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• W2012240515 abstract "All plant tissue is ultimately derived from the meristems, and the molecular mechanisms that control growth of apical meristems have been widely studied (reviewed in [1Traas J. Bohn-Courseau I. Cell proliferation patterns at the shoot apical meristem.Curr. Opin. Plant Biol. 2005; 8: 587-592Crossref PubMed Scopus (18) Google Scholar, 2Williams L. Fletcher J.C. Stem cell regulation in the Arabidopsis shoot apical meristem.Curr. Opin. Plant Biol. 2005; 8: 582-586Crossref PubMed Scopus (109) Google Scholar]). In contrast, much less attention has been paid to vascular meristems, such as the cambium and procambium, even though these meristems are the source of woody tissue and therefore generate the majority of plant biomass. Although biomass may represent a novel source of renewable energy [3Farrell A.E. Plevin R.J. Turner B.T. Jones A.D. O'Hare M. Kammen D.M. Ethanol can contribute to energy and environmental goals.Science. 2006; 311: 506-508Crossref PubMed Scopus (2073) Google Scholar], little is known about the molecular regulation of vascular-meristem activity. The vascular meristems participate in a highly ordered developmental process with a very prominent polarity. This polarity results in precisely orientated divisions of meristematic initials that generate files of cells, which differentiate into highly specialized and spatially separated xylem and phloem cells (Figure 1A). The factors that are necessary to establish and maintain this polarity remain unknown. This manuscript describes the identification of the pxy mutant in which the spatial organization of vascular development is lost and the xylem and phloem are partially interspersed. The PXY gene encodes for a receptor-like kinase (RLK) that defines a novel role for RLKs in the meristem where it functions to maintain the cell polarity required for the orientation of cell division during vascular development. All plant tissue is ultimately derived from the meristems, and the molecular mechanisms that control growth of apical meristems have been widely studied (reviewed in [1Traas J. Bohn-Courseau I. Cell proliferation patterns at the shoot apical meristem.Curr. Opin. Plant Biol. 2005; 8: 587-592Crossref PubMed Scopus (18) Google Scholar, 2Williams L. Fletcher J.C. Stem cell regulation in the Arabidopsis shoot apical meristem.Curr. Opin. Plant Biol. 2005; 8: 582-586Crossref PubMed Scopus (109) Google Scholar]). In contrast, much less attention has been paid to vascular meristems, such as the cambium and procambium, even though these meristems are the source of woody tissue and therefore generate the majority of plant biomass. Although biomass may represent a novel source of renewable energy [3Farrell A.E. Plevin R.J. Turner B.T. Jones A.D. O'Hare M. Kammen D.M. Ethanol can contribute to energy and environmental goals.Science. 2006; 311: 506-508Crossref PubMed Scopus (2073) Google Scholar], little is known about the molecular regulation of vascular-meristem activity. The vascular meristems participate in a highly ordered developmental process with a very prominent polarity. This polarity results in precisely orientated divisions of meristematic initials that generate files of cells, which differentiate into highly specialized and spatially separated xylem and phloem cells (Figure 1A). The factors that are necessary to establish and maintain this polarity remain unknown. This manuscript describes the identification of the pxy mutant in which the spatial organization of vascular development is lost and the xylem and phloem are partially interspersed. The PXY gene encodes for a receptor-like kinase (RLK) that defines a novel role for RLKs in the meristem where it functions to maintain the cell polarity required for the orientation of cell division during vascular development. Although the highly ordered nature of vascular differentiation is most readily apparent in secondary growth (Figures 1A and 1M), an analogous process occurs during primary vascular-tissue formation. In the Arabidopsis inflorescence stem, files of cells are generated by a coordinated process of orientated cell divisions and differentiation (Figure 1B) that result in vascular bundles forming a characteristic segment shape (Figure 1C). In order to identify factors that contribute to the polarity of vascular tissue, we screened a chemically mutated population of Arabidopsis Landsberg erecta (Ler) plants. By examining hand-cut sections of inflorescence stems, we identified a mutant in which the vascular bundles appeared more flattened around the stem (Figure 1D). Staining sections with Aniline blue, which causes the sieve plates of the phloem to fluoresce yellow under UV light, demonstrated that the phloem appeared to be adjacent to, or interspersed with, the xylem (Figures 1E and 1F). In view of the poor spatial separation of xylem and phloem, the mutant was named phloem intercalated with xylem (pxy). Mutant bundles were significantly narrower in the radial direction and wider in the tangential direction such that the ratio of these two measurements was 1.54 ± 0.04 in the wild-type (WT) and 4.53 ± 0.27 in pxy (p < 0.001). A phenotype was also evident in the organization of vascular bundles in petioles. Wild-type petioles exhibit collateral organization with the xylem and phloem, forming distinct layers that are separated by the procambium (Figure 1G). In pxy petioles, the vascular tissue is less organized (Figure 1H) and resembles the phenotype exhibited in vascular bundles of pxy stems. In contrast, the vascular tissue of roots from pxy seedlings exhibits no obvious phenotypic alterations (Figures 1I and 1J). In longitudinal sections, wild-type xylem vessels are straight, and all vessels in a bundle lie mostly parallel to one another within the plane of the section (Figure 1K). In contrast, pxy xylem vessels are irregularly shaped and move in and out of the plane of the section, and this results in adjacent vessels having different orientations (Figure 1L). Together, these data suggest that both the transverse and longitudinal organization of the vascular tissue is disrupted in pxy stems. The overall plant morphology of pxy is normal except that the inflorescence stem is shorter than the wild-type (Figure S1A in the Supplemental Data available online), a common feature of several mutants that exhibit altered vascular tissue in the stem [4Hanzawa Y. Takahashi T. Michael A.J. Burtin D. Long D. Pineiro M. Coupland G. Komeda Y. ACAULIS5, an Arabidopsis gene required for stem elongation, encodes a spermine synthase.EMBO J. 2000; 19: 4248-4256Crossref PubMed Scopus (216) Google Scholar, 5Parker G. Schofield R. Sundberg B. Turner S. Isolation of COV1, a gene involved in the regulation of vascular patterning in the stem of Arabidopsis.Development. 2003; 130: 2139-2148Crossref PubMed Scopus (53) Google Scholar, 6Pineau C. Freydier A. Ranocha P. Jauneau A. Turner S. Lemonnier G. Renou J.P. Tarkowski P. Sandberg G. Jouanin L. et al.hca: An Arabidopsis mutant exhibiting unusual cambial activity and altered vascular patterning.Plant J. 2005; 44: 271-289Crossref PubMed Scopus (34) Google Scholar], and the underside of the cotyledons is red in colors consistent with anthocyanin accumulation (Figure S1B). Despite the red color of the cotyledons, the pattern of venation was similar to the wild-type (Figure S1C). Similarly there was no obvious alteration in the pattern of leaf venation (data not shown). pxy therefore affects the organization of vascular tissue within a bundle rather than the organization of bundles within an organ. Map-based cloning was used to demonstrate that pxy was caused by a mutation in At5g61480 (Figure 2A), and this mutation resulted in a change from a glutamine to a premature stop codon at position 971, causing the predicted protein sequence to be truncated by 91 amino acids (Figure 2B). A clone containing only At5g61480 entirely complemented the pxy phenotypes and restored both normal vascular-tissue organization and plant growth (Figures 2C and 2D). Reverse genetics was used to identify four additional T-DNA insertions within At5g61480 in the Colombia (Col) background (Figure 2B). These T-DNA insertions also gave the pxy phenotype (Figures S3A–S3D), confirming that it is indeed caused by a mutation in At5g61480 (henceforth referred to as PXY). The predicted protein encoded by the PXY gene contains a putative kinase domain and a predicted extracellular domain with 21 leucine-rich repeats (eLRR) (Figure 2B). PXY falls within subgroup XI of the Arabidopsis receptor kinase-like (RLK) family that all contain putative eLRR domains [7Shiu S.H. Bleecker A.B. Receptor-like kinases from Arabidopsis form a monophyletic gene family related to animal receptor kinases.Proc. Natl. Acad. Sci. USA. 2001; 98: 10763-10768Crossref PubMed Scopus (964) Google Scholar] (Figure S2A). The pxy-2 allele has a T-DNA insertion that results in the loss of the entire predicted kinase domain (Figure 2B). Interestingly, this mutant exhibits a less severe phenotype than pxy-1 (Figures S3B and S3D), suggesting that pxy-1 may act as an antimorphic allele. There are two other putative eLRR kinases (encoded by At1g08590 and At4g28650) that are obviously very closely related to PXY and that we have named PXY-LIKE (PXL1 and PXL2, respectively) (Figure S2A and [7Shiu S.H. Bleecker A.B. Receptor-like kinases from Arabidopsis form a monophyletic gene family related to animal receptor kinases.Proc. Natl. Acad. Sci. USA. 2001; 98: 10763-10768Crossref PubMed Scopus (964) Google Scholar]). Insertion mutants in either pxl1 or pxl2 do not exhibit an obvious phenotype in the stem (Figures S3E and S3F). As judged by broader, flatter vascular bundles and a less clear distinction between xylem and phloem, double-mutant combinations of a Col allele, of pxy (pxy-3) with pxl1 and pxl2, generate a more severe vascular phenotype than pxy-3 alone, suggesting that these genes act synergistically with PXY in regulating vascular-tissue development in the stem (Figures S3C, S3H, and S3I). The triple-mutant combination did not enhance the double-mutant phenotype (data not shown), and in all mutants some polarity appears to be retained within the vascular bundle, suggesting that other factors maybe involved. It became evident during mapping that the pxy phenotype was clearly more pronounced in the Ler background in which it was isolated (Figure S3B; compare to Figures 1C–1F). Ler carries a mutation in the ER gene that also encodes LRR receptor kinase that affects a number of diverse plant developmental processes including organ shape and spacing of stomata [8Nadeau J.A. Sack F.D. Control of stomatal distribution on the Arabidopsis leaf surface.Science. 2002; 296: 1697-1700Crossref PubMed Scopus (366) Google Scholar, 9Shpak E.D. Lakeman M.B. Torii K.U. Dominant-negative receptor uncovers redundancy in the Arabidopsis ERECTA leucine-rich repeat receptor-like kinase signaling pathway that regulates organ shape.Plant Cell. 2003; 15: 1095-1110Crossref PubMed Scopus (164) Google Scholar]. The data presented here suggest that it may have some role in maintaining polarity of vascular tissue. PXY is related to the LRR-RLK CLV1 (see below); in common with PXY, the most severe clv1 alleles are apparently antimorphic and the clv1 phenotype is more evident in the Ler background [10Dievart A. Dalal M. Tax F.E. Lacey A.D. Huttly A. Li J.M. Clark S.E. CLAVATA1 dominant-negative alleles reveal functional overlap between multiple receptor kinases that regulate meristem and organ development.Plant Cell. 2003; 15: 1198-1211Crossref PubMed Scopus (147) Google Scholar]. Subgroup XI of the Arabidopsis LRR-receptor-kinase family also contains CLV1 and BAM1/2/3 (Figure S2A) that respectively regulate proliferation and maintenance of the dividing cells in the shoot apical meristem [11Clark S.E. Williams R.W. Meyerowitz E.M. The CLAVATA1 gene encodes a putative receptor kinase that controls shoot and floral meristem size in Arabidopsis.Cell. 1997; 89: 575-585Abstract Full Text Full Text PDF PubMed Scopus (1008) Google Scholar, 12DeYoung B.J. Bickle K.L. Schrage K.J. Muskett P. Patel K. Clark S.E. The CLAVATA1-related BAM1, BAM2 and BAM3 receptor kinase-like proteins are required for meristem function in Arabidopsis.Plant J. 2006; 45: 1-16Crossref PubMed Scopus (251) Google Scholar]. In order to determine whether pxy might function to regulate cell number in the procambium, we counted the number of cells per vascular bundle. The mean number of the metaxylem cells within each vascular bundle was significantly (p < 0.001) lower in pxy (12 ± 0.8) than in the wild-type (19 ± 0.8). In contrast, the total number of xylem cells (vessels and xylem parenchyma) per vascular bundle was not significantly (p > 0.05) different between wild-type plants (93 ± 4) and pxy (107 ± 4) plants, suggesting that although there is a change in the proportion of different cell types, the total number of cells in the xylem remains similar. These data suggest that pxy is not analogous to the clv1 mutant in which proliferation of stem cells leads to large increases in cell numbers. To further examine the distribution of procambial cells, we transformed wild-type and pxy plants with a construct in which the promoter of the ATHB-15/CORONA gene was used to drive expression of the GUS gene (ATHB15GUS). ATHB-15 encodes a presumed transcription factor and is a member of the class III homeodomain-leucine zipper (HD-Zip III) gene family involved in the regulation of early vascular development [13Ohashi-Ito K. Fukuda H. HD-Zip III homeobox genes that include a novel member, ZeHB-13 (Zinnia)/ATHB-15 (Arabidopsis), are involved in procambium and xylem cell differentiation.Plant Cell Physiol. 2003; 44: 1350-1358Crossref PubMed Scopus (127) Google Scholar, 14Prigge M.J. Otsuga D. Alonso J.M. Ecker J.R. Drews G.N. Clark S.E. Class III homeodomain-leucine zipper gene family members have overlapping, antagonistic, and distinct roles in Arabidopsis development.Plant Cell. 2005; 17: 61-76Crossref PubMed Scopus (521) Google Scholar]. In young vascular bundles, ATHB15GUS forms a relatively broad band of expression between the protoxylem and phloem. As vascular-bundle differentiation proceeds, expression is confined to a narrow band of cells in a manner that exactly matches the undifferentiated procambial cells (Figure 3A). In very old vascular bundles in which primary vascular development has ceased, GUS activity is also detected in the xylem cap cells that are located inside of the protoxylem (Figure S2B). In contrast, pxy plants transformed with the ATHB15 reporter, and GUS activity no longer formed a distinctive layer between the xylem and phloem but appeared interspersed with the xylem (Figure 3A), consistent with the histochemical analysis described above. This result was confirmed with an alternative marker for procambial-cell identity. BRL1 plays a role in regulating procambial-cell number in the stem [15Cano-Delgado A. Yin Y.H. Yu C. Vafeados D. Mora-Garcia S. Cheng J.C. Nam K.H. Li J.M. Chory J. BRL1 and BRL3 are novel brassinosteroid receptors that function in vascular differentiation in Arabidopsis.Development. 2004; 131: 5341-5351Crossref PubMed Scopus (359) Google Scholar]. Results obtained from analyzing plants in which the BRL1 promoter is used to drive the GUS gene (Figure S1D) are very similar to those observed for ATHB15GUS above marker and confirms the persistence of procambial cells throughout vascular development in pxy. Furthermore, the organization of vascular tissue is clearly affected early on during vascular development in pxy (Figure 4B) when relatively few xylem and phloem cells have differentiated and vascular bundles contain a high proportion of procambial cells. Consequently, data from both the cell counts and the ATHB15GUS and BRL1GUS reporter-gene analysis suggest that procambial cells are maintained throughout vascular development, indicating that pxy mutants do not behave like bam1, bam2, and bam3 and that the phenotype is not a result of inappropriate differentiation of procambial cells. So whereas PXY, CLV1, and BAM1, BAM2, and BAM3 are all essential for meristem function and are structurally similar, PXY appears to function as part of a quite different mechanism. Two other more distantly related LRR-RLKs regulate different aspects of vascular development. BRL1 is described above, and a second, VH1, is involved in the transition to a procambial-cell fate [16Clay N.K. Nelson T. VH1, a provascular cell-specific receptor kinase that influences leaf cell patterns in Arabidopsis.Plant Cell. 2002; 14: 2707-2722Crossref PubMed Scopus (106) Google Scholar]. In addition to regulating the orientation of cell division, vascular-cell polarity is also important in maintaining the continuity of the vascular network (reviewed in [17Turner S.R. Gallois P. Brown D. Tracheary element differentiation.Annu. Rev. Plant Biol. 2007; 58: 407-433Crossref PubMed Scopus (166) Google Scholar]). A proteoglycan named xylogen has been identified that promotes xylem-cell differentiation and is secreted from the end of the cell in a polar manner that is required for the continuity of the xylem [18Motose H. Sugiyama M. Fukuda H. A proteoglycan mediates inductive interaction during plant vascular development.Nature. 2004; 429: 873-878Crossref PubMed Scopus (243) Google Scholar].Figure 4Cell Divsions in Procambial Cells of pxy during Vascular DevelopmentShow full caption(A–D) Resin-embedded transverse sections of stem vascular bundles of young (A and B) and old (C and D) stems from wild-type (A and C) and pxy (B and D) plants. The phloem (ph), xylem (xy), and procambium (pc) and recent cell divisions (arrows) are indicated. Scale bars represent 30 μm.(E and F) A close-up picture of part of (A) and (B). Recent divisions of procambial cells, identified as a very thin cell wall dividing two cells, are marked with arrows and in the wild-type are confined to a narrow region close to the phloem (E). Similar divisions are also marked by arrows in pxy; however, arrowheads indicate undifferentiated, apparently recently divided cells, in pxy, that appear more widely distributed within the vascular bundle (F).View Large Image Figure ViewerDownload Hi-res image Download (PPT) (A–D) Resin-embedded transverse sections of stem vascular bundles of young (A and B) and old (C and D) stems from wild-type (A and C) and pxy (B and D) plants. The phloem (ph), xylem (xy), and procambium (pc) and recent cell divisions (arrows) are indicated. Scale bars represent 30 μm. (E and F) A close-up picture of part of (A) and (B). Recent divisions of procambial cells, identified as a very thin cell wall dividing two cells, are marked with arrows and in the wild-type are confined to a narrow region close to the phloem (E). Similar divisions are also marked by arrows in pxy; however, arrowheads indicate undifferentiated, apparently recently divided cells, in pxy, that appear more widely distributed within the vascular bundle (F). The very ordered pattern of vascular development seen in the stems is a result of orientated cell divisions that appear coordinated between adjacent procambial cells. The fact that cell files are no longer clearly identifiable in pxy suggests that the orientation of cell division is lost. Thin sections were used to more carefully examine procambial-cell divisions. In the wild-type, adjacent procambial cells divide in a coordinated manner along a defined division plane, and this results in a band of procambial cells separating the xylem and phloem (Figures 4A, 4C, and 4E). In pxy plants, however, divisions are not localized to a discrete domain. When dividing cells are observed in pxy vascular bundles, they are much more widely distributed within the vascular bundles (Figures 4B, 4D, and 4F). Individual procambial cells frequently occur adjacent to differentiating xylem, and the division plane exhibits no obvious correlation with adjacent cells, the organization of the vascular bundle, or the overall organization of the stem (Figure 4F). Interestingly, although the separation of xylem and phloem is lost in pxy, both tissues still form, suggesting that spatial separation of xylem and phloem is not necessary for their proper differentiation. This finding implies that the fate of procambial cells is controlled by some quite separate mechanism. A number of mutants, such as kan, abv, rev, and phb, that affect the polarity of plant organs have been identified [19Eshed Y. Baum S.F. Perea J.V. Bowman J.L. Establishment of polarity in lateral organs of plants.Curr. Biol. 2001; 11: 1251-1260Abstract Full Text Full Text PDF PubMed Scopus (470) Google Scholar, 20Kerstetter R.A. Bollman K. Taylor R.A. Bomblies K. Poethig R.S. KANADI regulates organ polarity in Arabidopsis.Nature. 2001; 411: 706-709Crossref PubMed Scopus (432) Google Scholar, 21McConnell J.R. Barton M.K. Leaf polarity and meristem formation in Arabidopsis.Development. 1998; 125: 2935-2942PubMed Google Scholar, 22McConnell J.R. Emery J. Eshed Y. Bao N. Bowman J. Barton M.K. Role of PHABULOSA and PHAVOLUTA in determining radial patterning in shoots.Nature. 2001; 411: 709-713Crossref PubMed Scopus (801) Google Scholar]. In some alleles of these mutants, vascular bundles of the stem may exhibit an amphivasal organization in which the xylem surrounds the phloem; however, they still exhibit ordered cell division of the procambium and have spatially separated xylem and phloem [23Emery J.F. Floyd S.K. Alvarez J. Eshed Y. Hawker N.P. Izhaki A. Baum S.F. Bowman J.L. Radial patterning of Arabidopsis shoots by class IIIHD-ZIP and KANADI genes.Curr. Biol. 2003; 13: 1768-1774Abstract Full Text Full Text PDF PubMed Scopus (810) Google Scholar]. These observations suggest that there are two polarity cues required for normal vascular development. One is controlled by genes such as kan, abv, rev, and phb that polarize xylem and phloem over a long distance and reflect the polarity of the organ. The other is shorter range, involves the PXY gene, and operates within the vascular tissue to ensure the segregation of the xylem and phloem by polarized division of the procambial cells. It is possible that loss of orientated cell divisions may result from a disruption to a pathway that acts directly to polarize cells or indirectly by affecting a pathway that is required to set up the original polarity of the vascular bundle. To examine these possibilities, we measured PXY mRNA levels by using real-time quantitative PCR. Although PXY expression was elevated in the pxy mutant, the pattern of expression was essentially the same as the wild-type and occurred throughout the course of stem development (Figure 3B). To gain a more detailed picture of PXY gene expression, we cloned the PXY promoter in front of the GUS reporter gene. PXY was expressed in the vascular tissue of a variety of organs including the veins of leaves and the stele of roots (Figure S2C). In the stem, the pattern of GUS expression was almost identical to that of ATHB15 and appeared to be confined to the procambial cells in developing vascular bundles throughout the course of vascular-bundle development (Figure 3C). This link between PXY and the procambial-specific marker ATHB15 is further strengthened by analysis of publicly-available expression data that show a very strong correlation between the expression of these two genes (data not shown). Consequently, both real-time quantitative PCR and reporter-gene data give an expression pattern that indicates that PXY may not only be required very early in vascular development to initialize the polarity of the vascular bundle but also is continuously required throughout the course of stem vascular-tissue development. PXY is expressed in dividing procambial cells, and given the close predicted domain structure and its similarity to CLV1, it is likely to act as receptor. The data presented here are consistent with a model in which expression of an extracellular ligand in either the adjacent developing xylem or phloem would result in an instructive signal to only one side of the procambial; this signal could be translated into cell-polarity information and used to determine the orientation of cell division (Figure 1M). In plants in which PXY is defective, this polarity information would be lost, resulting in random positioning of the division plane and a concomitant breakdown of the ordered pattern of cell division (Figure 1N). A role for PXY in a signaling pathway that transmits information required to determine the proper cell-division plane during vascular development is consistent with the known function of several plant receptor kinases. For example, SCM is essential for transmitting positional information necessary for the proper cell fate of root epidermal cells [24Kwak S.H. Shen R. Schiefelbein J. Positional signaling mediated by a receptor-like kinase in Arabidopsis.Science. 2005; 307: 1111-1113Crossref PubMed Scopus (163) Google Scholar]. Furthermore, the putative receptor kinase ER and the membrane-bound eLRR containing TMM protein are both essential components of a pathway that influences the division plane of meristemoid cells and regulates the correct spacing of stomata in the pavement epithelial cells of leaves [8Nadeau J.A. Sack F.D. Control of stomatal distribution on the Arabidopsis leaf surface.Science. 2002; 296: 1697-1700Crossref PubMed Scopus (366) Google Scholar, 25Shpak E.D. McAbee J.M. Pillitteri L.J. Torii K.U. Stomatal patterning and differentiation by synergistic interactions of receptor kinases.Science. 2005; 309: 290-293Crossref PubMed Scopus (404) Google Scholar]. A model in which PXY expressed in procambial cells receives information from adjacent tissue would be analogous to the mechanism that controls the division of EMS cells in C. elegans embryogenesis, where the division plane is dependent upon the position of the adjacent P2 cell [26Goldstein B. Cell contacts orient some cell-division axes in the Caenorhabditis-Elegans embryo.J. Cell Biol. 1995; 129: 1071-1080Crossref PubMed Scopus (110) Google Scholar] that confers information via the WNT signaling pathway [27Schlesinger A. Shelton C.A. Maloof J.N. Meneghini M. Bowerman B. Wnt pathway components orient a mitotic spindle in the early Caenorhabditis elegans embryo without requiring gene transcription in the responding cell.Genes Dev. 1999; 13: 2028-2038Crossref PubMed Scopus (177) Google Scholar]. The data presented here demonstrate that PXY, an LRR kinase, plays an essential role in maintaining polarity within the vascular meristems and represents an important step forward in our understanding of this relatively poorly studied type of meristematic tissue. It may also represent a more general pathway for all plant meristems in which LRR kinases function to both regulate the differentiation of stem cells and maintain the polarity that determines the orientation of cell division. The authors wish to thank Keith Brennan, Denis Headon, Ray Wightman, and particularly Joe Ogas for their comments on the manuscript; Frank Bedouin for his contribution to identifying the pxy-1 allele; and Carl Knipe for his help in complementation of the pxy gene. We also thank Dr. Kyoko Ohashi-Ito, Michigan State University for supplying the Athb15:GUS and BRL1:GUS constructs, Nottingham Arabidopsis Stock Centre and Arabidopsis Biological Resource Center for supplying seeds, and the BBSRC for a studentship to K.F. Download .pdf (.79 MB) Help with pdf files Document S1. Experimental Procedures and Three Figures" @default.
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• W2012240515 title "PXY, a Receptor-like Kinase Essential for Maintaining Polarity during Plant Vascular-Tissue Development" @default.
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