Abstract

The Three Amino acid Loop Extension (TALE) proteins constitute an ancestral superclass of homeodomain transcription factors conserved in animals, plants and fungi. In plants they comprise two classes, KNOTTED1-LIKE homeobox (KNOX) and BEL1-like homeobox (BLH or BELL, hereafter referred to as BLH), which are involved in shoot apical meristem (SAM) function, as well as in the determination and morphological development of leaves, stems and inflorescences. Selective protein-protein interactions between KNOXs and BLHs affect heterodimer subcellular localization and target affinity. KNOXs exert their roles by maintaining a proper balance between undifferentiated and differentiated cell state through the modulation of multiple hormonal pathways. A pivotal function of KNOX in evolutionary diversification of leaf morphology has been assessed. In the SAM of both simple- and compound-leafed seed species, downregulation of most class 1 KNOX (KNOX1) genes marks the sites of leaf primordia initiation. However, KNOX1 expression is re-established during leaf primordia development of compound-leafed species to maintain transient indeterminacy and morphogenetic activity at the leaf margins. Despite the increasing knowledge available about KNOX1 protein function in plant development, a comprehensive view on their downstream effectors remains elusive. This review highlights the role of TALE proteins in leaf initiation and morphological plasticity with a focus on recent advances in the identification of downstream target genes and pathways.

Highlights

  • Homeodomain (HD) transcription factors play major roles in the development of complex organisms, ranging from humans to plants

  • A set of studies on tomato, including the KN1 overexpression in mutants with different levels of leaf dissection [79] and the switch of KNOX1 proteins into repressors combined with the time-regulated expression in leaf primordia [55], revealed that: (a) the sensitivity to KNOX1 action varies during leaf development; (b) the KNOX1 activity is restricted to the developmental window of primary morphogensis [38]

  • Common genetic pathways have been unravelled in different developmental programs such as organ primordia initiation at the shoot apical meristem and the morphogenetic events that shape margins and complexity of leaves

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Summary

Introduction

Homeodomain (HD) transcription factors play major roles in the development of complex organisms, ranging from humans to plants. Increasing morphological complexity requires finely tuned processes of polarized growth along different axis to establish specialized organs with determinate shapes and symmetries. This process is fundamental during development and involves the integration of multiple signals from different parts of the organism’s body. Among the four subgroups of TALEs identified, the members of MEIS/PREP and Pre-B cell leukaemia transcription factors (PBX) form heterotrimers with homeobox proteins (Hox). Together they act within transcriptional regulation complexes to specify anteroposterior identities and regulate genetic programs during development [4]. Some KNOXs are able to move from cell to cell through plasmodesmata (D) [33,34,35,36]

Overview of the Role of TALEs in Shoot Apical Meristem
TALE Genes in Organ Primordia Initiation
TALE Genes in the Control of Simple Leaf Morphology
TALE Genes in the Control of Compound Leaf Morphology
ILRC Subgroup of Legumes
TALEs Work at the Boundaries
A Matter of Targets: the Secret Agents of TALEs
Links to Hormones
Transcription Factors
Cell Wall Proteins
Conclusions and Perspectives
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