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Phytochrome Effects in the Nyctinastic Leaf Movements of Albizzia Julibrissin and Some Other Legumes1 2 William S

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Phytochrome Effects in the Nyctinastic Leaf Movements of Albizzia Julibrissin and Some Other Legumes1 2 William S Plant Physiol. (1967) 42, 1413-1418 Phytochrome Effects in the Nyctinastic Leaf Movements of Albizzia julibrissin and Some Other Legumes1 2 William S. Hillman and Willard L. Koukkari Biology Department, Brookhaven National Laboratory, Upton, New York 11973 Received June 5, 1967. Summnary. Participation of phytochrome 'is evident in the nyctinastic responise of leaves of Albizzia julibrissin (silk-tree), Albizzia lophantha, Leucaena glauca, Poinciana gilliesi and Calliandra inequilatera; closure of excised pairs of pinnules upon darkening is rapid following red illumination and slow following far-red. Under good conditions the difiference is obvious within 10 minutes. These observations conifirm a report by Fondeville, Borthwick, and Hendricks on the sensitive plant, Mimosa pudica, but indicate that the efifect bears no necessary relationship to the anomalous sensitivity of Mimosa. In A. julibrissin, phytochrome control is mnarked in experiments conducted early in the daily 12-hour light period and appears absent, or nearly so, toward the end of the light period, perhaps due to interaction with an endogenous circadian rhythm. Effects of leaf maturity and of the position of a pinnule-pair within a leaf are also evident. Tih-ese results are not easily reconciled with hypotheses of phytochrome action through gene activation and nucleic acid synthesis, but are consistent with hypothess ibased onl permeability changes and membrane properties. The mgnitude and reproducibility of the response in A. jutlibrissin suggest its use as a lajboratory exercise; this and related systems should prove valuable for eventuai identification of the mechanism of phytochrome action. Fondeville, Borthwick, and Hendricks (2) re- pinnately twice-compound leaves generally similar in ported on a role of phytochrome in the nyctinastic character to those of Mimosa pudica, (but not obviously response of the sensitive plant, Mimnosa pudica: closure sensitive to the touch. Plants wiith 6 or more leaves of the pinnules upon darkening was rapid when the were moved at least 3 days before use to chambers darkness 'followed a brief red illumination and slower with 12 hours per day of cool white fluorescent light if the darkness followed far-red, with appropriate (about 900 ft-c, with a small incandescent supple- reversibility. The major significance of this report ment) at about 240. led us to see wrhether such effects could be obhserved In early experiments single pairs of pinnules were in plants other than the relatively anomalous Mimosa, cut with, 1 to 2 cm of raohilla, the end of which was our imniedFate objective being an experimental system placed in distilled water-agar in a covered staining more stable and quantitative than that orig.nally de- dish. These preparations were left in the chambers scribed. This objective has now been attained well with the 'plants until the following day, at which time enough to provide, among other things. a rapid and the experiment was condlucted. In later experiments, dependable elenentary laboratory exercise. indtividual pinnule-pairs were cut with only that portion of the rachilla (2-4 cm) to which they were attached Materials and Methods (7), floated in 2 ml of distilled water in a 5 cm petri dish, and the experiment started withiin 30 min- Albizzia julibrissin (silk-tree), A. loplianttha, Leu- utes of cutting. In the agar experiments, pinnule caena glauca, Pointciana gilliesi, and Calliandra in- closing was followed by measuring the aperture tin equilatera were propagated iin the \varm greenhouse nmmni between the tips of the pinnules, while in the from seeds and cuttings. These species all have water experiments the angle between the anterior edges of the pinnules was measured to the nearest 100 by lifting each pair with a forceps and moving it over a set of standards affixed to transparent plastic 1 Research carried out at Brookhavcn National Lab- (7). The damage inevitably done by such handling oratory under the auspices of the United States Atomic seems to have no effect on the 'response. All opera- Energy Commission. tions after the start of a dark treatment were con- 2 A report of this work was presented at the annual meeting of the Northeastern Section. ASPP, 'May 5 ducted in a dark room with minimal exposuire to a and 6, 1967 at Cambridge, MWassachusetts. dim green safe light (3-5). 1413 1414 PLANT PHYSIOLOGY Red light was obtained from a cool white fluores- cent tube separated from the tissue by a one-eighth inch thickness of Rohm an-d Haas Red 2444 "Plexi- PET IOLE glas" and, far-red from an incandescent spotlight behind at least 6 cm of water and a one-eighth inch thickness of Rohm and Haas "Black" FRF-700 "Plexiglas" (4). Intensities of the fluorescent and incandescent sources without the filters in place were respectively about 220 and 6000 foot candles. Illumi- nation constisting of various iproportions, of red and far-red to bring about various phlotostationarv states of phytochrome was provided by incandescent spot- lights behind at least 8 cm of water, red "Plexiglas," and an appropriate glass filter. As previouslv cali- brated by direct spectrophotometric assays of etiolated DI seedling tissue (3, 5) the sources used establish ap- proximately the following photostationary states (as percentage PFR) in 5 minutes: the standard red source, 88 to 92; incandescent with red "Plexiglas" C I only, 64; the same plus Corninz filter 1-61, 44; the same except with filter 4-77, 20: the same except with filter 1-64, 12; and the far-red plastic behind anv incandescent soutrce, 5. B' Results Al All the work described deals with A. julibrissin, A except as specifically mentioned. Initial attempts to FIG. 1. Diagramii of a 10-pinnate leaf of AlbizZia observe phytochrome control of nyctinasty gave ex- jullibrissint or a similar plant. tremnely variable,results. This variability was reduiced after several maior factors affecting nvctinastv were pinnae, bearing pinnules), and sonie pinnules. The identified, as indicated below. 10 pinnae are in 5 symmetrical pairs AA' to EE'; Rate of Closing at Various Tinies During the leaves used in various experiments have had from 2 Light Period. With plants held on a schedule of to 14 pinnae. The number of pinnules on various 12 hours li!ht-12 hours darkness, the rapiditv of portions of a leaf of the kind diagrammed might be, n,yctinastic closing upon darkening is greatest late in e.g., 18 to 25 for each pinna A(A') thru C, 12 to 18 the light period and least in its early hours. In an for I) and 7 to 14 for E. agar experiment, transfers to darkness were made at An agar experiment compared the phytochrome intervals from 'hour 2 of the light period to hour l1. response of pinnule-pairs from di'fferent pinnae and Effects of actual times of day were eliminated by from different positions wvithin each pinna.' Using simultaneously replicating the experiment on material figure 1 for orientation, pinnule-pairs I and II were g,rown in 2 chambers with schedulles set 6 hours taken from pinnae B, C, and D and their opposites. apart. With pinnule-pairs having initial apertures of Pinnule-pairs I and, II were the sixth and twelfth about 22 nmm, the decrease in aiperture (closing) from the base, respectively (n - m 6). Pinnule- produced by 10 miinutes of darkness averaged about pairs BI, BII .... DII received 2 minutes of red 2 mm at houtr 2 of the light period and' increased light before the (larkness whiile the corresponding in apparently monotonic fashion to about 14 at hour pairs of B'I . D'II received far-red. The results, 11. Such results show only that nyctinasty is more in table Ia, allow 3 conclusions. First, there is a rapid as the light iperiod proceeds; subsequent experi- clear phytochrome effect, closing being much more ments, however, also indicate an interaction between rapid with red than with far-red pretreatment. Sec- the time of the light period and the phytoclhrome ond, there is a clear position effect. wvith the II response. (twelfth) pairs closing more slowly than the more Effect of Positioni Within the Leaf Structure. basal leaflets. Third. there seems to be no pinlia When whole iplants are darkened, nyctinastic closure effect (e.g., B vs C vs D). Table lb, representing proceeds at obviously different rates in different an experiment as precisely similar as possible bitt done leaves and in different portions of the same leaf. later (hr 10 instead of hr 1) in the light period indi- For example, within a leaf, terminal leaflets generally cates vividly that closinig is faster at this time. In close more slowly than the others. Figure 1 is a addition, no phytochrome response is evident, though diagram of a 10-pinnate leaf indicating the rachis one could not conclude on the basis of these data (main axis, bearing pinnae), rachillae (axes of the alone that it is necessarily absent, since the rapidity HILLMAN AND KOUKKARI-PHYTOCHROME EFFECTS IN NYCTINASTIC MOVEMENTS 1415 Table I. Response to Darkness of Pinnule-Pairs from A. julibrissin Preilluininiated with Red (R) or Far-Red (F) Refer to figure 1 for orientation. Aperture is distance between pinnule tips in mm. Rachillae imbedded in agar, excised the day before use. Change in aperture at indicated time after start, following R or F 30 Min 180 Min Pinnae Pinnule-pairs Initial aperture R F R F A. Experiment started at hr 1 of light period B,B' I 19, 19 -4 0 -13 0 II 21, 23 -2 0 -7 0 C,C' I 18, 20 -3 -1 -12 -1 II 21, 22 -2 -1 -6 0 D,D' 1 18, 19 - 0 -13 +1 II 18, 21 -2 -1 -6 0 B.
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