Soil salinization and a degraded ecological environment are challenging agricultural productivity and food security. Rice (Oryza sativa), the staple food of much of the world’s population, is categorized as a salt-susceptible crop. Improving the salt tolerance of rice would increase the potential of saline-alkali land and ensure food security. Salt tolerance is a complex quantitative trait. Biotechnological efforts to improve the salt tolerance of rice hinge on a detailed understanding of the molecular mechanisms underlying salt stress tolerance. In this review, we summarize progress in the breeding of salt-tolerant rice and in the mapping and cloning of genes and quantitative trait loci (QTLs) associated with salt tolerance in rice. Furthermore, we describe biotechnological tools that can be used to cultivate salt-tolerant rice, providing a reference for efforts aimed at rapidly and precisely cultivating salt-tolerance rice varieties.

Summary: In early seedlings, the primary root adapts rapidly to environmental changes through the modulation of endogenous hormone levels. The phytohormone ethylene inhibits primary root elongation, but the underlying molecular mechanism of how ethylene‐reduced root growth is modulated in environmental changes remains poorly understood. Here, we show that a novel rice (Oryza sativa)DOFtranscription factor OsDOF15 positively regulates primary root elongation by regulating cell proliferation in the root meristem, via restricting ethylene biosynthesis. Loss‐of‐function ofOsDOF15impaired primary root elongation and cell proliferation in the root meristem, whereasOsDOF15overexpression enhanced these processes, indicating that OsDOF15 is a key regulator of primary root elongation. This regulation involves the direct interaction of OsDOF15 with the promoter ofOsACS1, resulting in the repression of ethylene biosynthesis. The control of ethylene biosynthesis by OsDOF15 in turn regulates cell proliferation in the root meristem.OsDOF15transcription is repressed by salt stress, and OsDOF15‐mediated ethylene biosynthesis plays a role in inhibition of primary root elongation by salt stress. Thus, our data reveal how the ethylene‐inhibited primary root elongation is finely controlled by OsDOF15 in response to environmental signal, a novel mechanism of plants responding to salt stress and transmitting the information to ethylene biosynthesis to restrict root elongation.

Rice (Oryza sativa) germination and seedling establishment, particularly in increasingly saline soils, are critical to ensure successful crop yields. Seed vigor, which determines germination and seedling growth, is a complex trait affected by exogenous (environmental) and endogenous (hormonal) factors. Here, we used genetic and biochemical analyses to uncover the role of an APETALA2-type transcription factor, SALT AND ABA RESPONSE ERF1 (OsSAE1), as a positive regulator of seed germination and salt tolerance in rice by repressing the expression of ABSCISIC ACID-INSENSITIVE5 (OsABI5). ossae1 knockout lines exhibited delayed seed germination, enhanced sensitivity to abscisic acid (ABA) during germination and in early seedling growth, and reduced seedling salt tolerance. OsSAE1 overexpression lines exhibited the converse phenotype, with increased seed germination and salt tolerance. In vivo and in vitro assays indicated that OsSAE1 binds directly to the promoter of OsABI5, a major downstream component of the ABA signaling pathway and acts as a major regulator of seed germination and stress response. Genetic analyses revealed that OsABI5-mediated ABA signaling functions downstream of OsSAE1. This study provides important insights into OsSAE1 regulation of seed vigor and salt tolerance and facilitates the practical use of OsSAE1 in breeding salt-tolerant varieties suitable for direct seeding cultivation.

Soil compaction represents a major agronomic challenge, inhibiting root elongation and impacting crop yields. Roots use ethylene to sense soil compaction as the restricted air space causes this gaseous signal to accumulate around root tips. Ethylene inhibits root elongation and promotes radial expansion in compacted soil, but its mechanistic basis remains unclear. Here, we report that ethylene promotes abscisic acid (ABA) biosynthesis and cortical cell radial expansion. Rice mutants of ABA biosynthetic genes had attenuated cortical cell radial expansion in compacted soil, leading to better penetration. Soil compaction-induced ethylene also up-regulates the auxin biosynthesis gene OsYUC8 . Mutants lacking OsYUC8 are better able to penetrate compacted soil. The auxin influx transporter OsAUX1 is also required to mobilize auxin from the root tip to the elongation zone during a root compaction response. Moreover, osaux1 mutants penetrate compacted soil better than the wild-type roots and do not exhibit cortical cell radial expansion. We conclude that ethylene uses auxin and ABA as downstream signals to modify rice root cell elongation and radial expansion, causing root tips to swell and reducing their ability to penetrate compacted soil.

Roots are important plant ground organs, which absorb water and nutrients to control plant growth and development. Phytohormones have been known to play a crucial role in the regulation of root growth, such as auxin and ethylene, which are central regulators of this process. Recent findings have revealed that root development and elongation regulated by ethylene are auxin dependent through alterations of auxin biosynthesis, transport and signaling. In this review, we focus on the recent advances in the study of auxin and auxin–ethylene crosstalk in plant root development, demonstrating that auxin and ethylene act synergistically to control primary root and root hair growth, but function antagonistically in lateral root formation. Moreover, ethylene modulates auxin biosynthesis, transport and signaling to fine-tune root growth and development. Thus, this review steps up the understanding of the regulation of auxin and ethylene in root growth.

Rice is an important monocotyledonous crop worldwide; it differs from the dicotyledonous plant Arabidopsis in many aspects. In Arabidopsis, ethylene and auxin act synergistically to regulate root growth and development. However, their interaction in rice is still unclear. Here, we report that the transcriptional activation of OsEIL1 on the expression of YUC8/REIN7 and indole-3-pyruvic acid (IPA)-dependent auxin biosynthesis is required for ethylene-inhibited root elongation. Using an inhibitor of YUC activity, which regulates auxin biosynthesis via the conversion of IPA to indole-3-acetic acid (IAA), we showed that ethylene-inhibited primary root elongation is dependent on YUC-based auxin biosynthesis. By screening phenotypes of seedling primary root from mutagenesis libraries following ethylene treatment, we identified a rice ethylene-insensitive mutant, rein7-1, in which YUC8/REIN7 is truncated at its C-terminus. Mutation in YUC8/REIN7 reduced auxin biosynthesis in rice, while YUC8/REIN7 overexpression enhanced ethylene sensitivity in the roots. Moreover, YUC8/REIN7 catalyzed the conversion of IPA to IAA, truncated version at C-terminal end of the YUC8/REIN7 resulted in significant reduction of enzymatic activity, indicating that YUC8/REIN7 is required for IPA-dependent auxin biosynthesis and ethylene-inhibited root elongation in rice early seedlings. Further investigations indicated that ethylene induced YUC8/REIN7 expression and promoted auxin accumulation in roots. Addition of low concentrations of IAA rescued the ethylene response in the rein7-1, strongly demonstrating that ethylene-inhibited root elongation depends on IPA-dependent auxin biosynthesis. Genetic studies revealed that YUC8/REIN7-mediated auxin biosynthesis functioned downstream of OsEIL1, which directly activated the expression of YUC8/REIN7. Thus, our findings reveal a model of interaction between ethylene and auxin in rice seedling primary root elongation, enhancing our understanding of ethylene signaling in rice. Rice is an important crop worldwide and is grown in water-saturated environments during its life cycle. This unique feature confers that rice might have different aspects from Arabidopsis in ethylene signaling. Although the crosstalk between ethylene and auxin is well understood in Arabidopsis, however, the interaction in rice is largely unclear. Here, we show that YUC8/REIN7, a member of the YUC gene family, catalyzing the conversion of IPA to IAA in auxin biosynthesis, is transcriptionally modulated by ethylene signaling component OsEIL1, and mainly participates in auxin biosynthesis and ethylene-inhibited root growth. We first identified that ethylene-inhibited root elongation is suppressed by the inhibitor of YUC activity, and YUC8/REIN7 is required for IPA-dependent auxin biosynthesis, indicating that YUC8/REIN7 is involved in ethylene-inhibited root elongation in rice early seedlings. Moreover, ethylene induced YUC8/REIN7 transcription and promoted auxin accumulation in roots. Addition of low concentrations of IAA rescued the ethylene response in the rein7-1, demonstrating that ethylene stimulates auxin biosynthesis dependent on YUC8/REIN7 function. Further evidence revealed that OsEIL1 transcriptionally activates the expression of YUC8/REIN7, and YUC8/REIN7-mediated auxin biosynthesis genetically acts downstream of OsEIL1. Our data in the present report identified an interaction between ethylene and auxin in rice seedling primary root elongation, increasing our understanding of ethylene signaling in rice root growth.