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Asymmetric cell divisionsCell-cycle entry will lead to the actual division process to form two new daughter cells. Two fundamentally different types of cell division occur during plant growth and development. One type exclusively generates new cells with the same cell fate and is designated 'proliferative division'. Proliferative cell divisions are primarily responsible for growth maintenance of the plant body by providing new cells and do not result in the formation of new cell types, tissues or organs. By contrast, for the onset of differentiation, either in the meristem itself or in quiescent cells outside the meristems, a special type of cell division -- 'generative' or 'asymmetric' division -- is required, in which the two daughter cells acquire distinct fates. In plants, asymmetric divisions are explicitly involved in developmental processes, such as the start of embryogenesis, stomata and pollen development, and lateral root initiation. Asymmetric divisions are also essential for stem-cell maintenance: a stem cell must divide asymmetrically to give rise to a stem-cell daughter cell and a second cell that is committed to a new cell fate.It is currently unclear what causes the distinction between a symmetric and an asymmetric division at the molecular level. In Drosophila melanogaster , evidence is accumulating that some of the key cell-cycle regulators themselves (such as the CDK-activating phosphatase string (also known as cdc25 )) function as intrinsic factors for the asymmetric localization of cell-fate determinants [21]. Although similar links between cell-cycle and cell-fate specification mechanisms have not yet been firmly established in plants, the basic cell-cycle machinery is probably also involved in controlling asymmetric division. For example, whereas new roots and shoots can be obtained easily from wild-type A. thaliana leaf explants, the regeneration of new tissues is completely impaired in plants that ectopically express the A-type cyclin CYCA3;2 , indicating that CYCA3;2-CDK activity might be crucial for the divergence between proliferative and generative asymmetric divisions [22].Another example of a putative direct link between cell-fate specification and cell division can be observed during stomata development, and involves the plant-specific CDKB1;1. This particular B-type CDK is highly active in stomatal precursor cells that divide asymmetrically. Evidence for its potential function in stomatal asymmetric division has been demonstrated by downregulation of CDKB1;1 activity in transgenic A. thaliana plants, resulting in a strong decrease in the number of stomata because of an arrest of the stomatal precursor divisions [23].
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