Cell cycling plays an important role in plant development, including: (1) organ morphogenesis, (2) cell proliferation within tissues, and (3) cell differentiation. In this study we use a cyclin::beta-glucuronidase reporter construct to characterize spatial and temporal patterns of cell cycling at each of these levels during wild-type development in the model genetic organism Arabidopsis thaliana (Columbia). We show that a key morphogenetic event in leaf development, blade formation, is highly correlated with localized cell cycling at the primordium margin. However, tissue layers are established by a more diffuse distribution of cycling cells that does not directly involve the marginal zone. During leaf expansion, tissue proliferation shows a strong longitudinal gradient, with basiplastic polarity. Tissue layers differ in pattern of proliferative cell divisions: cell cycling of palisade mesophyll precursors is prolonged in comparison to that of pavement cells of the adjacent epidermal layers, and cells exit the cycle at different characteristic sizes. Cell divisions directly related to formation of stomates and of vascular tissue from their respective precursors occur throughout the period of leaf extension, so that differing tissue patterns reflect superposition of cycling related to cell differentiation on more general tissue proliferation. Our results indicate that cell cycling related to leaf morphogenesis, tissue-specific patterns of cell proliferation, and cell differentiation occurs concurrently during leaf development and suggest that unique regulatory pathways may operate at each level.

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Summary Primary vascular pattern determines pathways of long distance transport for water, nutrients and signalling molecules within plant shoot systems. Expression of an Arabidopsis thaliana homeobox gene 8::GUS construct is restricted to the procambium and provides a molecular marker for vascular pattern at early developmental stages. Primary vascular pattern and phyllotaxis are highly coordinated, with vascular sympodia corresponding to phyllotactic parastichies. During vegetative development, primary vasculature forms a reticulate pattern, with each leaf trace derived from two vascular sympodia. Shoot phase change is marked by alterations of this fundamental pattern. Formation of leaf trace procambial strands is temporally coordinated with primordium initiation, but ATHB‐8::GUS expression is discontinuous in these strands at early stages. The interconnection of vascular sympodia provides alternate pathways for long distance transport in shoots of A. thaliana and reflects the limited secondary growth in this species. ATHB‐8::GUS expression identifies a prepattern that precedes anatomical definition of procambium. The longitudinal discontinuity in ATHB‐8::GUS expression indicates that one function of this gene is to define the xylem components of vascular radial pattern.

The unusual perforations in the leaf blades of Monstera obliqua (Araceae) arise through programmed cell death early in leaf development. At each perforation site, a discrete subpopulation of cells undergoes programmed cell death simultaneously, while neighboring protoderm and ground meristem cells are unaffected. Nuclei of cells within the perforation site become terminal deoxynucleotidyl transferase-mediated dUTP nick end labeling (TUNEL)-positive, indicating that DNA cleavage is an early event. Gel electrophoresis indicates that DNA cleavage is random and does not result in bands that represent multiples of internucleosomal units. Ultrastructural analysis of cells at the same stage reveals misshapen, densely stained nuclei with condensed chromatin, disrupted vacuoles, and condensed cytoplasm. Cell walls within the perforation site remain intact, although a small disk of dying tissue becomes detached from neighboring healthy tissues as the leaf expands and stretches the minute perforation. Exposed ground meristem cells at the rim of the perforation differentiate as epidermal cells. The cell biology of perforation formation in Monstera resembles that in the aquatic plant Aponogeton madagascariensis (Aponogetonaceae; Gunawardena et al. 2004), but the absence of cell wall degradation and the simultaneous execution of programmed cell death throughout the perforation site reflect the convergent evolution of this distinct mode of leaf morphogenesis in these distantly related plants.

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