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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 [2325]
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 [3134]
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 [4749]
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]