Details of development and the phenology of postdormancy cone-bud development, microsporogenesis, pollen development, and pollination were similar for Pinus contorta var. contorta and var. latifolia growing near Victoria, B.C., but comparable stages of development for var. latifolia occurred about 1 month later near Prince George, B.C. Several developmental aspects were found which affect the reproductive potential of the species. Only 25% of the ovuliferous scales, mostly in the distal part of the cone, bear fertile ovules. Secretions formed on the ovules and micropylar arms which caused pollen to adhere to these surfaces. Pollination is by means of pollination drops which began to be exuded from the ovules about 2 weeks after the conelets began to emerge from their bud scales. Pollination drops were present within each conelet for 2 to 4 days. At that time conelets were most widely open. Pollination drops were then withdrawn as ovuliferous scales enlarged and sealed the conelets. Pollination drop exudation and withdrawal were affected by humidity and water stress within the tree. Cells lining the micropylar canal enlarged and sealed the micropyle after the conelet closed. Pollen settled into a pollen chamber in the nucellus tip where it germinated about 2 months after pollination. Ovules lacking germinating pollen aborted after megasporogenesis and before free nuclear division began. If many ovules aborted within a conelet, the conelet aborted before winter dormancy. Ovules began free nuclear female gametophyte development and pollen tubes extended into the nucellus before conelets stopped developing in mid-August.
The pollination mechanism of Picea engelmannii (Parry) was studied on small potted scions under natural conditions and in controlled environment chambers. Six stages of conelet development were recognized and related to pollen receptivity. Cone-lets appeared receptive for about 2 weeks but were actually receptive for only about 1 week. Secretory droplets appearing on the micropylar arms collected pollen for several days before pollination drops formed. Pollination drops formed acropetally in the conelet and only once from each ovule. Pollination caused rapid recession of the pollination drop, whereas the drop remained for several days on unpollinated ovules. There was some decrease in size of pollination drops during midday and reemergence the following night. Pollination drops were secreted by the nucellar tip in a manner similar to nectaries. Secretory cells collapsed following secretion. The drop contained 4.3% glucose and 3.8% fructose but no sucrose. High humidity increased the longevity and decreased the diurnal fluctuation in size of pollination drops. Conelets from trees with low leaf water potential developed more slowly and produced smaller and more viscous pollination drops. Cones averaged 103 ovuliferous scales, 90% of which were fertile. However, usually less than 50% of the potential seed set was achieved. One of the major causes for low seed set is inadequate pollination. A better understanding of the pollination mechanism and the receptive period may improve seed efficiency in controlled and supplemental mass pollinations.
Postdormancy development of the ovule, fertilization, and embryo and seed development of Pinus contorta (Dougl.) are described. Reproductive development in lodgepole pine (P. contorta var. latifolia) growing within and outside its natural range is described in detail and compared with the phenology of shore pine (P. contorta var. contorta) growing within its natural range. The female gametophyte of lodgepole pine overwinters at the free nuclear stage, resumes development in early April, and is mature by mid-June. Fertilization occurs in mid-June and the embryo and seed are mature by early August. Seed cones of lodgepole and shore pines growing in the coastal region resume development in late March, develop more slowly at first, then complete each subsequent stage of development about 1 month earlier than lodgepole pine grown in the interior of British Columbia. Stages of the reproductive cycle as they affect potential seed production are discussed.
Mature, dry, one-celled pollen was formed before pollen cones became dormant in the fall. Pollen averaged 27 μm in diameter, was irregular in shape, nonsaccate, and the surface was reticulate to tegillate-baculate and irregularly covered with orbicules. The pollen contained several large lipid droplets and no starch. No changes occurred in pollen during dormancy and pollen was shed at the one- or two-celled stage during the last half of March.All ovules were initiated and became flask shaped before seed-cone dormancy. No changes occurred in ovules during dormancy. Seed cones ended dormancy in early March, enlarged and opened, exposing the ovules. A pollination drop was produced by a breakdown of cells at the tip of the nucellus. A large pollination drop was exuded from each ovule in a cone but exudation did not occur at the same time in all ovules. Each ovule exuded and withdrew a pollination drop two to four times before the pollination drop was permanently withdrawn. Each ovule was receptive for a few days and each cone was receptive for about 1 week. The pollination drops were withdrawn in the presence or absence of pollen but were withdrawn more rapidly after pollen entered the pollination drop. The cuticular surface of the bract-scales prevented wetting of the surface and caused the beading of water droplets, which in turn could carry pollen to the micropyle. Pollen grains entering a pollination drop were withdrawn inside the drop into the micropyle. Cells lining the micropylar canal enlarged and sealed the canal while bract-scales enlarged and buried the ovules within the cone.
The development of the pollination mechanism and the engulfment of pollen by the stigmatic tip is described for Douglas-fir (Pseudotsugamenziesii (Mirb.) Franco) based on scanning electron microscopy. This information is used to determine and explain the optimal time of pollination and amount of pollen needed for maximum seed set. After dormancy the integument tip of the ovule developed into an unequally two-lobed stigmatic tip covered with long unicellular hairs. Most ovules had fully developed stigmatic tips when the seed cone emerged from the bud scales in early April. The conelets remained open and the stigmatic tip was most receptive for at least 4 days. Pollen freely sifted down between the bracts and ovuliferous scales and adhered to the stigmatic hairs. Six days after the conelets became receptive, stigmatic hairs around the micropyle began to collapse and were ungulfed with the entangled pollen into the micropyle. Also, ovuliferous scales began to thicken, restricting movement of pollen to the stigmatic tips. By 8–10 days after conelets became receptive, the stigmatic tips were completely engulfed, the ovuliferous scales had thickened enough to close the conelet, and the conelet had begun to bend down.Maximum seed set occurred when (1) cones were pollinated within 4 days after seed-cone buds had emerged half of the way out of their bud scales; (2) a minimum of 0.2 g of pollen was used per pollination bag; (3) a minimum of 11 pollen grains adhered to each stigmatic tip; and (4) at least 3 pollen grains were taken into each micropylar canal. The engulfing process occurred at the same rate and in the same manner regardless of whether living or heat-killed pollen was present or absent on the stigmatic surface. Poor seed set as it related to the pollination mechanism is discussed.
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