Ethylene aplenty signals soil compaction: It's tough to drive a spade through compacted soil, and plant roots seem to have the same problem when growing in compacted ground. Pandey et al. found that the problem is not, however, one of physical resistance but rather inhibition of growth through a signaling pathway. The volatile plant hormone ethylene will diffuse through aerated soil, but compacted soil reduces such diffusion, increasing the concentration of ethylene near root tissues. The cellular signaling cascades triggered by too much ethylene stop root growth. Therefore, gaseous diffusion serves as a readout of soil compaction for plant roots growing in search of productive nutrition. Science , this issue p. 276

Root traits such as root angle and hair length influence resource acquisition particularly for immobile nutrients like phosphorus (P). Here, we attempted to modify root angle in rice by disrupting the OsAUX1 auxin influx transporter gene in an effort to improve rice P acquisition efficiency. We show by X-ray microCT imaging that root angle is altered in the osaux1 mutant, causing preferential foraging in the top soil where P normally accumulates, yet surprisingly, P acquisition efficiency does not improve. Through closer investigation, we reveal that OsAUX1 also promotes root hair elongation in response to P limitation. Reporter studies reveal that auxin response increases in the root hair zone in low P environments. We demonstrate that OsAUX1 functions to mobilize auxin from the root apex to the differentiation zone where this signal promotes hair elongation when roots encounter low external P. We conclude that auxin and OsAUX1 play key roles in promoting root foraging for P in rice.

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.

Flag leaf angle impacts the photosynthetic capacity of densely grown plants and is thus an important agronomic breeding trait for crop architecture and yield. The hormone auxin plays a key role in regulating this trait, yet the underlying molecular and cellular mechanisms remain unclear. Here, we report that two rice (Oryza sativa) auxin response factors (ARFs), OsARF6 and OsARF17, which are highly expressed in lamina joint tissues, control flag leaf angle in response to auxin. Loss-of-function double osarf6 osarf17 mutants displayed reduced secondary cell wall levels of lamina joint sclerenchymatous cells (Scs), resulting in an exaggerated flag leaf angle and decreased grain yield under dense planting conditions. Mechanical measurements indicated that the mutant lamina joint tissues were too weak to support the weight of the flag leaf blade, resembling the phenotype of the rice increased leaf angle1 (ila1) mutant. We demonstrate that OsARF6 and OsARF17 directly bind to the ILA1 promoter independently and synergistically to activate its expression. In addition, auxin-induced ILA1 expression was dependent on OsARF6 and OsARF17. Collectively, our study reveals a mechanism that integrates auxin signaling with the secondary cell wall composition to determine flag leaf angle, providing breeding targets in rice, and potentially other cereals, for this key trait.

Primary root growth in cereal crops is fundamental for early establishment of the seedling and grain yield. In young rice (Oryza sativa) seedlings, the primary root grows rapidly for 7–10 days after germination and then stops; however, the underlying mechanism determining primary root growth is unclear. Here, we report that the interplay of ethylene and gibberellin (GA) controls the orchestrated development of the primary root in young rice seedlings. Our analyses advance the knowledge that primary root growth is maintained by higher ethylene production, which lowers bioactive GA contents. Further investigations unraveled that ethylene signaling transcription factor ETHYLENE INSENSITIVE3-LIKE 1 (OsEIL1) activates the expression of the GA metabolism genes GIBBERELLIN 2-OXIDASE 1 (OsGA2ox1), OsGA2ox2, OsGA2ox3, and OsGA2ox5, thereby deactivating GA activity, inhibiting cell proliferation in the root meristem, and ultimately gradually inhibiting primary root growth. Mutation in OsGA2ox3 weakened ethylene-induced GA inactivation and reduced the ethylene sensitivity of the root. Genetic analysis revealed that OsGA2ox3 functions downstream of OsEIL1. Taken together, we identify a molecular pathway impacted by ethylene during primary root elongation in rice and provide insight into the coordination of ethylene and GA signals during root development and seedling establishment.