Development of the quadripolar meiotic cytoskeleton in spore mother cells of the moss Funaria hygrometrica

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Abstract
Evidence presented in the accompanying paper that plastids function as microtubule (MT)-organizing centres for development of the quadripolar cytoskeleton of pre-meiotic spore mother cells (SMCs) in the moss Funaria hygrometrica is complemented here by observations on the MT system in these cells. Early in meiotic prophase numerous MTs align progressively along the two plastids as they elongate. Concomitant with (and perhaps causal for) plastid rotation, new MT arrays grow from each tip of each plastid to both tips of the other plastid. The ‘along-plastid’ and ‘between-plastid’ arrays ultimately form the edges of a tetrahedron, enclosing the prophase nucleus. MT breakdown at the centre of each edge leaves four cones of MTs, one emanating from each vertex, located at the plastid tips. These partially fuse in between-plastid pairs to give a twisted spindle with broad knife-edge poles oriented at right angles to one another, i.e. a condensed form of the quadripolar precursor. The twist causes the metaphase plate and the subsequent phragmoplast and organelle band to be saddle-shaped, and the daughter nuclei to be elongated perpendicular to one another along the two knife edges. The tetrahedral array returns during interkinesis and again breaks down into four cones of MTs centred on the plastid tips; these, however, now become individual half spindles for the two perpendicularly arranged second division spindles. When meiosis is completed the four haploid nuclei thus come to lie at the vertices of a tetrahedron that was established by MT-mediated plastid positioning during meiotic prophase. The tetrahedral cage of MTs precedes meiosis yet predicts the planes of division, and in these two respects it is the meiotic counterpart of the preprophase band of MTs, which develops before mitosis in most higher plant cells.