Abstract

Auxin contributes to almost every aspect of plant development and metabolism as well as the transport and signalling of auxin-shaped plant growth and morphogenesis in response to endo- and exogenous signals including stress conditions. Consistently with the common belief that auxin is a central trigger of developmental changes in plants, the auxin treatment of explants was reported to be an indispensable inducer of somatic embryogenesis (SE) in a large number of plant species. Treating in vitro-cultured tissue with auxins (primarily 2,4-dichlorophenoxyacetic acid, which is a synthetic auxin-like plant growth regulator) results in the extensive reprogramming of the somatic cell transcriptome, which involves the modulation of numerous SE-associated transcription factor genes (TFs). A number of SE-modulated TFs that control auxin metabolism and signalling have been identified, and conversely, the regulators of the auxin-signalling pathway seem to control the SE-involved TFs. In turn, the different expression of the genes encoding the core components of the auxin-signalling pathway, the AUXIN/INDOLE-3-ACETIC ACIDs (Aux/IAAs) and AUXIN RESPONSE FACTORs (ARFs), was demonstrated to accompany SE induction. Thus, the extensive crosstalk between the hormones, in particular, auxin and the TFs, was revealed to play a central role in the SE-regulatory network. Accordingly, LEAFY COTYLEDON (LEC1 and LEC2), BABY BOOM (BBM), AGAMOUS-LIKE15 (AGL15) and WUSCHEL (WUS) were found to constitute the central part of the complex regulatory network that directs the somatic plant cell towards embryogenic development in response to auxin. The revealing picture shows a high degree of complexity of the regulatory relationships between the TFs of the SE-regulatory network, which involve direct and indirect interactions and regulatory feedback loops. This review examines the recent advances in studies on the auxin-controlled genetic network, which is involved in the mechanism of SE induction and focuses on the complex regulatory relationships between the down- and up-stream targets of the SE-regulatory TFs. In particular, the outcomes from investigations on Arabidopsis, which became a model plant in research on genetic control of SE, are presented.

Highlights

  • The process of somatic embryogenesis (SE) demonstrates the unique developmental capacity of plants for switching on the embryogenic programme of development in somatic cells that have already differentiated

  • Distinct progress in deciphering the molecular mechanism that governs the embryogenic transition of plant somatic cells has been made in the last decade and the role of auxin in SE induction has been intensively studied at the molecular level

  • The transcripts of the transcription factor genes (TFs) genes that are related to hormone responses, in particular those that control auxin metabolism and signalling are overrepresented in the SE-transcriptomes of many plants

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Summary

Introduction

The process of somatic embryogenesis (SE) demonstrates the unique developmental capacity of plants for switching on the embryogenic programme of development in somatic cells that have already differentiated. There are some differences between the in vitro culture conditions that are applied to induce SE in different plants, treating explants with auxin and auxin-like substances seems to be commonly required for SE induction In support of this statement, auxin treatment was applied in 74 of the 80 protocols reviewed in Table S1 and auxin was frequently combined with cytokinins (65%). 2,4-D treatment activated the core regulators of the auxin-signalling AUXIN RESPONSE FACTORs (ARFs) and the TRYPTOPHAN AMINOTRANSFERASE OF ARABIDOPSIS1/YUCCA, TAA1/YUC-dependent auxin biosynthesis pathway in SE-induced explants of Arabidopsis [36,37] Both the auxin- and stressor-like intrinsic function of 2,4-D are closely related in promoting embryogenic development in plant cells

Biosynthesis and Accumulation of Auxin During SE
Complex Interactions Between the TF Genes that Control Auxin-Induced SE
Findings
Conclusions and Future Prospects
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