植物科学最前線 7:220 (2016)(#% $*)ǪǦŁ vťåƽė ƍĭòûòûǵŵûƊƔƎ ŷůûĄĹ §ĕƆƍĭĎŨÀ¥źÌŻȊ Ȋ
Mechanisms controlling lateral root formation in Arabidopsis
Key words: lateral root formation, Arabidopsis thaliana
Tatsuaki Goh1, Hidehiro Fukaki1
1

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植物科学最前線 7:225 (2016)ǨùƀŴbXwuǍĂA@)>:ĸĳA'@ȇMalamy & Benfey, 1997ȉ
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Maizel >-Mrwe.ȉpG]Qw]ȂģǯC:$'ȉŎ¹÷Ƣƶ>Ŏ2)ƀǦ@6(CƥłƂ,ǍĂȉƢƶ*-=,¯ǃ'ȉŎ-3 Ř¡ĚűCĚī@-CŁ>,"ȇMaizelet al, 2011ȉVermeer et al, 2014ȉvon Wangenheim et al, 2016Ȉ ğȉŎƀŷǤƑ-pGdGkwRuM)]puSLqe]wjǎŊȇLavenus et al, 2015ȉVoss et al, 2015Ȉ+*CƤ7ÏB'ȉƢƶ¯ǃȉƢƶĚű-ï¿ȉƢƶ¯¿ȉǨùƀŴCƨÏƂ,ǎŊ@),=%'ȉŎƀŷ-¯ùŗœ,&'ȉ>,ŵǎǡ8)ŅĝA@>8  ŇƓ(.ȉQtG`_T_-ŎĚī-¯ùŗœ,&'Œǔ" ĺđ-ǍĂįš-ǡĉ,=?ȉƢƶsfr>Ŏƞsfr6(-ĐēŴǖ,&'ȉ66+ŗœŁ>,A'"

T. Goh, H. Fukaki-6
BSJ-Review 7:225 (2016)

植物科学最前線 7:226 (2016)ŎĚī.ȉŐů-Üüƀŷ- A"l\rƞ)Ʊ>AȉÜüĚű-µĢŗœȉƢƶ-EG\u[F[F-Šýŗœ,&'ȉ>,ŵǎǡ8)ŅĝA@ 6"ȉŐů.ȉŞ¯;ŌȄ+*àîŶë,ħ'ȉŎĚīCǕƚ'ȉǧ°+Ěű-ŎƞCœƜ@ A.¾)(+Őů,)%'ȉǫǇ+ŷúĬŽ)+%'@ ğȉŶëħƘ)ŎĚī-µĢŗœ)-ǳCŁ>,@),=%'ȉ66+Ŷë,ǧħ"ŎƞœǟC:&Őů-®2)&+@)ŅĝA@?8".',Atkinson, J. A., Rasmussen, A., Traini, R., Voss, U., Sturrock, C., Mooney, S. J., Wells, D. M., & Bennett, M. J. 2014.
Branching out in roots: uncovering form, function, and regulation. Plant Physiol. 166: 538-550.
Bao, Y., Aggarwal, P., Robbins, N. E., Sturrock, C. J., Thompson, M. C., Tan, H. Q., Tham, C., Duan, L., Rodriguez, P. L.,
Vernoux, T., Mooney, S. J., Bennett, M. J., & Dinneny, J. R. 2014. Plant roots use a patterning mechanism to position
lateral root branches toward available water. Proc. Natl. Acad. Sci. USA 111: 9319-9324.
Barlow, P. W., & Adam, J. S. 1988. The Position and Growth of Lateral Roots on Cultured Root Axes of Tomato,
Lycopersicon-Esculentum (Solanaceae). Plant Systematics and Evolution 158: 141-154.
Berckmans, B., Vassileva, V., Schmid, S. P., Maes, S., Parizot, B., Naramoto, S., Magyar, Z., Alvim Kamei, C. L., Koncz, C.,
Bogre, L., Persiau, G., De Jaeger, G., Friml, J., Simon, R., Beeckman, T., & De Veylder, L. 2011. Auxin-dependent cell
cycle reactivation through transcriptional regulation of Arabidopsis E2Fa by lateral organ boundary proteins. Plant Cell
23: 3671-3683.
Charlton, W. A. 1983. Patterns of Distribution of Lateral Root Primordia. Annals of Botany 51: 417-427.
Charlton, W. A. (1996). Lateral Root Initiation. In Plant Roots: The Hidden Half, 2nd edn. (ed. Y. Waisel, A. Eshel & U.
Kafkafi), pp. 149-173. New York: Marcel Dekker Inc.
De Rybel, B., Audenaert, D., Xuan, W., Overvoorde, P., Strader, L. C., Kepinski, S., Hoye, R., Brisbois, R., Parizot, B.,
Vanneste, S., Liu, X., Gilday, A., Graham, I. A., Nguyen, L., Jansen, L., Njo, M. F., Inze, D., Bartel, B., & Beeckman, T.
2012. A role for the root cap in root branching revealed by the non-auxin probe naxillin. Nat Chem Biol 8: 798-805.
De Rybel, B., Vassileva, V., Parizot, B., Demeulenaere, M., Grunewald, W., Audenaert, D., Van Campenhout, J., Overvoorde,P., Jansen, L., Vanneste, S., Möller, B., Wilson, M., Holman, T., Van Isterdael, G., Brunoud, G., Vuylsteke, M., Vernoux,T., De Veylder, L., Inzé, D., Weijers, D., Bennett, M. J., & Beeckman, T. 2010. A novel Aux/IAA28 signaling cascade
activates GATA23-dependent specification of lateral root founder cell identity. Curr. Biol. 20: 1697-1706.
De Smet, I., Tetsumura, T., De Rybel, B., Frey, N. F. D., Laplaze, L., Casimiro, I., Swarup, R., Naudts, M., Vanneste, S.,
Audenaert, D., Inzé, D., Bennett, M. J., & Beeckman, T. 2007. Auxin-dependent regulation of lateral root positioning in
the basal meristem of Arabidopsis. Development 134: 681-690.
De Smet, I., Vassileva, V., De Rybel, B., Levesque, M. P., Grunewald, W., Van Damme, D., Van Noorden, G., Naudts, M.,
Van Isterdael, G., De Clercq, R., Wang, J. Y., Meuli, N., Vanneste, S., Friml, J., Hilson, P., Jürgens, G., Ingram, G. C.,
Inzé, D., Benfey, P. N., & Beeckman, T. 2008. Receptor-like kinase ACR4 restricts formative cell divisions in the
Arabidopsis root. Science 322: 594-597.
Ditengou, F. A., Teale, W. D., Kochersperger, P., Flittner, K. A., Kneuper, I., van der Graaff, E., Nziengui, H., Pinosa, F., Li, X.,
Nitschke, R., Laux, T., & Palme, K. 2008. Mechanical induction of lateral root initiation in Arabidopsis thaliana. Proc.

T. Goh, H. Fukaki-7
BSJ-Review 7:226 (2016)

植物科学最前線 7:227 (2016)

Natl. Acad. Sci. USA 105: 18818-18823.
Dubrovsky, J. G., Sauer, M., Napsucialy-Mendivil, S., Ivanchenko, M. G., Friml, J., Shishkova, S., Celenza, J., & Benková, E.
2008. Auxin acts as a local morphogenetic trigger to specify lateral root founder cells. Proc. Natl. Acad. Sci. USA 105:
8790-8794.
Fukaki, H., Nakao, Y., Okushima, Y., Theologis, A., & Tasaka, M. 2005. Tissue-specific expression of stabilized
SOLITARY-ROOT/IAA14 alters lateral root development in Arabidopsis. Plant J. 44: 382-395.
Fukaki, H., Tameda, S., Masuda, H., & Tasaka, M. 2002. Lateral root formation is blocked by a gain-of-function mutation in
the SOLITARY-ROOT/IAA14 gene of Arabidopsis. Plant J. 29: 153-168.
Goh, T., Joi, S., Mimura, T., & Fukaki, H. 2012. The establishment of asymmetry in Arabidopsis lateral root founder cells isregulated by LBD16/ASL18 and related LBD/ASL proteins. Development 139: 883-893.
Hirota, A., Kato, T., Fukaki, H., Aida, M., & Tasaka, M. 2007. The auxin-regulated AP2/EREBP gene PUCHI is required formorphogenesis in the early lateral root primordium of Arabidopsis. Plant Cell 19: 2156-2168.
Hodge, A., Berta, G., Doussan, C., Merchan, F., & Crespi, M. 2009. Plant root growth, architecture and function. Plant and Soil
321: 153-187.
Inukai, Y., Sakamoto, T., Ueguchi-Tanaka, M., Shibata, Y., Gomi, K., Umemura, I., Hasegawa, Y., Ashikari, M., Kitano, H., &Matsuoka, M. 2005. Crown rootless1, which is essential for crown root formation in rice, is a target of an AUXINRESPONSE FACTOR in auxin signaling. Plant Cell 17: 1387-1396.
Ito, J., Fukaki, H., Onoda, M., Li, L., Li, C., Tasaka, M., & Furutani, M. 2016. Auxin-dependent compositional change inMediator in ARF7- and ARF19-mediated transcription. Proc. Natl. Acad. Sci. USA 113: 6562-6567.
Kircher, S., & Schopfer, P. 2016. Priming and positioning of lateral roots in Arabidopsis. An approach for an integrating
concept. J Exp Bot 67: 1411-1420.
Kumpf, R. P., Shi, C. L., Larrieu, A., Sto, I. M., Butenko, M. A., Peret, B., Riiser, E. S., Bennett, M. J., & Aalen, R. B. 2013.
Floral organ abscission peptide IDA and its HAE/HSL2 receptors control cell separation during lateral root emergence.
Proc. Natl. Acad. Sci. USA 110: 5235-5240.
Laskowski, M., Grieneisen, V., Hofhuis, H., Hove, C., Hogeweg, P., Marée, A., & Scheres, B. 2008. Root System Architecturefrom Coupling Cell Shape to Auxin Transport. PLoS Biol. 6: e307.
Laskowski, M. J., Williams, M. E., Nusbaum, H. C., & Sussex, I. M. 1995. Formation of lateral root meristems is a two-stage
process. Development 121: 3303-3310.
Lavenus, J., Goh, T., Guyomarc'h, S., Hill, K., Lucas, M., Voss, U., Kenobi, K., Wilson, M. H., Farcot, E., Hagen, G., Guilfoyle,
T. J., Fukaki, H., Laplaze, L., & Bennett, M. J. 2015. Inference of the Arabidopsis lateral root gene regulatory networksuggests a bifurcation mechanism that defines primordia flanking and central zones. Plant Cell 27: 1368-1388.
Lavenus, J., Goh, T., Roberts, I., Guyomarc'h, S., Lucas, M., De Smet, I., Fukaki, H., Beeckman, T., Bennett, M., & Laplaze, L.
2013. Lateral root development in Arabidopsis: fifty shades of auxin. Trends Plant Sci. 18: 450-458.
Lucas, M., Guédon, Y., Jay-Allemand, C., Godin, C., & Laplaze, L. 2008. An auxin transport-based model of root branching in
Arabidopsis thaliana. PLoS ONE 3: e3673.
Lucas, M., Kenobi, K., von Wangenheim, D., Vobeta, U., Swarup, K., De Smet, I., Van Damme, D., Lawrence, T., Peret, B.,
Moscardi, E., Barbeau, D., Godin, C., Salt, D., Guyomarc'h, S., Stelzer, E. H., Maizel, A., Laplaze, L., & Bennett, M. J.
2013. Lateral root morphogenesis is dependent on the mechanical properties of the overlaying tissues. Proc. Natl. Acad.

T. Goh, H. Fukaki-8
BSJ-Review 7:227 (2016)

植物科学最前線 7:228 (2016)

Sci. USA 110: 5229-5234.
Maizel, A., von Wangenheim, D., Federici, F., Haseloff, J., & Stelzer, E. H. 2011. High-resolution live imaging of plant growthin near physiological bright conditions using light sheet fluorescence microscopy. Plant J. 68: 377-385.
Malamy, J. E. 2005. Intrinsic and environmental response pathways that regulate root system architecture. Plant Cell Environ
28: 67-77.
Malamy, J. E., & Benfey, P. N. 1997. Organization and cell differentiation in lateral roots of Arabidopsis thaliana.
Development 124: 33-44.
Mallory, T. E., Chiang, S. H., Cutter, E. G., & Gifford, E. M. 1970. Sequence and Pattern of Lateral Root Formation in 5
Selected Species. American Journal of Botany 57: 800-&.
Moreno-Risueno, M. A., Van Norman, J. M., Moreno, A., Zhang, J., Ahnert, S. E., & Benfey, P. N. 2010. Oscillating gene
expression determines competence for periodic Arabidopsis root branching. Science 329: 1306-1311.
Okushima, Y., Fukaki, H., Onoda, M., Theologis, A., & Tasaka, M. 2007. ARF7 and ARF19 regulate lateral root formation via
direct activation of LBD/ASL genes in Arabidopsis. Plant Cell 19: 118-130.
Okushima, Y., Overvoorde, P. J., Arima, K., Alonso, J. M., Chan, A., Chang, C., Ecker, J. R., Hughes, B., Lui, A., Nguyen, D.,
Onodera, C., Quach, H., Smith, A., Yu, G., & Theologis, A. 2005. Functional genomic analysis of the AUXINRESPONSE FACTOR gene family members in Arabidopsis thaliana: unique and overlapping functions of ARF7 andARF19. Plant Cell 17: 444-463.
Richter, G. L., Monshausen, G. B., Krol, A., & Gilroy, S. 2009. Mechanical stimuli modulate lateral root organogenesis. Plant
Physiol. 151: 1855-1866.
Scheres, B., & Laskowski, M. 2016. Root patterning: it takes two to tangle. J Exp Bot 67: 1201-1203.
Swarup, K., Benková, E., Swarup, R., Casimiro, I., Péret, B., Yang, Y., Parry, G., Nielsen, E., De Smet, I., Vanneste, S.,
Levesque, M. P., Carrier, D., James, N., Calvo, V., Ljung, K., Kramer, E., Roberts, R., Graham, N., Marillonnet, S., Patel,
K., Jones, J. D. G., Taylor, C. G., Schachtman, D. P., May, S., Sandberg, G., Benfey, P., Friml, J., Kerr, I., Beeckman, T.,
Laplaze, L., & Bennett, M. J. 2008. The auxin influx carrier LAX3 promotes lateral root emergence. Nat. Cell Biol. 10:
946-954.
Tian, H., De Smet, I., & Ding, Z. 2014a. Shaping a root system: regulating lateral versus primary root growth. Trends Plant Sci.
19: 426-431.
Tian, H., Jia, Y., Niu, T., Yu, Q., & Ding, Z. 2014b. The key players of the primary root growth and development also function
in lateral roots in Arabidopsis. Plant Cell Rep 33: 745-753.
Van Norman, J. M., Xuan, W., Beeckman, T., & Benfey, P. N. 2013. To branch or not to branch: the role of pre-patterning inlateral root formation. Development 140: 4301-4310.
Vanneste, S., De Rybel, B., Beemster, G. T. S., Ljung, K., De Smet, I., Van Isterdael, G., Naudts, M., Iida, R., Gruissem, W.,
Tasaka, M., Inzé, D., Fukaki, H., & Beeckman, T. 2005. Cell cycle progression in the pericycle is not sufficient for
SOLITARY ROOT/IAA14-mediated lateral root initiation in Arabidopsis thaliana. Plant Cell 17: 3035-3050.
Vermeer, J. E., von Wangenheim, D., Barberon, M., Lee, Y., Stelzer, E. H., Maizel, A., & Geldner, N. 2014. A spatialaccommodation by neighboring cells is required for organ initiation in Arabidopsis. Science 343: 178-183.
von Wangenheim, D., Fangerau, J., Schmitz, A., Smith, R. S., Leitte, H., Stelzer, E. H., & Maizel, A. 2016. Rules andSelf-Organizing Properties of Post-embryonic Plant Organ Cell Division Patterns. Curr. Biol. 26: 439-449.

T. Goh, H. Fukaki-9
BSJ-Review 7:228 (2016)

植物科学最前線 7:229 (2016)

Voss, U., Wilson, M. H., Kenobi, K., Gould, P. D., Robertson, F. C., Peer, W. A., Lucas, M., Swarup, K., Casimiro, I., Holman,
T. J., Wells, D. M., Peret, B., Goh, T., Fukaki, H., Hodgman, T. C., Laplaze, L., Halliday, K. J., Ljung, K., Murphy, A. S.,
Hall, A. J., Webb, A. A., & Bennett, M. J. 2015. The circadian clock rephases during lateral root organ initiation in
Arabidopsis thaliana. Nat. Commun. 6: 7641.
Xuan, W., Audenaert, D., Parizot, B., Moller, B. K., Njo, M. F., De Rybel, B., De Rop, G., Van Isterdael, G., Mahonen, A. P.,
Vanneste, S., & Beeckman, T. 2015. Root Cap-Derived Auxin Pre-patterns the Longitudinal Axis of the Arabidopsis Root.
Curr. Biol. 25: 1381-1388.
Xuan, W., Band, L. R., Kumpf, R. P., Van Damme, D., Parizot, B., De Rop, G., Opdenacker, D., Moller, B. K., Skorzinski, N.,
Njo, M. F., De Rybel, B., Audenaert, D., Nowack, M. K., Vanneste, S., & Beeckman, T. 2016. Cyclic programmed cell
death stimulates hormone signaling and root development in Arabidopsis. Science 351: 384-387.

T. Goh, H. Fukaki-10
BSJ-Review 7:229 (2016)