Bud development in western hemlock. II. Initiation and early development of pollen cones and seed cones

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Seed cones in Tsuga heterophylla (Raf.) Sarg. are found at the tips of distal lateral branches and form as a result of the transition of a previously vegetative apex. Pollen cones may be formed similarly and are then found at the tips of less-vigorous proximal branches but more commonly they develop from newly initiated axillary buds on short proximal shoots. In all cases, apices undergo transition to the reproductive state after a period of bud-scale initiation. Some apices initiate many bud scales, then either initiate leaves or undergo transition to a seed-cone apex in July. Other apices initiate fewer bud scales, then late in June undergo transition to a pollen-cone apex. Transition to a reproductive apex is marked by an increase in mitotic activity and apical size and loss of the vegetative pattern of zonation. Zonation reappears during the slower period of late bract and microsporophyll initiation but is not as prominent as it was in vegetative apices. In seed-cone buds, all bracts, ovuliferous scales, and megaspore mother cells are formed before dormancy. In pollen-cone buds all microsporophylls and microsporangia are initiated before dormancy and pollen mother cells begin meiosis and remain in the diffuse diplotene stage during dormancy. Pollen- and seed-cone buds become dormant in December. The time of cone initiation and sexuality of cones may be influenced by day length. The pattern of reproduction in western hemlock is compared in some respects with that of other conifers.