Our understanding of the molecular controls regulating the identity of the vascular cambium and the development of secondary xylem and phloem have not yet benefited much from the use of Arabidopsis as a genetic system. Under appropriate growth conditions Arabidopsis undergoes extensive secondary growth in the hypocotyl, with the development of both a vascular and a cork cambium. The secondary xylem of the hypocotyl develops in two phases, an early phase in which only vessel elements mature and a later stage in which both vessel elements and fibres are found. During this second phase the secondary xylem of Arabidopsis closely resembles the anatomy of the wood of an angiosperm tree, and can be used to address basic questions about wood formation. The development of the vascular cambium and secondary growth in Arabidopsis hypocotyl is described and its utility as a model for wood formation in trees is considered.

Rearrangements of cortical microtubules (CMTs) during the dierentiation of axial secondary xylem elements within taproots and shoots of Aesculus hippocastanum L. (horse-chestnut) are described. A correlative approach was employed using indirect immuno¯uorescence microscopy of a-tubulin in 6-to 10-lm sections and transmission electron microscopy of ultrathin sections. All cell types ± ®bres, vessel elements and axial parenchyma ± derive from fusiform cambial cells which contain randomly oriented CMTs. At the early stages of development, ®bres and axial parenchyma cells possess helically arranged CMTs, which increase in number as secondary wall thickening proceeds and simple pits develop. In contrast, incipient vessel elements are distinguished by the marking out of sites of bordered pits; these sites ®rst appear as microtubule-free regions within the reticulum of randomly oriented CMTs that characterises their precursor fusiform cambial cells. Subsequently, the ring of CMTs which develops at the periphery of the microtubule-free region decreases in diameter as the over-arching pit border is formed. Like bordered pits, large-diameter, non-bordered pits (contact pits) which develop between vessel elements and adjacent contact ray cells originate as microtubule-free regions and are also associated with development of a ring of CMTs at the periphery. In the case of contact pits, however, there is no reduction in the diameter of the CMT ring during pit development. Tertiary cell wall thickenings are also a feature of vessel elements and appear to form at sites where bands of laterally associated, transversely oriented CMTs, separated from each other by microtubule-free zones, are found. Later, these bands of CMTs become narrower, and separate into pairs of microtubule bundles located on each side of the developing wall thickening. Development of perforations between vessel elements is also associated with the presence of a ring of CMTs at their periphery.

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SummaryDespite new techniques for studying the cell biology of plant development in recent years, the secondary vascular system has been neglected. Why is this? Here it is argued that some of the barriers that have prevented more widespread study of the tree secondary vascular system are no longer valid. Some of the more intriguing aspects of the secondary vascular system include the recent discovery of a putative plant muscle and identification of a cytoskeleton-facilitated three-dimensional symplasmic transport pathway that permeates the tree. There are great merits in the recently adopted model tree species, poplar, and a new model system -wood formation in Arabidopsis . The time is now right for much greater exploitation of the possibilities that exist for study of the secondary vascular system of trees.© New Phytologist (2002) 153 : 213 -223

The immunolocalisation of unconventional myosin VIII ('myosin') in the cells of the secondary vascular tissues of angiosperm (Populus tremula L. x P. tremuloides Michx. and Aesculus hippocastanum L.) and gymnosperm (Pinus pinea L.) trees is described for the first time and related to other cytoskeletal elements, as well as to callose. Both myosin and callose are located at the cell plate in dividing cambial cells, whereas actin microfilaments are found alongside the cell plate; actin and tubulin are both associated with the phragmoplast. Myosin and callose also localise to the plasmodesmata-rich pit fields in the walls of living cells, which are particularly abundant within the common walls between ray cells and between ray cells and axial parenchyma cells in the phloem and xylem. In those xylem ray cells that contact developing vessel elements and tracheids, myosin, tubulin, actin and callose are localised at the periphery of developing contact and cross-field pits; the respective antibodies also highlight the bordered pits between vessels and between tracheids. The aperture of the bordered pits, whose diameter diminishes as the over-arching border of these pits develops, also houses myosin, actin and tubulin. Myosin, actin and callose are also found together around the sieve pores of sieve elements and sieve cells. We suggest that an acto-myosin contractile system (a 'plant muscle') is present at the cell plate, the sieve pores, the plasmodesmata within the walls of long-lived parenchyma cells, and at the apertures of bordered pits during their development.