Ethylene-mediated reactive oxygen species signalling is involved in adaptive responses of wheat seedlings to waterlogged conditions through controlling formation of lysigenous aerenchyma and expression of genes encoding ethanol fermentation enzymes in roots

A radial oxygen loss (ROL) barrier in roots of waterlogging-tolerant plants promotes oxygen movement via aerenchyma to the root tip, and impedes soil phytotoxin entry. The molecular mechanism and genetic regulation of ROL barrier formation are largely unknown. Zea nicaraguensis, a waterlogging-tolerant wild relative of maize (Zea mays ssp. mays), forms a tight ROL barrier in its roots when waterlogged. We used Z. nicaraguensis chromosome segment introgression lines (ILs) in maize (inbred line Mi29) to elucidate the chromosomal region involved in regulating root ROL barrier formation. A segment of the short-arm of chromosome 3 of Z. nicaraguensis conferred ROL barrier formation in the genetic background of maize. This chromosome segment also decreased apoplastic solute permeability across the hypodermis/exodermis. However, the IL and maize were similar for suberin staining in the hypodermis/exodermis at 40 mm and further behind the root tip. Z. nicaraguensis contained suberin in the hypodermis/exodermis at 20 mm and lignin at the epidermis. The IL with ROL barrier, however, did not contain lignin in the epidermis. Discovery of the Z. nicaraguensis chromosomal region responsible for root ROL barrier formation has improved knowledge of this trait and is an important step towards improvement of waterlogging tolerance in maize.

SUMMARYSuberin is a complex polymer composed of aliphatic and phenolic compounds. It is a constituent of apoplastic plant interfaces. In many plant species, including rice (Oryza sativa), the hypodermis in the outer part of roots forms a suberized cell wall (the Casparian strip and/or suberin lamellae), which inhibits the flow of water and ions and protects against pathogens. To date, there is no genetic evidence that suberin forms an apoplastic transport barrier in the hypodermis. We discovered that a rice reduced culm number1 (rcn1) mutant could not develop roots longer than 100 mm in waterlogged soil. The mutated gene encoded an ATP-binding cassette (ABC) transporter named RCN1/OsABCG5. RCN1/OsABCG5 gene expression in the wild type was increased in most hypodermal and some endodermal roots cells under stagnant deoxygenated conditions. A GFP-RCN1/OsABCG5 fusion protein localized at the plasma membrane of the wild type. Under stagnant deoxygenated conditions, well suberized hypodermis developed in wild types but not in rcn1 mutants. Under stagnant deoxygenated conditions, apoplastic tracers (periodic acid and berberine) were blocked at the hypodermis in the wild type but not in rcn1, indicating that the apoplastic barrier in the mutant was impaired. The amount of the major aliphatic suberin monomers originating from C 28 and C 30 fatty acids or x-OH fatty acids was much lower in rcn1 than in the wild type. These findings suggest that RCN1/OsABCG5 has a role in the suberization of the hypodermis of rice roots, which contributes to formation of the apoplastic barrier.

Plants that are adapted to waterlogged conditions develop aerenchyma in roots for ventilation. Some wetland plant species also form an apoplastic barrier at the outer cell layers of roots that reduces radial oxygen loss (ROL) from the aerenchyma and prevents toxic compounds from entering the root. The composition of the apoplastic barrier is not well understood. One potential component is suberin, which accumulates at the hypodermal/exodermal cell layers of the roots under waterlogged soil conditions or in response to other environmental stimuli. However, differences in suberin content and composition between plant species make it difficult to evaluate whether suberin has a role in preventing ROL. In this article, we summarize recent advances in understanding apoplastic barrier formation in roots and, between various plant species, compare the chemical compositions of the apoplastic barriers in relation to their permeability to oxygen. Moreover, the relationship between suberin accumulation and the barrier to ROL is discussed.

The formation of a barrier to radial oxygen (O2 ) loss (ROL) in the root is an important adaptation of plants to root flooding, but the biochemical changes in plant roots where the barrier is formed are unclear. In this study, we analysed metabolic profiles and gene expression profiles in roots of rice (Oryza sativa L.) plants grown under stagnant deoxygenated conditions, which induce suberization in the outer cell layers of the roots and formation of barrier to ROL. Under these conditions, two distinctive biochemical features of the roots were the accumulations of malic acid and very long chain fatty acids (VLCFAs). We also showed that the expressions of some genes encoding plastid-localized enzymes, which convert malic acid to acetyl coenzyme A (AcCoA), were simultaneously up-regulated under stagnant conditions. The expression levels of these genes in specific root tissues isolated by laser microdissection suggested that malic acid is converted to AcCoA predominantly in the plastids in the outer cell layers of rice roots. We propose that the physiological role of malic acid accumulation in rice roots grown under stagnant conditions is to provide a substrate for the biosynthesis of fatty acids, which, in turn, are used in the biosynthesis of suberin.

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Or ri ig gi in na al l r re es se ea ar rc ch h a ar rt ti ic cl le e R Re ev vi ie ew w a ar rt ti ic cl le e S Sh ho or rt t r re ep po or rt t Nishiuchi S, Watanabe K, Sato S, Takahashi H and Nakazono M 2021 Expression analysis of genes for cytochrome P450 CYP86 and glycerol-3-phosphate acyltransferase related to suberin biosynthesis in rice roots under stagnant deoxygenated conditions.