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Do cell-intrinsic (lineage) or cell-extrinsic (neighbours) signals drive plant development? Intrinsic vs. Extrinsic signals drive development? •In animals, simpler animals driven by intrinsic signals, more complex animal by extrinsic •Intrinsic = Signals that are contained in the cell, often arising from factors passed down from ancestor cells (parentage is critical, position less so) •Extrinsic signals = cell-cell signaling between neighbours (position is critical, parentage less so) 1 Plant cell division: •Plant cells are surrounded by walls (cellulose microfibrils in matrix of pectins and hemicelluloses (1o) or lignin (2o)) •Lignin is hydrophobic and prevents cell expansion •little cell migration (cell sliding possible) Cells of same parentage are also neighbours Figure 2-1 Raven, Fig 8-12 •Preprophase band - microtubules that predict position of cell plate •Cell plate forms through fusion of golgi derived vesicles 2 •Fusion of golgi vesicles to form cell membrane and cell wall Raven, Fig 8-10 Strands of ER may be trapped between fusing vesicles, resulting in plasmodesmata 3 Intrinsic (lineage) and extrinsic (position) difficult to separate in plants: •daughter cells connected by cell wall - therefore are neighbours •Positional information could support either lineage or neighbour as a signaling mechanism How to assess importance of lineage? •Clonal analysis •Do the same cells always have the same fate? Figure 3-1 4 Chimeric plants - sectors of genetically different cells •Sectorial vs. periclinal Figure 3-2 Cells of a meristematic sector are not restricted a particular fate (e.g. leaf/not-leaf) Cells of a meristematic layer generally give rise to particular cell layer fimbriata floricaula deficiens L2 chimera for the gene fimbriata floricaula fimbriata L1 chimera for floricaula Schultz et al., 2001 5 Cell killing - can one cell adopt the fate of another? •Ablate QC, procambium divides to fill QC •Assessed by QC, procambium, columella specific markers •suggests extrinsic rather than intrinsic Sabatini et al., 1999, Figure 3-3 Figure 3.4 Van den Berg, 1995 If kill cortical initial, pericycle cell divides periclinally to fill cortical space But what if consequence of ablation? Cells are totipotent - not surprising if this pathway is induced by laser 6 Chimeric plants provide a situation where layer invasions can be monitored without ablation •Normally L1 = epidermis (no chlorophyll), L2 = subepidermis, L3 everything else •GWG chimeras •Fringe - L2 has adopted normally L3 fate •Pale green - L1 (green) has invaded L2 •Therefore, layers are not always clonally distinct, but when invasions occur, cells adopt fate according to position, not origin Figure 3.4, Stewart, 1978 Colchicine induced meristematic chimeras in Datura •Determined that cell layers usually maintain distinct lineages •Cell sizes are very different, but co-ordination of cell division between layers compensates Figure 3.6 Satina et al., 1940 7 Mutations that affect cell shape do not affect gross morphology of organs •Therefore, plant can compensate for defects in normal lineage •Changes in cell expansion compensate for defects in division Maize leaf More transverse divisions Maize leaf Figure 3.7 Cleary and Smith, 1998 Cell division and expansion are coordinated •Gamma irradiating wheat embryos stops cell division, but does not stop initiation of primordia •Primordium is initiated by expansion of cells (cannot form further) Figure 3.8 Foard, 1971 8 fass mutants •Cell expansion and cell division are abnormal •Gross morphology is abnormal •Cell fates are maintained Figure 3.9 Torrez-Ruiz and Jurgens, 1994 Cell age and cell position are closely linked in plants Heterochronic - change in time of cell fate acquisition Homeotic - change in position of cell fate acquistion Not easy to distinguish in plants ap2-6, sepals become carpels 9 paused mutant - cell death in CZ •slow leaf initiation, but leaf fate is not slowed •Plant size and position of organ are changed, fate is not •Suggests age is critical Figure 3-10 Telfer et al., 1997 altered meristem programming (amp1) • rate of leaf initiation is increased •Fate of leaves is like wild type at a similar age •i.e. more leaves of each type Psd and amp1 support the idea that age, not position, is critical •GAs as age monitor? •Application of GA accelerate abaxial trichomes •Defects in GA delay abaxial trichomes 10