Sexual reproduction of Larix occidentalis

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Abstract
Pollen-cone and seed-cone buds broke dormancy about 2 weeks before vegetative buds on the same tree. Pollen mother cells, which had over-wintered at pachytene or the diffuse stage of meiosis, resumed meiosis and tetrads of microspores were formed by mid-March. Wingless five-celled mature pollen developed by mid-to late April when pollination occurred.When development resumed after dormancy a ring of meristematic tissue formed the integument around the nucellus. The integument tip developed a short abaxial tip and a large adaxial lobe on which developed numerous long stigmatic hairs. A slit-like micropyle remained between the two lips. Several pollen grains usually adhered to the stigmatic hairs and then the two lips grew into the micropyle, engulfing the pollen. No pollination drop was observed. Within the micropylar canal, pollen greatly elongated then formed a pollen tube when the elongated pollen contacted the nucellus.Megaspore mother cells underwent meiosis at the time of pollination. Female gametophyte development, which was the same as in most other members of the Pinaceae, was completed in early June and two to five archegonia were formed. Fertilization occurred in early June, 6 to 8 weeks after pollination. A 16-celled proembryo developed. Simple polyembryony was common but cleavage polyembryony was not observed. Embryo development was similar to other members of the Pinaceae. Embryos and seeds were mature by mid-August.Normal appearing but inviable seed is common in L. occidentalis because the ovule is fully enlarged and the seed coat well developed at fertilization. Inviable seed commonly resulted from the absence of pollination, inviable pollen, lack of fertilization, later ovule abortion, or embryo abortion, primarily during early embryonic stages. Flat empty seed also occurred and resulted from abortion of the megaspore mother cell or early female gametophyte.