Dark blue boxes mark MITE-like elements. Molecular functions of D and ANAC074. S6B, ANAC074pro::ANAC074-GFP) restored cell death in the pith parenchyma of anac074 inflorescence stems (Fig. We also investigated the effect of expressing the rice D homolog, OsD (SI Appendix, Fig. âµ5Present address: Department of Human Life Science and Design, Faculty of Contemporary Human Life Science, Tokyo Kasei Gakuin University, Machida, Tokyo 194-0292, Japan. OsD has 70% identity (76% similarity) with D and 41% identity (50% similarity) with ANAC074. 5A), similar to the observations for wild-type cells, and did not increase the percentage of cells lacking plastids and nuclei (SI Appendix, Fig. S4). By contrast, juicy-stem varieties have nonfunctional D, do not induce PCD in the stem pith parenchyma, and retain water within protoplasts of pith parenchyma cells, leading to the juicy-stem trait. 7). Identifying D as the gene involved in programmed death of plant pith parenchyma cells will provide an approach to breeding crops for sugar and ethanol production. Our transactivation assay in yeast cells showed that D or its C-terminal half fused with the GAL4 DNA-binding domain (GAL4 BD) activated the expression of reporter genes (Fig. Second, VNS subfamily proteins are conserved from mosses to flowering plants and are involved in the differentiation of water-conductive and supporting tissues such as hydroid cells, xylem vessels, and xylem fibers (31). S5A), xylem of 65-d-old hypocotyls (SI Appendix, Fig. In animals, the bulk of functional cells in any organ is called the “parenchyma.” This distinguishes the cells which perform the organ’s primary function from “structural” … S8, dark ocher squares), suggesting that these cells had undergone PCD. These cells are especially important for producing sugar and ethanol from the sugar juice of grass stems. Error bars show SD (n = 3). The percentage of cells lacking plastids and nuclei reached 55, 24, and 33% at 48 h after estrogen addition to LexA::D, LexA::ANAC074, and LexA::OsD cell lines, respectively (SI Appendix, Fig. Even Arabidopsis NAC1, which is phylogenetically closest to the D subfamily, does not induce ectopic PCD in Arabidopsis culture cells (Fig. Vascular plant stems contain three main types of tissues: the outermost dermal tissue protects the internal tissues, vascular tissue transports nutrients and water, and ground tissue stores nutrients and water. Thus, among these functionally differentiated group II NAC subfamily proteins, so far, only D subfamily proteinsâat least those in sorghum, Arabidopsis, and riceâhave been shown to have the ability to induce PCD. 3A). (B) Fine mapping of the D locus. Blue vertical bars show molecular marker positions. Analysis of the SKS and MS3B BAC clones and reference genome sequences corresponding to the 18.99-kb region identified only one gene, Sobic.006G147400 (Fig. âµ6Present address: Department of Bioscience, Tokyo University of Agriculture, Tokyo 156-8502, Japan. By contrast, at 48 h after estrogen addition, many of the cells expressing LexA::D, LexA::ANAC074, and LexA::OsD lacked nuclei and plastids (Fig. The unifying characteristic of all parenchyma cells is that they are living at maturity and capable of cell division, giving them important roles in regeneration and wound healing. Arrowheads indicate cells with violet staining derived from transcript-specific hybridization. A graduate student makes a call to action. (B) Histochemical staining of Arabidopsis expressing the GUS reporter gene under control of the ANAC074 promoter. Common sorghum varieties include grain sorghum, forage sorghum, biomass sorghum, and sweet sorghum (10). (A) Quantitative RT-PCR analysis of ANAC074 mRNA levels (normalized with respect to UBQ10 mRNA levels) in seedlings, young leaves, mature leaves, flower buds, and flowers in 50- and 65-d-old hypocotyls and in 35-, 50-, and 65-d-old inflorescence stems. (Lower) Growth of yeast cells expressing each effector in the presence or absence of histidine and adenine. These results indicated that functional and nonfunctional D alleles were widely distributed in Asian and African germplasms (SI Appendix, Fig. 6C). Parenchyma cells can function as storage sites for starches, proteins, oils, and so on, and they contribute support to the plant if they are turgid. Furthermore, unlike VNS subfamily proteins, D subfamily proteins do not appear to be involved in the differentiation of xylem vessels because their loss of function does not affect the PCD of stem tracheary elements, at least in sorghum and Arabidopsis. S5B), and xylem and degenerating pith parenchyma of 65-d-old inflorescence stems (Fig. Parenchyma cells are also located in flowers and fruits. Error bars show SD (n = 3, *P < 0.005, **P < 0.0001). These combined results indicate that ANAC074 is the master transcription factor that induces PCD in Arabidopsis. (C) Percentage of Evans blue-stained cells immediately after (0 h) and 48 h after the addition of estrogen. The loss of nuclei and plastids suggests that expression of D, ANAC074, or OsD induced cell death. 1D, Upper). S4), can induce PCD (26). 5C), whereas the percentage of dead cells in these cell lines in the absence of estrogen was â¼10%, which was comparable with that in wild-type cells and LexA::NAC1 cell lines (Fig. Parenchyma is the tissue made up of cells and intercellular spaces that fills the interior of the body of a flatworm, which is an acoelomate. Nucleotide primers used in this study are listed in Dataset S3. Some of the chief function of the parenchyma cells are storage, transporting, gas exchange, protection, photosynthesis, repairing the damaged tissues and in generating other specialised cells. Expression of ANAC074 under the control of the ANAC074 promoter (SI Appendix, Fig. S8, light red, light blue, light green triangles, respectively). Phloem fibres are flexible long cells that make up the soft fibresâ¦. Error bars show SD (n = 3, *P < 0.05, **P < 0.005, ***P < 0.001). Light blue lines show NAC domain-coding regions. (D) Luciferase (LUC)-based transactivation assay in Arabidopsis culture cell protoplasts. 5A, arrowheads). Copyright © 2020 National Academy of Sciences. Arrowheads indicate nuclei. In Arabidopsis culture cells, GFP-fused D or GFP-fused ANAC074 localized to the nucleus before cell death (Fig. At 48 h after estrogen addition, Evans blue stain marked cells lacking nuclei and plastids in LexA::D, LexA::ANAC074, and LexA::OsD cell lines (Fig. The entire function of the parenchyma seems often to be in the filling of the gaps between more specialized cells. D expression during the formation of dry pith parenchyma in sorghum stems. Parenchyma cells are a type of cell found within most plants. Pith parenchyma cells store water in various plant organs. Some of these arose in geographical areas near where sorghum is proposed to have been initially domesticated around 4000â3000 BC (9, 22) (SI Appendix, Fig. (Scale bars: 50 µm in A, B, and D.). Given that cell death is widely observed in pith parenchyma of flowering plant stems, we speculated that D may have a similar role in other flowering plants. General definition and function of parenchyma cells in plants In plants, parenchyma cells (Gr. Thus, these two types of alleles are candidates for ancient nonfunctional D alleles that were selected during the course of sorghum domestication. âµ3Present address: Research and Development Department, EARTHNOTE Co. Ltd., Nago, Okinawa 905-1152, Japan. In this study, we identified a gene, long referred to as D, in a promising energy grass, Sorghum bicolor, that is responsible for reducing stem water content. Yoneda, K.-i.H., F.I., T.M., S.K., and J.-i. This gene contains three exons and encodes a protein with homology to transcription factors containing a plant-specific DNA-binding domain (Fig. (B) Sugar concentration (Brix value) in stem juice of SKS and MS3B at 30 DAH. (C) Stem sugar content of SKS and MS3B at 30 DAH. These results strongly suggest that expression of D, ANAC074, or OsD is sufficient to induce cell death in Arabidopsis cells, but NAC1 expression is not sufficient. 3G). In this scenario, PCD is specifically induced in dry-stem varieties expressing functional D, which reduces their water content and leads to the dry-stem trait. One of the most important functions of parenchyma cells is that of healing and repair. These results indicate that D and ANAC074 have transactivation activity. S4). Details for plant materials, calculation of stem juice content, measurement of sugar concentration (Brix), calculation of stem sugar content, air porosity measurement, map-based cloning, phylogenetic analysis, polymorphism analysis, quantitative RT-PCR, plasmid construction, analysis of amino acid sequence homology, Arabidopsis transformation, in situ hybridization, histochemical GUS staining, microscopic observations, transcriptional activation assay in yeast cells, luciferase-based transactivation assay in Arabidopsis protoplasts, yeast one-hybrid assay, 3â²-tag digital gene expression profiling, gene ontology analysis, statistical analysis, and data availability are described in SI Appendix, Materials and Methods. Living and dead pith parenchyma cells of 65-d-old hypocotyls ( SI Appendix, Fig is located chromosome... Not detected or was much Lower than D expression in each organ 0.0001 ) the functional cells in organ! 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