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. 2017 May 12;174(3):1728–1746. doi: 10.1104/pp.17.00413

Table I. Developmental and cytological changes during lamina joint (LJ) growth.

Stage Morphology Cytology
S1 (Initiation) A ligule and a pair of auricles differentiate at the LJ initiation site No aerenchyma in LJ, which is full with parenchyma cells, except vascular bundles arranged at the abaxial side
LJ is hollow and transparent Small parenchyma cell clusters at the adaxial side and abaxial region between developing vascular bundles
The periphery of abaxial vascular bundles protrudes to form ridges
S2 (Young) LJ, leaf blade, and sheath are white or creamy yellow LJ size increases mainly due to the extensive expansion of parenchyma cells
LJ protrudes and becomes larger PCD of mature large parenchyma cells occurs to form small holes
Longitudinal holes connect and fuse to form intact aerenchyma
S3 (Young) Leaf continues to elongate, and leaf blade expands Rapid cell division between the vascular bundles and epidermis cells at the abaxial domain
LJ is still enclosed by the prior leaf sheath Small parenchyma cell clusters are formed at the adaxial side
Leaf blade and sheath become green, while LJ is white (distinguishable from each other) Parenchyma cells continue expanding
A pair of horn-shaped auricles and a tongue-like ligule can be observed Large and small vascular bundles locating at the abaxial side develop continuously
S4 (Maturation) Leaf blade and sheath fully develop Thickening of cell walls of small cell clusters results in the formation of sclerenchyma cell clusters
LJ emerges from prior leaf sheath and is exposed in the air Newly formed sclerenchyma cells at both sides perforate through longitudinally to form sclerenchyma fibers
LJ is firm and plump Vascular bundles and nearby wall-thickened small cell clusters at the adaxial side form ridges along the longitudinal direction
LJ starts to bend Parenchyma cells stop proliferation
The periphery of whole LJ becomes smooth and the transverse size of the LJ achieves its maximum
S5 (Postmaturation) LJ adaxial side elongates and bends to the abaxial side to form a curvature, resulting in the increasing leaf angle Ongoing thickening of cell walls of small cell clusters on both sides
Parenchyma cells at the adaxial side undergo longitudinal elongation
Asymmetric cell growth between the adaxial and abaxial sides leads to increased leaf inclination
S6 (Senescence) LJ approaches the maximum angle and begins to wither due to the loss of water Larger aerenchyma and fewer layers of parenchyma cells
LJ is dehydrated and distorted