The roots of future rice harvests

The authors thank the Global Rice Science Partnership and Agropolis Fondation (Special grant n° 1400–009 and Rhizopolis grant n° 1001–005) benefiting from a national ANR Investissement d’Avenir” grant ANR-10-LABX-001-01) for supporting the workshop. They acknowledge the assistance of Nathalie Pivot, Cirad and Véronique Rafin, INRA in workshop organization. The root research at Cirad and University of Aberdeen is supported by the European Grant (FP7/2007-2013) under grant agreement n° 289300.27 EURoot “Enhancing resource Uptake from ROOTs under stress in cereal crops”. Research at IRRI is supported by the Generation Challenge Program and the Bill and Melinda Gates Foundation. J.X. is supported by the AcRF Tier 2 grant (MOE2009-T2-1-060) from the Ministry of Education of Singapore and National Research Foundation Singapore under its Competitive Research Programme (CRP Award No. NRF2010 NRF-CRP002-018). Doan Trung Luu is supported by the EU Marie Curie International Outgoing Fellowship 'ORYZAQUA – Cell Biology of Rice Aquaporins' (PIOF-GA-2011-300150). AP acknowledges the Generation Challenge Programme funded project “Targeting drought avoidance root traits to enhance rice productivity under water limited environments”. Financial support for A.G. Diedhiou was provided by the Université Cheikh Anta Diop (UCAD, VE12/13, CpVIII-Ar4 ) and GRISP. *This paper is dedicated to the late memory of Pr Ping Wu who passed away in a tragic car accident on June 12th, 2014.Peer reviewedPublisher PD

Molecular mechanism of crown root initiation and the different mechanisms between crown root and radicle in rice

Monocot plants produce numerous adventitious (crown) roots. The plant hormone auxin has positive effects on crown root formation, while cytokinin suppresses it. We have demonstrated that auxin-induced CROWN ROOTLESS5 (CRL5) regulates crown root initiation in rice through the induction of OsRR1, a negative regulator of cytokinin signaling. CRL5 overexpressing calli formed adventitious roots, although CRL5 overexpressing plants did not induce ectopic roots, suggesting that CRL5, which promotes de novo root initiation, might function only in de-differentiated cells. A radicle initiated normally in a crl5 mutant, in spite of the defect in crown root initiation, whereas crown roots, but not a radicle, were produced in a radicleless1 (ral1) mutant. A crl5 ral1 double mutant displayed an additive phenotype, showing that the formation of each root is regulated by different genetic mechanisms in rice

Genotypic Variations in Responses of Lateral Root Development to Transient Moisture Stresses in Rice Cultivars

Soil water regimes under field conditions inevitably tend to fluctuate ranging from drought to waterlogging. Genotypes that adapt better to such changing hydrologic conditions are assumed to have the ability to maintain root system development under such conditions. This study aimed to evaluate the responses of root system development based on lateral root production to transient moisture stresses, and the contribution of the elongation of seminal and nodal root axes and their lateral, root branching, and aerenchyma development in the seminal root axis, to root system development. The seedlings of two aerobic genotypes (UPLRi7 and NSICRc9) and one irrigated-lowland genotype (PSBRc82), and two parental genotypes (Nipponbare and Kasalath) of chromosome segment substitution lines (CSSLs) were grown by hydroponics. The seedlings were exposed to a drought condition by adding polyethylene glycol to the solution for 7 days and then to an O2-deficient stagnant condition for 7 days (drought-to-stagnant condition), or to reverse successive conditions (stagnant-to-drought condition). Under both conditions, the aerobic genotypes showed greater ability to produce lateral roots than the irrigated-lowland genotype. Under the transient stagnant-to-drought condition, the root traits that contributed to greater lateral root production in the aerobic genotypes were faster seminal root elongation that was closely associated with branching of lateral roots, and greater nodal root production. Under transient drought to stagnant condition; these were faster seminal root elongation mediated by higher aerenchyma formation, and greater nodal root production. Kasalath showed much greater ability to produce lateral roots under both transient moisture stress conditions than Nipponbare. This indicates the potential utility of the CSSLs for precise identification of desirable root traits with less genetic confounding

Utilizing Chromosome Segment Substitution Lines (CSSLs) for Evaluation of Root Responses to Transient Moisture Stresses in Rice

Drought and waterlogging that occur sequentially under field conditions are important abiotic stresses affecting plant growth and development. The ability to maintain the root system development during the contrasting moisture stresses may be one of the key traits for plant adaptation. This study aimed to identify the key root traits that contributed to the above ability by comparatively examining the effects of the two moisture stresses in succession on root system development. The chromosome segment substitution lines (CSSLs) from the crosses between the japonica rice cultivar Nipponbare and indica rice cultivar Kasalath were used for precise comparison of root system development. The rice seedlings were grown by hydroponics under a continuously well-aerated condition for 14 days (non-stressed), a drought condition for 7 days followed by an oxygen (O2)-deficient (stagnant) condition for 7 days (drought-to-stagnant, D-S), or a stagnant condition for 7 days followed by drought condition for 7 days (stagnant-to-drought, S-D). CSSL43 and 47 did not show any significant differences in growth from Nipponbare under the non-stressed condition, but exhibited greater lateral root production under the stresses. Lateral root production was most closely related to faster seminal root elongation mediated by higher aerenchyma formation in the D-S condition, and to more branching of lateral roots on the seminal root axis in the S-D condition. The D-S condition severely affected lateral root production due to reduced seminal root elongation and aerenchyma formation. These results confirmed the fact that those root traits previously identified using different cultivars greatly contribute to plant adaptation. Oxygen deficiency preceded by drought (D-S) was more stressful to roots than drought preceded by O2 deficiency (S-D), because drought reduced root aerenchyma formation during the subsequent stagnant condition

A Role for Auxin in Ethylene-Dependent Inducible Aerenchyma Formation in Rice Roots

Internal oxygen diffusion from shoot to root tips is enhanced by the formation of aerenchyma (gas space) in waterlogged soils. Lysigenous aerenchyma is created by programmed cell death and subsequent lysis of the root cortical cells. Rice (Oryza sativa) forms aerenchyma constitutively under aerobic conditions and increases its formation under oxygen-deficient conditions. Recently, we have demonstrated that constitutive aerenchyma formation is regulated by auxin signaling mediated by Auxin/indole-3-acetic acid protein (AUX/IAA; IAA). While ethylene is involved in inducible aerenchyma formation, the relationship of auxin and ethylene during aerenchyma formation remains unclear. Here, we examined the effects of oxygen deficiency and ethylene on aerenchyma formation in the roots of a rice mutant (iaa13) in which auxin signaling is suppressed by a mutation in the degradation domain of IAA13 protein. The results showed that AUX/IAA-mediated auxin signaling contributes to ethylene-dependent inducible aerenchyma formation in rice roots. An auxin transport inhibitor abolished aerenchyma formation under oxygen-deficient conditions and reduced the expression of genes encoding ethylene biosynthesis enzymes, further supporting the idea that auxin is involved in ethylene-dependent inducible aerenchyma formation. Based on these studies, we propose a mechanism that underlies the relationship between auxin and ethylene during inducible aerenchyma formation in rice roots

Mechanical Stimulus-Sensitive Mutation, rrl3Affects the Cell Production Process in the Root Meristematic Zone in Rice

Genetic studies on the response of plant root to environmental stimuli are important for elucidating the mechanism of the stress tolerance of plants. We isolated and characterized a recessive rice mutant, rrl3,which was highly sensitive to mechanical stimulus and has short roots. No significant difference was observed between the seminal roots of rrl3mutant and wild type in the mean axial and radial length of mature cortical cells. On the other hand, meristematic zone of the root was smaller and the cortical cell flux in the growing zone of the root was significantly lower in the mutant than in the wild type. In addition, the rrl3mutant and the wild type did not differ in sensitivity to ethylene, IAA or ABA. These results suggest that the RRL3gene specifically regulates the cell production process in the root meristematic zone under a mechanically impeded condition and does not regulate the sensitivities to ethylene, IAA and ABA