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A study on adaxial-abaxial polarity-dependent control of vascular stem cell fates

NURANI, Alif Meem 東京大学 DOI:10.15083/0002004758

2022.06.22

概要

Cell division and differentiation are fundamental for constructing the complex body plan of living organisms. These processes are carried out by a population of stem cells with the capacity of selfrenewal as well as transformation to other specialized cell types. Plant vascular stem cells are recognized for their infinite proliferative capacity, causing the girth of a tree bark to grow thicker over a lifetime. Moreover, the immobility of plant stem cells signifies that cell fate decision must occur before the differentiation process, to locate the daughter cells in their functional position. My research aims to elucidate how vascular stem cells in plants choose their cell fate to develop xylem vessels elements that conduct water and phloem sieve elements for transportation of photoassimilates. I used a culture system named as in vitro Vascular cell Induction culture System Using Arabidopsis Leaves (VISUAL) in which mesophyll cells in Arabidopsis cotyledons reprogram into vascular stem cells, subsequently inducing trans-differentiation of ectopic xylem vessel element-like cells and phloem sieve element-like cells. First, I carried out a chemical screen in VISUAL and successfully isolated a secondary cell wall fluorescent probe BF-170, that allows high-resolution depth imaging of in vitro and in situ xylem development. Next, I used BF-170 to monitor the three-dimensional vascular differentiation pattern in VISUAL. I discovered that ectopic xylem cells were predominantly transdifferentiated from mesophyll cells on the adaxial side. On the contrary, ectopic phloem cells visualized by phloem markers are mostly generated from abaxial mesophyll cells. This suggests that adaxial-abaxial polarity of leaves may be involved during cell fate decision of vascular stem cells. To test this, I further analyzed the effect of abaxial polarity mutants on the xylem differentiation pattern in VISUAL. Among the mutants, YABBY gene family mutant (yab3-12) displayed enhanced xylem differentiation on the abaxial side, indicating that YAB3 may repress xylem and promote phloem cell identity on the abaxial domains in VISUAL. My results using in vitro vascular differentiation system was also supported by the influence of YABBY family genes during secondary growth of vascular tissues in vivo. Based on these results, I discussed the possibility that spatial cues may be relayed by such abaxial mediators to assist in the correct cell fate decision of vascular stem cells. To analyze how adaxial-abaxial polarity cues affect vascular differentiation, I opted for transcriptome analysis of adaxial and abaxial cells isolated by Laser Capture Microdissection (LCM) before and after 33h induction in VISUAL. Interestingly, the expression of xylem-specific genes was severely diminished on the abaxial side, whereas phloem-specific genes were not differentially expressed between adaxialabaxial sides. This provides novel evidence that the abaxial domains primarily suppress xylem differentiation in VISUAL. In search of abaxial regulators of vascular cell fate, I further identified the CLE41 gene, encoding TDIF peptide, as a potential candidate downstream of YAB3, to be abundantly expressed on the abaxial side. Mutant analysis using its receptor TDR exhibited excessive abaxial xylem differentiation. TDIF-TDR is widely known for its role in inhibiting xylem differentiation. Finally, I propose a concept of such peptide-receptor signaling acting downstream of positional cues to assist in vascular patterning.

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