Plant Physiol.

(1978) 62, 779-780

Seasonal Variation in the Hormone Content of Willow

II.

EFFECT OF PHOTOPERIOD ON GROWTH AND ABSCISIC ACID CONTENT OF TREES UNDER FIELD CONDITIONS'
Received for publication October 25, 1977 and in revised form June 15, 1978

RONALD ALVIM Centro de Pesquisas do Cacau-CEPLA C, Caixa Postal 7, Itabuna, Bahia, Brazil SHEILA THOMAS AND PETER F. SAUNDERS Department of Botany and Microbiology, University College of Wales, Aberystwyth, Wales
ABSTRACT
Levels of abscisic acid were followed in the xylem sap, mature leaves, and apices of field-grown willow (Salix vihnalis L.) during the summer months, under natural and artificially extended photoperiods. Although the long day treatment prevented the general onset of dormancy, the plants grown under natural daylengths showed lower concentration of abscisic acid than those kept under long days.

leaves were harvested from 20 randomly selected shoots, 12 leaves being taken along the whole length of every stem on each sampling date. Sixty shoot tips were harvested at random on each occasion. Extension growth at the apex was periodically measured in 10 randomly selected shoots per treatment, from May 25 until the plants given extended photoperiods showed significant frost injury, when the experiment was terminated. Quantitative analysis of ABA was carried out by the method described by Alvim (1, 2).

RESULTS The pattern of seasonal changes in ABA levels previously Correlations between high ABA levels and the dormant period in woody plants have recently been found in the xylem sap of detected in xylem sap by Alvim et al. (2) was confirmed, but peach (4) and in buds of apple (6) and birch (5). Marked seasonal maximum concentrations were found earlier, on July 9. ABA fluctuations in ABA content of xylem sap of willow (Salix vimin- levels varied seasonally in all parts of the plant studied. Extracts alis L.) which could also be associated with the stage of dormancy from plants given extended photoperiods had higher ABA conwere reported by Alvim et al. (2). The concentration found in tents than those from the controls, the difference being particularly July, 4 weeks before plants stopped elongating, represents about accentuated in apical tissues (Fig. 1). Time courses of extension growth at the apex in both photoper10 times that found in spring. Leaf buds opened well after ABA iodic treatments and maximum and minimum air temperatures levels started to decrease at the end of December. It is tempting and pertinent to question whether the changes in are shown in Figure 2. Plants under natural photoperiods ceased ABA levels in the summer were caused by the shortening of elongating at the same time of the year as in the previous studies. daylength. It may be relevant that the levels of inhibitory sub- Extended daylengths prevented apex abortion, leaf senescence, stances in the sap extracts began to increase shortly after the and the cessation of extension growth until November 15, when longest day (June 21). Despite the fact that several experiments the experiment was terminated. From May 25 to July 31 the mean carried out under controlled environments (1, 3) did not show any fresh weight per apex for both treatments increased linearly, from evidence of an effect of short photoperiods on the ABA content of 0.26 g in May to 0.59 g at the end of July. On August 20, a willow rooted cuttings, it was decided to attempt to detect possible considerable difference could be detected between the treatments, when apex unit weight was 0.67 g in the plants under extended effects of photoperiodic changes under field conditions. photoperiods and only 0.22 g in the controls. Visible changes in apex size, however, occurred suddenly, within the period of 1 MATERIALS AND METHODS week, in the first days of August. In the course of the experimental period it was observed that Daylength was artificially extended in part of the same row of lateral buds initiated growth following decapitation until July 31 trees used in the previous year in the first part of this work (2). A minimum of 17 hr of daily illumination was applied in this in the controls, whereas in plants under extended photoperiods, treatment, using two fluorescent tubes (6.5 lumen/m at the shoot they continued to burst until mid-September. Even later in the year, in spite of the low temperature, there was a clear increase in tip level) standing 20 to 50 cm above the plants. ABA levels from both long-day-treated and control plants were bud length in response to decapitation and some of them actually periodically determined by GLC in extracts of the apices, mature opened, but were killed by frost. leaves, and sap of willow (S. viminalis L.). Xylem sap was obtained by the method described in the first paper in this series (2). Mature DISCUSSION Lights in the field were inadvertently switched off from July 6 'This paper represents part of the Ph.D. dissertation of R. A., whose to 11, when daylengths decreased from 16:24 to 16:18 hr. For this work was supported by the "Conselho Nacional de Pesquisa" (Brazil), by reason it was not possible to harvest plants under extended phothe Ministry of Overseas Development (Great Britain), and by the Cacao toperiods on July 9, when the peak of ABA occurred. The question Research Center (CEPLAC, Brazil). arises as to whether the highest levels of ABA encountered on this
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Plant Physiol. Vol. 62, 1978

date would also have occurred in plants under extended daylengths. _sop (gg/lOO mi) In spite of the short period of interrupted daylength extension, Apex (ag/100 opices) the effectiveness of the photoperiodic treatment cannot be argued, C2l Leaf (/ug100gf wt1 Extended photoperiods since the plants did not become dormant. Although the influence of photoperiod on the ABA content of willow could not be fully 50 estimated, it can be observed that the levels found in all plants on July 31 were still much higher than those of spring and clearly 40 had not been reduced by long day treatment. On the contrary, SO plants under extended photoperiods had consistently higher ABA contents than the control plants. At least part of the seasonal changes detected cannot be associated with photoperiod. These results reinforce the conclusions of Alvim et al. (3) that '20 ABA levels are not significantly changed by photoperiod and that this hormone may not play a role in bud dormancy. The increased 10 levels found in the apices of plants growing under extended photoperiods probably resulted from higher amounts of leaf ABA reaching the apex due to differences in the rates of leaf fall 20 23 N 9 25 between the two treatments. JUNE r : uf: 1i U T 1EPE MAYj In view of the marked effects of daylength on the growth FIG. 1. Seasonal changes in ABA contents of xylem sap, apices, and behavior of the willow plant and in particular on dormancy of mature leaves of willow plants growing under extended and natural lateral buds, it would be interesting to follow possible seasonal photoperiods. changes in the ABA content of such buds. However, due to the exeso rowt ixtnde wihow pelants udrntrl( adetne large variation in size of lateral buds as summer progresses and l20 also due to the difficulty in distinguishing leaf buds from flower buds at early stages of development, this experiment was not carried out.
E~~~~~~~~~~~~
20 10~~~~~~~~~~~ LITERATURE CITED
1. ALVIM R. 1973 Studies on endogenous hormones in relation to dormancy in woody plants. PhD thesis. Univ Wales, UK 2. ALVIM R, EW HEWErT, PF SAUNDERS 1976 Seaaonal variation in the hormone content of willow. 1. Changes in abscisic acid content and cytokinin activity in the xylem sap. Plant Physiol 57: 474-476 3. ALVIM R, PF SAUNDERS, RS BARROS 1978 Abscisic acid and the photoperiodic induction of dormancy in Salix viminalis L. Plant Physiol. In press 4. DAVISON RM, H YOUNG 1973 Abscisic acid content of xylem sap. Planta 109: 95-98 5. HARRISON MA, PF SAUNDERS 1975 The abscisic acid content of dormant birch buds. Planta 123: 291-298 6. SEELEY SD 1971 Electron capture gas chromatography of plant hormones with special reference to abscisic acid in apple bud dormancy. PhD thesis. Comell Univ, Ithaca, NY

i Lv~~~~~~~~~
0 20~~~~~

photoperiods.

CMay

June

July

August

Sept. October

FIG. 2. Maximum and minimum air temperatures and time courses of extension growth in willow plants under natural (0) and extended (0)

photoperiods.