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

1974 ◽  
Vol 52 (2) ◽  
pp. 283-294 ◽  
Author(s):  
John N. Owens ◽  
Marje Molder

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.


1984 ◽  
Vol 62 (3) ◽  
pp. 484-494 ◽  
Author(s):  
John N. Owens

Seed cones of Tsuga mertensiana (Bong) Carr. occur terminally on distal lateral branches and form from the differentiation of a terminal, previously vegetative apex, into a seed-cone apex. Pollen cones commonly occur on lateral branches and form from the differentiation of an undetermined axillary apex about 6 weeks after axillary bud initiation. Pollen cones also occasionally occur terminally. All cone buds began differentiation in late July after bud-scale initiation was complete and at about the end of lateral shoot elongation. Seed-cone buds initiated bracts and ovuliferous scales, but not ovules, before they became dormant at the end of October. Pollen-cone buds initiated all microsporophylls by early September. Microsporangia containing microspore mother cells differentiated before pollen-cone buds became dormant in mid-October. The time of cone-bud differentiation is related to vegetative bud and shoot development. The time and method of cone-bud differentiation is discussed in relation to T. heterophylla and other conifers having similar bud development.



1974 ◽  
Vol 52 (9) ◽  
pp. 2075-2084 ◽  
Author(s):  
John N. Owens ◽  
Marje Molder

Vegetative shoots initiate leaves and lateral branches continuously from mid-April until the end of September. No buds with bud scales are formed and the vegetative apex is enclosed by leaf primordia at various stages of development. Pollen cones are initiated on proximal vegetative shoots during a 3-week period from mid-June to early in July. Transition to a pollen-cone apex is marked by an increase in mitotic activity in the apex and by the formation of a lateral branch in the axil of one of the last-formed leaf primordia, causing the apex to appear to branch dichotomously. The lateral branch remains at the base of the pollen cone and may resume growth the next year after the pollen cone is shed. Pollen-cone development continues until the end of September. Meiosis occurs during the last 2 weeks of August, and pollen develops during September. Seed cones are initiated on newly formed, distal axillary vegetative shoots during a 3-week period from late June to mid-July. Transition to a seed-cone apex is marked by an increase in mitotic activity followed by bract-scale initiation. Usually three ovules are initiated in the axil of each bract scale. Seed-cone development is complete by early September and the seed cones become dormant. The pattern of reproduction in yellow cedar is compared to other conifers and the possible relationships are discussed between time of cone initiation, sexuality of cones, and day length.



1976 ◽  
Vol 54 (8) ◽  
pp. 766-779 ◽  
Author(s):  
John N. Owens ◽  
Marje Molder

Pollen-cone and seed-cone buds of Picea sitchensis (Bong.) Carr. are found as either terminal or axillary buds. Pollen cones are most likely to develop from small axillary apices on vigorous distal shoots or small terminal apices on less vigorous, proximal shoots. Seed cones are most likely to develop from large, distal axillary apices on vigorous shoots or smaller terminal apices on less vigorous shoots. All apices became mitotically active late in March, passed through a 3.5-month period of bud-scale initiation, and in mid-July became differentiated as vegetative, pollen-cone, or seed-cone apices. Potentially pollen-cone apices were smaller, had a lower mitotic frequency during bud-scale initiation, and produced fewer bud scales than apices which developed into seed-cone or vegetative buds. During bud-scale initiation all apices had a few strands of cells containing phenolic compounds in the developing pith. At the time of bud differentiation, the pith of vegetative apices accumulated more phenolic compounds and non-phenolic ergastic materials, whereas the pith of reproductive apices did not. This was followed by a marked increase in mitotic frequency in reproductive apices, resulting in changes in apical size and shape. Leaf, bract, and microsporophyll initiation began about the end of July. All microsporophylls were initiated by the end of August. Sporogenous cells developed, but meiosis did not occur before the pollen cones became dormant at the end of October. Two-thirds of the bracts were initiated by the end of August. The remaining bracts were initiated more slowly until dormancy. Ovuliferous scales were initiated for 3 months beginning in September, and megaspore mother cells appeared but did not undergo meiosis before seed cones became dormant at the end of November. There was no difference in the time of vegetative, pollen-cone, and seed-cone bud differentiation, which occurred at the end of lateral shoot elongation.



1989 ◽  
Vol 19 (11) ◽  
pp. 1422-1428 ◽  
Author(s):  
R. C. Bower ◽  
S. D. Ross ◽  
B. G. Dunsworth

Experiments on field-grown and potted trees investigated the effect of timing of gibberellin A3 treatment in relation to natural day length on flowering and sex expression in Chamaecyparisnootkatensis. When gibberellin A3 treatment began was more important than the number of weekly sprays (two to eight), and the optimal time of treatment was similar for promotion of seed and pollen cone buds. The optimal treatment time was independent of photoperiod or its direction of change, but instead appeared to be related to a specific stage of ontogeny during early development of the bud apex. Trees growing at progressively lower elevations attained this stage earlier in spring, suggesting that the mechanism triggering initiation of bud development involved heat sums. A higher concentration of gibberellin A3 was required to initiate seed cone buds (200 mg•L−1) than pollen cone buds (100 mg•L−1), and the average weekly dosage was more important than whether gibberellin A3 was applied weekly or biweekly. Young, potted trees within a greenhouse flowered more profusely than larger but similarly treated field-grown trees, indicating that the species is a good candidate for containerized seed orchards.



1971 ◽  
Vol 49 (7) ◽  
pp. 1165-1175 ◽  
Author(s):  
J. N. Owens ◽  
R. P. Pharis

Both seed and pollen cones in Thuja plicata Donn. are found at the tips of small lateral branches and form as a result of the transition of previously vegetative apices. The vegetative apex shows a cytohistological zonation similar to that found in other Cupressaceae and vegetative buds are not enclosed in scales. The first sign of pollen cone initiation occurred 13 days after the initial GA3 treatment under long days. Transition of the apex was marked by a slight increase in rate of cell division in all zones resulting in the formation of a mantle of small uniform cells several cells thick covering the surface of the apex. The apex increased in height and the long internode between the last-formed pair of leaf primordia and first pair of microsporophylls formed a short stalk at the base of the cone. The transition of a vegetative apex to a seed cone apex occurred 27 days after initial GA3 treatment and was similar to that for pollen cones. Bract primordia are initiated closer to the summit of the apex and are larger in initial stages than microsporophyll primordia, and longer internodes remain between successive pairs of bracts. Ovule initiation occurs when bracts have just begun upward growth. All microsporophylls, bracts, and ovules are formed during the few weeks after cone initiation and before cones become dormant. No anatomical changes occurred in either cone type during subsequent short-day cold treatment. A return to long-day warm conditions promoted normal cone development. Under natural conditions in the Victoria area, pollen cones are initiated early in June under long days and increasing day length while seed cones are initiated early in July under similar long days but decreasing day length. Development of both pollen and seed cones occurs during long days and decreasing day length. The possible relationship between sexuality of cones, gibberellin, auxin, and day length is discussed.



1979 ◽  
Vol 57 (14) ◽  
pp. 1557-1572 ◽  
Author(s):  
John N. Owens ◽  
Marje Molder

The time and method of cone-bud differentiation and the phenology of cone-bud development were studied in 10- to 20-year-old trees growing outside their natural range and three 50-year-old trees growing within their natural range.Both pollen-cone and seed-cone buds of western larch (Larix occidentalis Nutt.) normally differentiated on short shoots that were at least 1 year old. Pollen-cone buds were commonly on proximal nonvigorous, often pendant vegetative long shoots in lower regions of the crown, whereas seed-cone buds were usually found on distal short shoots on vigorous but less pendant vegetative long shoots in upper regions of the crown.All potential cone buds were indistinguishable from potential vegetative short shoot buds during bud-scale initiation. In early June, when vegetative short shoots had begun to initiate leaves, cone-bud apices entered a period of differentiation during which time the mitotic frequency of the apices greatly increased followed by a marked increase in apical size. During differentiation, pollen-cone apices did not initiate any basal foliar organs and a short stalk resulted at the base of the cone, whereas seed-cone apices initiated a few basal foliar primordia before bract initiation began. Microsporophyll initiation began during the last half of June and initiation occurred rapidly until the end of July. Micros porangial development occurred from August to late October when fully developed pollen-cone buds became dormant. Pollen mother cells began meiosis before dormancy and overwintered at the diffuse stage. Bract initiation began about the end of June, was rapid until mid-August, then continued more slowly until seed-cone buds became dormant in late October. Ovuliferous scales were initiated acropetally from mid-August until dormancy. Cone-bud differentiation occurred at about the end of the period of vegetative lateral long shoot elongation at all locations.



1989 ◽  
Vol 19 (8) ◽  
pp. 1051-1058 ◽  
Author(s):  
John N. Owens ◽  
Anna M. Colangeli

Cone buds were induced on container-grown and field-grown western hemlock (Tsugaheterophylla (Raf.) Sarg.) clones during a 3-year period to study the effects of time and duration of gibberellin A4/7 treatment on cone induction, sexuality of cones, and to relate these results to bud and shoot development. The most effective treatment times preceded anatomical differentiation. The most abundant pollen cones and seed cones were produced when trees were sprayed with gibberellin A4/7 before vegetative bud burst and early shoot elongation. Two to three weekly gibberellin A4/7 applications starting at preswollen and swollen-bud stages were adequate for pollen-cone production. Pollen-cone production decreased when the applications were started at vegetative bud burst or during early shoot elongation. A minimum of three weekly applications were required for seed-cone production, and applications were equally effective when started at preswollen, swollen, and vegetative bud burst stages. Seed-cone production decreased when three weekly applications were started during early shoot elongation; however, this was overcome by increasing the number of applications.



1991 ◽  
Vol 21 (6) ◽  
pp. 788-793 ◽  
Author(s):  
Derek L. S. Harrison ◽  
Mike U. Slee

Gibberellin A4/7 was applied to field-grown grafts of sexually mature poor and good flowering clones of Pinuscaribaea Morelet var. hondurensis. Barr. & Golf. Continuous applications of gibberellin A4/7 over a 6-week period significantly enhanced seed-cone production in poor flowering clones but had no effect on good flowering clones. Poor flowering clones also showed slight decreases in pollen-cone production with gibberellin A4/7, while clones that normally flowered well did not. Gibberellin A4/7 increased final shoot lengths in the lower (male) and upper (female) crown region of both poor and good flowering clones. A series of timed gibberellin A4/7 applications suggested that a single treatment near the onset of reproductive bud development may be used to enhance seed-cone production.



1988 ◽  
Vol 66 (5) ◽  
pp. 907-914 ◽  
Author(s):  
Anna M. Colangeli ◽  
John N. Owens

Pollen-cone development, which was divided into nine phenological stages, was compared with pollen cytology in eight field-grown western hemlock (Tsuga heterophylla (Raf.) Sarg.) clones in 1983 and three container-grown clones in 1984. Phenology proved to be an accurate indicator of cytology, independent of collection dates and rate of development. The breaking of dormancy, the resumption of development, and meiosis occurred during stage 1 (quiescent bud). During stage 2 (swollen pollen-cone bud) the tetrad of microspores separated. The exine wall was completely developed during stage 3 (bud burst). The one-celled microspores began to expand as a result of accumulation of reserves during stage 4, when the pollen cones were one-quarter to one-half emerged through the bud scales. The first cell division occurred during stage 5, when the cones were greater than half emerged. The second and third cell divisions occurred during stage 6, after the cones had completely emerged through the bud scales. The fourth and final cell division, resulting in mature five-celled pollen grains occurred during stage 7 (stalk elongation). Mature pollen consisted of two prothallial cells and a stalk, a body, and a tube cell. Pollen shed occurred at stage 8. The empty cones (stage 9) remained on the trees until the following winter. Some practical implications of relating pollen-cone phenology to cytology are discussed.



1988 ◽  
Vol 18 (1) ◽  
pp. 43-53 ◽  
Author(s):  
Conor O'Reilly ◽  
John N. Owens

Reproductive growth and development were studied in 1983 in seven provenances of Pinuscontorta Dougl. ssp. latifolia Engelm. growing in a provenance trial near Prince George, B.C. Stages of pollen release and seed-cone receptivity were scored by indices of cone development. Pollen- and seed-cone numbers were estimated and the distribution of seed cones within the upper crown and on annual growth increments of fourth-whorl branches was assessed. Pollen-and seed-cone bud development was followed in sectioned long-shoot buds taken at 2- to 3-week intervals. The times of maximum seed-cone receptivity and pollen release differed slightly among provenances, indicating that there was a high chance of cross-pollination. Differences among provenances in pollen-cone numbers were large, but smaller differences in seed-cone numbers were noted. No mature pollen cones or developing pollen-cone buds were found in the Yukon provenance. Seed-cone production varied with whorl position and was influenced by polycyclic long-shoot development. Potential pollen-cone buds were initiated from May until late June. Pollen cones first differentiated in early to mid July in all provenances. Potential seed-cone apices were noted from mid-June to late July and differentiation occurred in mid-July to early August, depending on provenance. Seed-cone bud development began first in the northern provenances.



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