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Table 1 Genes involved in major developmental events of the leaf

From: What determines a leaf's shape?

Developmental events

Genes

Description

Biological function

Mutant phenotype

Plant species

References

Leaf initiation

PIN-FORMED1 (PIN1)

Transmembrane protein

Auxin efflux

Develop naked, pin-shaped inflorescences; leaves become fused; phyllotaxy disrupted

Arabidopsis thaliana

[23–25]

Class-1 KNOTTED-like homeobox (KNOX1)

Homeodomain protein

Maintain stem cell identity

Loss-of-function mutants failed to develop SAM; gain-of-function mutants showed ectopic SAMs on leaves

Arabidopsis thaliana; Zea mays

[26, 27]

WUSCHEL (WUS)

Homeodomain protein

Maintain shoot and floral central meristem identity

Delayed growth; disorganized rosette leaves; inflorescence meristem defective

Arabidopsis thaliana

[28]

CLAVATA (CLV)

CLV1 (receptor kinase); CLV2 (transmembrane protein); CLV3 (extracellular protein)

Maintain stem cell size

Enlarged shoot and floral meristems; stem overgrowth; additional floral organs

Arabidopsis thaliana

[29, 30]

ASYMMETRIC LEAVES1/ ROUGH SHEATH2/ PHANTASTICA (ARP)

MYB domain protein

Stem cell differentiation

Stunted growth; polarity defects; unlike as1 and rs2, phan leaves are radialized

Arabidopsis thaliana; Zea mays; Antirrhinum majus

[31–34]

Leaf outgrowth: proximodistal patterning

KNOTTED1 (KN1)

Homeodomain protein

Maintain stem cell identity

Gain-of-function mutants displayed flaps of sheath tissue at leaf blade margin; leaf bifurcated

Zea mays

[35]

LIGULELESS NARROW-REFERENCE (LGN-R)

Serine/threonine kinase

Establishment of blade/sheath boundary

Heterozygotes displayed narrower and shorter leaves; homozygotes failed reproductive development

Zea mays

[36]

Leaf outgrowth: dorsoventral patterning

PHANTASTICA (PHAN)

MYB domain protein

Stem cell differentiation

Loss-of-function phan mutants develop needle-like leaves lacking dorsoventrality

Antirrhinum majus

[37]

ASYMMETRIC LEAVES2 (AS2)

LOB domain protein

Leaf venation pattern and lamina development

Develop narrow and curly leaves with alteration in adaxial/abaxial polarity

Arabidopsis thaliana

[38]

class III HOMEODOMAIN-LEUCINE ZIPPER (HD-ZIPIII)

Homeodomain and leucine zipper domain protein

Leaf polarity, meristem function

Lateral organs radialized with adaxial cell fate transformation; modification in vascular patterning

Arabidopsis thaliana

[39, 40]

KANADI (KAN)

GARP domain protein

Leaf polarity specification

Develop narrow adaxialized lateral organs; ectopic outgrowths on leaves; gain-of-function mutants displayed abaxialized cell types; blade expansion inhibited

Arabidopsis thaliana

[41, 42]

APUM23

PUF RNA-binding protein

Leaf polarity specification

Radialized leaves; disorganized vascular pattern

Arabidopsis thaliana

[43]

AUXIN RESPONSE FACTOR (ARF)

Protein with N-terminal DNA binding domain, activator/repressor domain, C-terminal dimerization domain

Leaf polarity specification

Narrow leaves with ectopic blade outgrowths

Arabidopsis thaliana

[44]

miR165

21-nucleotide non-coding RNAs

Leaf polarity specification, meristem function, vascular development

Loss of SAM; altered organ polarity; defective vascular development

Arabidopsis thaliana

[45]

miR166

21-nucleotide non-coding RNAs

Leaf polarity specification, meristem function, vascular development

Enlarged SAM; enhanced vascular development

Arabidopsis thaliana

[46]

RNA-DEPENDENT RNA POLYMERASE6 (RDR6)/DICER-LIKE4 (DCL4)

RNA-dependent RNA polymerase/RNase-III enzyme

Leaf polarity specification

Accelerated juvenile-to-adult phase transition; early development of adult lateral organs; lack ta-siRNAs

Arabidopsis thaliana

[47–49]

Leaf outgrowth: mediolateral patterning

YABBY (YAB)

Protein with zinc-finger and helix-loop-helix domains

Leaf polarity specification, lamina expansion

Minuscule and bushy plants with loss of lamina expansion and polarity defects

Arabidopsis thaliana

[50, 51]

Narrow sheath (ns)

Homeodomain protein

Leaf founder cell recruitment, leaf expansion

Develop extremely narrow leaves; short internode

Zea mays

[52, 53]

PRESSED FLOWER (PRS)

Homeodomain protein

Marginal cell proliferation

Smaller sepals; defective marginal regions

Arabidopsis thaliana

[54]

MAEWEST (MAW)

Homeodomain protein

Organ fusion and lateral expansion

Severe leaf blade reduction, thickened leaf margins; petal expansion reduced; defective carpel fusion

Petunia × hybrida

[55]

YUCCA (YUC)

Flavin monooxygenase

Leaf and vascular development, floral patterning

Stunted growth with curved leaves; smaller inflorescence meristem; defective floral and leaf vasculature

Arabidopsis thaliana

[56, 57]

Leaf expansion and maturation

ANGUSTIFOLIA (AN3)/GRF-INTERACTING FACTOR1(GIF1)

Transcription coactivator

Cell expansion

Reduced leaf width and length; petal width reduction; more leaf number

Arabidopsis thaliana

[58, 59]

GROWTH-REGULATING FACTOR5 (GRF5)

Transcription activators containing N-terminal QLQ or WRC domain

Cell proliferation

Loss-of-function mutants displayed narrow leaves and petals; gain-of-function mutants develop

Arabidopsis thaliana

[58, 59]

CINCINNATA (CIN)

TCP domain protein

Cell proliferation

Develop large crinkly leaves

Antirrhinum majus

[60]

Leaf margin alterations

miR164A

Non-coding miRNA

Leaf margin development

Enhanced leaf margin serration in loss-of-function mutants; gain-of-function mutants develop leaves with smooth margins

Arabidopsis thaliana

[61]

 

CUP-SHAPED COTYLEDON2 (CUC2)

Protein containing the NAC DNA-binding domain

Shoot meristem formation; organ boundary specification; leaf margin development

Produced leaves with smooth margins

Arabidopsis thaliana

[61]

 

PIN-FORMED1 (PIN1)

Transmembrane protein

Auxin efflux

Loss-of-function mutants develop smooth leaf margins

Arabidopsis thaliana

[62]

 

DEVELOPMENT-RELATED PcG TARGET IN THE APEX (DPA)

RAV transcription repressor

Organ initiation and development; leaf margin development

Loss-of-function mutants showed increased leaf margin serrations and enlarged petals; gain-of-function mutants possessed smooth margins

Arabidopsis thaliana

[63]