Hormonal control of second flushing in Douglas-fir shoots.

Spring-flushing, over-wintered buds of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) produce new buds that may follow various developmental pathways. These include second flushing in early summer or dormancy before flushing during the following spring. Second flushing usually entails an initial release of apical dominance as some of the current-season upper lateral buds grow out. Four hypotheses concerning control of current bud outgrowth in spring-flushing shoots were tested: (1) apically derived auxin in the terminal spring-flushing shoot suppresses lateral bud outgrowth (second flushing); (2) cytokinin (0.5 mM benzyladenine) spray treatments given midway through the spring flush period induce bud formation; (3) similar cytokinin spray treatments induce the outgrowth of existing current lateral buds; and (4) defoliation of the terminal spring-flushing shoot promotes second flushing. Hypothesis 1 was supported by data demonstrating that decapitation-released apical dominance was completely restored by treatment with exogenous auxin (22.5 or 45 mM naphthalene acetic acid) (Thimann-Skoog test). Hypothesis 2 was marginally supported by a small, but significant increase in bud number; and Hypothesis 3 was strongly supported by a large increase in the number of outgrowing buds following cytokinin applications. Defoliation produced similar results to cytokinin application. We conclude that auxin and cytokinin play important repressive and promotive roles, respectively, in the control of second flushing in the terminal spring-flushing Douglas-fir shoot.

A possible role of cytokinin in mediating long-distance nitrogen signaling in the promotion of sylleptic branching in hybrid poplar.

Nitrogen fertilization of roots enhances shoot growth in plants and cytokinins are known to initiate bud outgrowth in shoots. Is it possible that root-derived cytokinins may play a role in long-distance signaling for nitrogen availability in the promotion of sylleptic branching in hybrid poplar? Nitrogen fertilization in the form of 5 mM NH4NO3, KNO3 or NH4Cl was applied to roots of three hybrid poplar clones exhibiting contrasting degrees of sylleptic branching. Cytokinin (0.1-1 mM benzyladenine, BA) was applied directly to lateral buds of shoots. Glutamate, asparagine and glutamine were also applied as drops to buds or as foliar sprays. NH4NO3, KNO3 and NH4Cl all usually enhanced sylleptic branching within a week in the high sylleptic clone (11-11) but in four out of five trials there was no effect in the low sylleptic clone (47-174). NH4NO3 added directly to buds had no effect. Also, glutamate, asparagine and glutamine had no effect. However, 1 mM BA promoted lateral bud outgrowth in all three clones. These results are consistent with the long-distance nitrogen signaling hypothesis of Forde and Sakakibara wherein nitrogen is transduced to cytokinin via enhanced ipt activity in the roots and is translocated up the shoot with the subsequent promotion of leaf/bud outgrowth.

A reappraisal of the role of abscisic acid and its interaction with auxin in apical dominance.

BACKGROUND AND AIMS: Evidence from pea rms1, Arabidopsis max4 and petunia dad1 mutant studies suggest an unidentified carotenoid-derived/plastid-produced branching inhibitor which moves acropetally from the roots to the shoots and interacts with auxin in the control of apical dominance. Since the plant hormone, abscisic acid (ABA), known to inhibit some growth processes, is also carotenoid derived/plastid produced, and because there has been indirect evidence for its involvement with branching, a re-examination of the role of ABA in apical dominance is timely. Even though it has been determined that ABA probably is not the second messenger for auxin in apical dominance and is not the above-mentioned unidentified branching inhibitor, the similarity of their derivation suggests possible relationships and/or interactions. METHODS: The classic Thimann-Skoog auxin replacement test for apical dominance with auxin [0.5 % naphthalene acetic acid (NAA)] applied both apically and basally was combined in similar treatments with 1 % ABA in Ipomoea nil (Japanese Morning Glory), Solanum lycopersicum (Better Boy tomato) and Helianthus annuus (Mammoth Grey-striped Sunflower). KEY RESULTS: Auxin, apically applied to the cut stem surface of decapitated shoots, strongly restored apical dominance in all three species, whereas the similar treatment with ABA did not. However, when ABA was applied basally, i.e. below the lateral bud of interest, there was a significant moderate repression of its outgrowth in Ipomoea and Solanum. There was also some additive repression when apical auxin and basal ABA treatments were combined in Ipomoea. CONCLUSION: The finding that basally applied ABA is able partially to restore apical dominance via acropetal transport up the shoot suggests possible interactions between ABA, auxin and the unidentified carotenoid-derived branching inhibitor that justify further investigation.

Is auxin the repressor signal of branch growth in apical control?

"Apical control" is the repression of branch growth by a higher dominating branch or shoot. There has been some confusion in the literature concerning the meaning and causal mechanisms of this correlative phenomenon with those of "apical dominance," which term is often used in a strict sense to connote the repression of the initiation of axillary bud outgrowth by an active shoot apex. Although the term "apical control" is most commonly employed with respect to woody species, this phenomenon also widely occurs in herbaceous plants. Because of the strong evidence for a role of auxin as a repressor signal in apical dominance and partly because of this lack of distinction in terminology, a similar role for auxin in apical control is often assumed in spite of the obvious acropetal auxin transport difficulty and the lack of direct evidence for the acropetal transport of any inhibitor influence. In the present study with the herbaceous Ipomoea nil, it has been clearly demonstrated that while exogenous auxin (1% NAA) strongly restores apical dominance in the Thimann-Skoog experiment, auxin treatments to decapitated dominant shoots do not, in any observable way, restore apical control in lower dominated branches. Hence, in this fast-growing species, the hypothesis for the role of auxin as a repressor signal for apical control is not supported.

A preliminary investigation of the role of auxin and cytokinin in sylleptic branching of three hybrid poplar clones exhibiting contrasting degrees of sylleptic branching.

Sylleptic branches grow out from lateral buds during the same growing season in which the buds are formed. This type of branching is present in poplar and in many tropical species. It results in the production of more branches, more leaves and expanded photosynthetic capacity and is thought to assist in increasing the overall growth and biomass of the tree at a young age. However, very little is known about the physiology of sylleptic branching in poplar, which is an extremely important source of fibre and fuel. In the present study of three hybrid poplar clones (11-11, 47-174 and 49-177) of Populus trichocarpa x P. deltoides exhibiting contrasting degrees of sylleptic branching, an analysis was carried out on parent shoot elongation and sylleptic branching, together with a preliminary comparison of the parent shoots' sensitivity to auxin (naphthaleneacetic acid) as a repressor of lateral bud outgrowth, and cytokinin (benzyladenine) as a promoter. Suggestive evidence was found for an inverse correlation between parent shoot sensitivity to auxin and the degree of sylleptic branching, as well as a partially positive correlation with respect to sensitivity to cytokinin. The present data are consistent with the hypothesis that auxin and cytokinin may play repressive and promotive roles, respectively, in the sylleptic branching of hybrid poplar.