Age-related and origin-related control of the numbers of plasmodesmata in cell walls of developing Azolla roots

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Abstract
Plasmodesmata were counted in the longitudinal and transverse walls in developmental sequences of merophytes in roots of Azolla pinnata R.Br. The differences between certain categories of longitudinal wall were traced to factors that govern the surface area of the cell plates, the density of plasmodesmata (number per unit area of cell plate), and the amount by which each type of plate expands. No evidence for secondary augmentation of plasmodesmatal numbers after the cell-plate stage of development was found, but plasmodesmata are lost from the walls of sieve and xylem elements during their differentiation. Losses caused by cell separation occur in other tissues. The relatively high density of plasmodesmata in transverse walls is based not so much on a high density in the cell plates as on the relatively low expansion that these walls undergo. There appears to be a compensatory mechanism that relates initial plasmodesmatal density to the future expansion of the cell plate. The root shows determinate growth, the apical cell dividing about 55 times. Beginning at about the 35th division there is a progressive failure to maintain the plasmodesmatal frequencies that were developed in earlier cell divisions in the apical cell. The divisions that occur within the later-produced merophytes also show progressive diminution of plasmodesmatal numbers. The result is that the apex of the root, and particularly the apical cell, becomes more and more isolated symplastically, a phenomenon which could account for its limited lifespan and the determinate growth pattern of the root.