Abstract

Suberin is a specialized cell wall modifying polymer comprising both phenolic-derived and fatty acid-derived monomers, which is deposited in below-ground dermal tissues (epidermis, endodermis, periderm) and above-ground periderm (i.e., bark). Suberized cells are largely impermeable to water and provide a critical protective layer preventing water loss and pathogen infection. The deposition of suberin is part of the skin maturation process of important tuber crops such as potato and can affect storage longevity. Historically, the term “suberin” has been used to describe a polyester of largely aliphatic monomers (fatty acids, ω-hydroxy fatty acids, α,ω-dioic acids, 1-alkanols), hydroxycinnamic acids, and glycerol. However, exhaustive alkaline hydrolysis, which removes esterified aliphatics and phenolics from suberized tissue, reveals a core poly(phenolic) macromolecule, the depolymerization of which yields phenolics not found in the aliphatic polyester. Time course analysis of suberin deposition, at both the transcriptional and metabolite levels, supports a temporal regulation of suberin deposition, with phenolics being polymerized into a poly(phenolic) domain in advance of the bulk of the poly(aliphatics) that characterize suberized cells. In the present review, we summarize the literature describing suberin monomer biosynthesis and speculate on aspects of suberin assembly. In addition, we highlight recent advances in our understanding of how suberization may be regulated, including at the phytohormone, transcription factor, and protein scaffold levels.

Highlights

  • Plants have developed effective processes to facilitate their survival, including the production of secondary metabolites [1,2]

  • This review provides an overview of the current state of knowledge of suberin monomer biosynthesis, the assembly of monomers into the suberin macromolecule, and the coordination of suberin deposition by phytohormones, transcription factors (TF) and Casparian strip membrane domain proteins (CASPs)

  • Arabidopsis gpat5 loss-of-function mutants demonstrate substantial decreases in C20-C24 very-long-chain fatty acids (VLCFA) and their ω-hydroxy and dicarboxylic acid derivatives in suberin found in roots and seed coats, and overexpression of AtGPAT5 led to accumulation of sn-2 monoacylglycerols in the wax of Arabidopsis stems [20,21]

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Summary

Introduction

Plants have developed effective processes to facilitate their survival, including the production of secondary metabolites [1,2] The bioactivities of these metabolites involve regulating plant growth, and enabling plants to cope with stressful conditions including biotic threats and environmental hazards [3]. This review provides an overview of the current state of knowledge of suberin monomer biosynthesis, the assembly of monomers into the suberin macromolecule, and the coordination of suberin deposition by phytohormones, TFs and CASPs. An increased fundamental understanding of the role of suberin in response to various stressors, and of the mechanisms that regulate the suberization process, may have important implications for crop improvement efforts, including enhanced tuber storage and resistance to drought stress and pathogen infection [27]

The Suberin Enigma
Suberin Biosynthesis and Assembly
Biosynthesis and Polymerization of Phenolic Monomers
Phenylpropanoid Metabolism during Suberization
Biosynthesis of Aliphatic Monomers
Elongation
Oxidation
Reduction
Acyl-CoA Dependent Aliphatic Monomer Esterification
Possible Mechanisms for Aliphatic Monomer Polymerization
The Temporal Deposition of Suberin
Regulation of Suberization
Other Phytohormones
Transcription Factors
MYB and MYC Transcription Factors
NAC Transcription Factors
WRKY Transcription Factors
Interaction between Phytohormones and Transcription Factors
A Role for Casparian Strip Membrane Domain Proteins
Findings
Conclusions

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