCR analyses of the genes investigated showed their expression profiles and patterns similar to those observed in stress-challenged plants. To find genes specifically responsive to 3-D clinorotation, I selected 10 genes that were highly expressed under short-term (1 h) 3-D clinorotation, and their expression patterns were validated by qRT- PCR analysis. Nine out of the 10 genes were up-regulated due to not only clinorotation but also touch and/or wounding in 3-week-old Arabidopsis seedlings. However, WRKY46 expression was elevated only by 3-D clinorotation. Also, to clarify the characteristics of the genes expressed at high level in response to 3-D clinorotation, 20 cis-elements in the promoters of the 40 selected genes, including the 10 3-D clinorotation-inducible genes, six WRKY genes, and abiotic stress-inducible genes, were analyzed, and their spatial positions on each promoter were determined. Four cis-elements (M/T-G-T-P from MYB1AT or TATABOX5, GT1CONSENSUS, TATABOX5, and POLASIG1) showed a unique spatial arrangement in most 3-D clinorotation-inducible genes including WRKY46. It was suggested that the M/T-G-T- P cis-element pattern identified in the promoter of WRKY46 could play a role in regulating gene expression due to 3-D clinorotation.

To understand the role of the WRKY46 gene in morphological change under clinorotated conditions, the responses of wrky46 knockout mutants, constructed by T- DNA insertion, to 3-D clinorotation and gravistimulation by reorientation were investigated. Morphological changes in roots, such as multidirectional growth patterns, were observed in the wild type (WT) seedlings from day 3 to day 4 after the start of 3- D clinorotation, while in the wrky46 mutant seedlings the changes observed were minimal. Interestingly, the expressions of genes involved in auxin transport, such as AUX1, PIN2, PIN3, PIN4, PIN7, and ARG1, appeared to be lower in WT than wrky46 mutant seedlings under 3-D clinorotation. In addition, the gravitropic curvature of wrky46 roots at 10 to 15 h following gravistimulation was smaller than that of WT, and it became the same in WT and wrky46 by 20 h after gravistimulation. Because auxin- resistance genes AXRs are known to play a role in gravitropic bending of roots, I analyzed their expressions. The results showed that AXR1, AXR3, and AXR4 tended to be higher in WT seedlings compared to those in wrky46 mutant seedlings after gravistimulation.

Thus, this study showed that roots of Arabidopsis seedlings display multiple- directional growth together with coiling under 3-D clinorotated conditions. These morphological changes due to clinorotation could result from loss of gravitropic response, phenocopying root growth in microgravity. Analyses of gene expression with microarray and qRT-PCR identified a clinorotation-responsive gene WRKY46 that could play a role in the clinorotaion-inducible changes of root growth pattern. The results of functional analysis of WRKY46 suggested its involvement in root bending by gravitropic response or by multi-directional growth during 3-D clinorotation, probably via regulating auxin transport and/or auxin response.

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