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How do wounded plants repair or regenerate? Key words: regenerative capacity, repair, wounding

Momoko Ikeuchi, Akira Iwase, Keiko Sugimoto RIKEN Center for Sustainable Resource Science

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M. Ikeuchi, A. Iwase & K. Sugimoto-2

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for root gravitropism13,14. By one day post cut, Lugol staining confirmedde novo starch accumulation above the cut site (Fig. 2a andSupplementary Fig. 4). More intense staining was observed two dayspost cut (Fig. 2a and Supplementary Fig. 4). To test for recovery ofcolumella function, we subjected regenerating roots to a standard gravi-tropism assay by reorienting them perpendicularly to the gravity vectorand scoring the response over time. All wild-type roots showed a cleargravitropic response within 12 h. Although cut roots did not respond togravity in the first 12 h after excision when cut at 130mm, 13.8% of thecut roots exhibited a clear gravitropic response at 1 day post cut, 55.4%at 2 days post cut and 89.2% at 3 days post cut (n 5 65, for all timepoints). However, the quiescent-centre-specific marker WUSCHELRELATED HOMEOBOX 5 (WOX5) was either ectopically expressedin the endodermal file or, at times, expressed in differentiated columellacells at one day post cut (Fig. 2a and Supplementary Fig. 4). Thus, asearly as one day after complete columella excision, a new set of cellsexpressed columella markers and performed columella-specific func-tions while the morphology of the stem cell niche had not yet recovered.

Given the early re-establishment of a differentiated cell type, wetested the requirement for functional stem cells by using mutants inwhich post-embryonic root growth ceases due to the failure to main-tain the stem cell niche. The PLETHORA (PLT) gene family has beenshown to be critical for root formation15, with the double mutantplt1plt2 showing differentiation of stem cells at three days post ger-mination5, as verified under our conditions (Fig. 2b, note the lack ofstem cell layer between the quiescent centre and the starch-stainedcolumella). The uncut double mutant root has abnormal tip and stemcell niche morphology but normal gravitropism and convergent lon-gitudinal cell files5. Surprisingly, plt1plt2 roots cut at four days post

germination quickly regenerated by re-establishing the U-shaped con-vergent pattern of longitudinal cell files at the tip (Fig. 2c, e andSupplementary Fig. 5a). Moreover, starch granules accumulated inthe regenerating double mutants (Fig. 2d) and the gravitropic responsewas re-established (Fig. 2c and Supplementary Fig. 6), indicating thatfunctional columella cells were re-specified during regeneration.Similarly, scarecrow (scr) mutants, which fail to maintain root stem cellfunction through a pathway independent of PLT1 and PLT2 (refs 5, 16,17), were also able to restore their pre-cut pattern, starch staining andgravitropism (Fig. 2b–d, f and Supplementary Figs 5b and 6). PLT1 andPLT2 are expressed early in regeneration in wild-type roots (Supple-mentary Fig. 7). However, using microarray comparison of plt1plt2mutant and wild-type roots, we ruled out that PLT1- and PLT2-dependent genes were induced by alternative mechanisms in regenerat-ing double mutants (Supplementary Fig. 8). We note that a lowerpercentage of plt1plt2 and scr mutants regenerated compared to wild-type roots (Fig. 2c), which we hypothesize is due to the documentedeffect of both mutants in reducing cell divisions in the meristematiczone15,17—the pool of cells recruited for regeneration. Together, theseresults show that stem cell niche activity is not necessary for early root-tip regeneration and imply the existence of an independent mechanismfor cell specification and patterning in the meristematic region.

Several results suggest that auxin, which has been shown to positionthe root stem cell niche and to form a potentially instructive concen-tration gradient18,19, may be a critical component of the mechanism thatcoordinates organogenesis20. First, roots failed to regenerate beyond theearliest stages when we blocked auxin transport during regenerationusing N-1-naphthylphthalamic acid (NPA; Supplementary Fig. 9).Second, auxin efflux carriers and an auxin-responsive reporter

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Figure 1 | Root-tip regeneration and cell fate re-specification in wild type. a, Schematic ofArabidopsis root apical meristem with quiescentcentre (70 mm) and standard excision point(130mm) positions; LRC, lateral root cap.b, Regeneration frequency in wild type (Col-0);n 5 102 (70 mm), 57 (130mm), 111 (200mm), 32(270mm); error bars, standard error of theestimate of the proportion (Methods).c–e, Confocal time-lapse of single regeneratingroots in CYCB1;1::GFP (c), the columella markerPET111 (d) and the quiescent-centre-specificpromoter fusion pWOX5::GFP(ER) (e), atconsecutive days post cut (d.p.c.); scale bars,50mm. f, Expression levels of columella-enriched(n 5 103, top panels) and quiescent-centre-enriched (n 5 95, bottom panels) transcriptsduring regeneration; left, expression in cell typesof uncut roots9–11; right, expression in uncut tipsor regenerating stumps at the time pointsindicated.

NATURE | Vol 457 | 26 February 2009 LETTERS

1151 Macmillan Publishers Limited. All rights reserved©2009

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