Flower initiation of Lemna perpusilla under continuous low-intensity light

Lemna perpusilla 6746, a short-day plant, flowered under low-intensitywhite light (10 lux)irrespective of the photoperiod. Red lightof about 20 ergs/cm²/sec also permitted flowering under continuousillumination. The effect of the low-intensity light employedwas not equivalentto that of darkness but similar to that ofblue or far-red light in photoperiodic system. (Received June 15, 1973; )     

Photoreversibility of flower initiation in Lemna perpusilla as affected by the light quality of the main light period

Reversible floral responses of Lemna perpusilla to red and far-redlights appeared only at the beginning of the inductive darkperiod when the 8 hr photoperiod consisted of white or red light.When blue or far-red light was given during the 8 hr photoperiod,the far-red given at the beginning of the dark period scarcelyinhibited flowering; red/far-red reversibility newly appearedat the middle of the dark period. This indicates that the photoregulationsystem in the flowering of L. perpusilla can be converted fromthe Pharbitis type to the Xanthium type by changing the lightquality of the main photoperiod from white or red to blue orto far-red, which is known to be effective for the so-calledhigh-energy photoreaction of photomorphogenesis. (Received July 2, 1975; )     

RED LIGHT, BLUE LIGHT, AND COPPER ION IN THE PHOTOPERIODIC CONTROL OF FLOWERING IN LEMNA PERPUSILLA 6746

L. perpusilla, 6746 grown in HUTNER's medium (containing a highlevel of EDTA) responds as a short-day plant under white orred light, but is almost completely daylength-indifferent underblue. In a medium containing ca. 2µmoles/liter of unchelatedcopper ion, it flowers rapidly under all photoperiods, white,blue or red. Although certain long-day blue schedules permitflowering even on HUTNER's medium, blue is not simply "inactive"photoperiodically, since inclusion of a single brief red exposurein certain blue schedules is sufficient to prevent floweringas though the whole schedules were red. The indicated blue-redinteraction should prove important for further analyses of photoperiodismin this plant. ¹Research carried out at Brookhaven National Laboratory underthe auspices of the U.S. Atomic Energy Commission.     

Dark reactions in the flowering ofLemna perpusilla 6746

Summary This study concerns the effects of red and far-red light on flowering in the short day plantLemna perpusilla 6746. The critical day length for maximum flowering was found to be 10 hours. Exposure to red light near the middle of the dark period inhibited flowering, and the time of maximum sensitivity to red light occurred 9 hours after the beginning of dark periods of either 14 or 17 hours. The inhibition by red light was not reversible by far-red light, which also inhibited flowering, especially when given early in the dark period. Flowering inhibited by exposure to far-red light at the beginning of the dark period could be restored by subsequent exposure to red light. It appears that two photoperiodic partial processes in some plants may be controlled by the red, far-red reversible pigment system.With 5 Figures in the Text     

Inhibitory Action of Blue and Far-Red Light in the Flowering of Lemna paucicostata

In a new strain of short-day duckweed (Lemna paucicostata T-101), blue and far-red light-induced inhibition of flowering was investigated. Flowering of this strain failed to be induced under a short-day photoperiod of blue and far-red light, although it responded as a typical short-day plant in red and white light. When the short-day photoperiod of blue or far-red light was terminated by a 15 min red light pulse, flowering recovered completely. This inducing effect of red light was reversed by subsequent exposure to far-red light. Furthermore, it could be demonstrated that 30 min of blue light completely reversed the flowering inductive effect of 5 min red light and vice versa. Evidence is presented suggesting that the inhibitory action of blue and far red light may be due to the lowering of phytochrome P_fr levels below those required to start the dark reactions which lead to flowering. These results are discussed in relation to the time measurement system of photoperiodism.     

Photoinduction of Spore Germination in a Fern, Mohria caffrorum

Irradiation of spores of the fern Mohria caffrorum Sw. witheither red light (67.4 µW cm^–2) or far-red light(63.2 µW cm^–2) for a period of 24 h induced maximumlevels of germination. Brief irradiations with blue light (127.6µW cm^–2) administered before or after photoinductioncompletely nullified the effects of red or far-red light; however,with prolonged exposure to blue light, germination levels roseto near maximum. The similar effects of red and far-red lightin promoting spore germination makes the involvement of phytochromein this process questionable. Based on energy requirements,the promotive and inhibitory phases of blue light appear toinvolve independent modes of action. Mohria caffrorum, ferns, spore germination, photoinduction, phytochrome     

Lack of flowering responses to DGMU in a mutant of Lemna perpusilla 6746

DCMU, in a sucrose supplemented medium, promoted short and longday flowering and inhibited long day frond production of wildtype Lemna perpusilla 6746, but not of mutant strain 1073. Resultssuggest a defect in the mutant that mimics DCMU poisoning. ¹ This work was supported by National Science Foundation GrantGB-12955. (Received December 11, 1972; )     

The relationship between red and far-red light on flowering of the long-day plant, Lemna gibba

Lemna gibba, a long-day duckweed, can be induced to flower whenthe 10 hr white photoperiod is extended with red or far-redlight. The 10 hr red photoperiod is also effective in inducingflowering when followed by a far-red extension, but a red extensionis ineffective. When 2 hr of far-red light are given immediately after the 10hr red photoperiod, the following red as well as the far-redextension can induce flowering, indicating that the 2 hr far-redlight plays an important role as a starting factor for induction.This red or far-red extension is effectively replaced by a redbreak given at a proper time in the darkness which follows the2 hr far-red light as the starting factor. The effect of thered break in not cancelled by subsequent exposure to far-red,which synergistically promotes flowering. However, a red break given immediately after a proper periodof far-red extension further promotes flowering. The phase sensitiveto the red break coincides with that sensitive to the red breakgiven in darkness. The effect of the red break is reversed bysubsequent exposure to far-red, contrary to the effect of thered break in darkness. Using these results, relation between red and far-red lighton flowering in L. gibba is discussed. (Received July 17, 1971; )     

Flowering responses of Lemna perpusilla and L. gibba in relation to nitrate concentration in the culture medium

Flowering responses of Lemna perpusilla strain 6746, a short-dayplant, and L. gibba strain G3, a long-day plant, to nitrateconcentration in Hoagland's type medium with or without EDTA,were compared. Maximum flowering of L. perpusilla under SD occurredat higher nitrate concentrations than did colony proliferation.Even under CL, L. perpusilla grown at sub-optimal nitrate concentrationsfor colony proliferation, flowered irrespective of the presenceof EDTA which reduces flowering. Unlike L. perpusilla, L. gibba failed to flower under SD atany nitrate concentration whether or not EDTA was added. UnderCL, however, L. gibba flowered at almost any nitrate concentrationwith or without EDTA. Double optima for nitrate concentrationwas exhibited in the presence of EDTA; optimal concentrationfor colony proliferation came between the two optima for flowering. We concluded that the nitrogen level of the medium is importantin regulating flowering of duckweeds, and that the effect ofEDTA, if any, may primarily be on colony proliferation and onlysecondarily or antagonistically on flowering. ¹ Present address: Institute for Agricultural Research, TohokuUniversity, Sendai 980, Japan. (Received September 25, 1971; )     

Effect of far-red light pulse on induction and production of flowers in Lemna paucicostata 6746 in darkness

A short-day duckweed, Lemna paucicostata 6746, was exposed tocontinuous darkness at 26^?C, and the changes in the floral parameters(3) due to far-red and/or red light pulse given at various timesof the dark period were studied. Parameters a (vegetative growth rate) and (flowering ratio)were respectively decreased and increased with a far-red lightpulse given at the outset of the dark period. The decreaseda and the increased remained almost unchanged until the 7thhour, but returned to their initial levels thereafter. The far-redlight actions on a and were reversed by subsequent exposureto red light. Parameter P₁ (pre-flower induction period) wasextended by 1 day when far-red and/or red pulse was given atabout the 7th hour of the dark period. A far-jed pulse givenat the outset of the dark period only affected parameter P₂(flower induction period). Although the sensitivity of P₂ tored light increased with time, its sensitivity to far-red lightremained constant and at about the 7th hour was equally sensitiveto far-red and red lights. Both red and far-red pulses givenlater than the 7th hour were increasingly ineffective on P₂.The red/far-red reversibility occurred only for the action onP₂ of the far-red pulse applied during the early dark period.Parameter P₄ (flower production period) varied rhythmicallyin length with a far-red puke, the maximum shortening and extensionbeing induced by the pulse given at about the 7th and 19th hours,respectively. The sensitivity of P₄ to red light also changedrhythmically with an inverse phase angle to the rhythmic responseto farred light, and the far-red and red light actions werereversed respectively by subsequent red and far-red lights. These findings suggested that multiple timing devices includingan hourglass-type clock and a circadian clock are involved induckweed flowering. (Received October 25, 1978; )     

Reversal of sucrose inhibition of Lemna flowering by adenine derivatives

Sucrose inhibition of flowering in Lemna perpusilla 6746 wasat least partially reversed by 5'-AMP, cyclic 3',5'-AMP, 5'-ADP,5'-ATP and K₂HPO₄. These results are in contrast to those reportedfor L. gibba in which reversal was effected by cyclic AMP, butnot by other adenine derivatives. ¹ This work was supported by National Science Foundation GrantGB-12955. (Received June 11, 1973; )     

Red and blue light-stimulated proton efflux by epidermal leaf cells of the Argenteum mutant of Pisum sativum

Light stimulates leaf expansion in dicotyledons by increasingapoplastic acidification, cell wall loosening and solute accumulationfor turgor maintenance. Red and blue light enhance growth viadifferent photo-systems, but the cellular location and modesof action of these systems is not known. Here, the effect of red and blue light was studied on transportprocesses in epidermal cells of expanding leaves of the Argenteummutant of Pisum satlvum. Both red and blue light caused extraceiiuiaracidification by isolated epidermal tissue, which was stimulatedby extracellular K⁺ and inhibited by DCCD at 0.1 mol m^–3.Acidification induced by red compared with blue light showeddifferent saturating kinetics in fluence rate-response curves.Under near saturating light conditions the effects of red andblue light were additive. The red light-induced acidificationwas inhibited by far-red light while the blue light-inducedacidification was not. Light caused a hyperpoianzation of themembrane potential in epidermal strips, and stimulated ⁸⁶Rb⁺uptake by epidermal protoplasts. These results show that phytochromeand an additional blue light-photoreceptor function in isolatedepidermal cells to promote proton efflux, hyperpolarization,and cation uptake. Key words: Pisum sativum, light-induced acidification, ion transport, epidermis, photoreceptor     

THE RELATION BETWEEN RED AND BLUE OR FAR-RED LIGHTS IN THE NIGHT-INTERRUPTION OF THE PHOTOPERIODIC TUBERIZATION IN BEGONIA EVANSIANA

Effects of night-interruption on the aerial tuber formationof Begonia evansiana Andr. were investigated. Among coloredlights tested, red light was most effective to reduce the photoperiodicresponse. It inhibited tuberization almost completely at lowintensities. The red lightaction was partially reversed by subsequentblue or far-red irradiation under the 12-hour daylength, andthe relation between the red and the blue or far-red lightswas reversible. No reversal, however, was observed under the8-hour daylength. The inhibitory action of red light remainedunchanged on irradiating with red, blue or far-red light beforethe night periods. ¹Present address: Department of Horticulture, Purdue University,Lafayette, Indiana, USA     

Studies on the Photoreceptors for the Promotion and Inhibition of Flowering in Dark-Grown Seedlings of Pharbitis nil Choisy

Three-day-old etiolated seedlings of Pharbitis nil were exposedto red light for 10 min and sprayed with N⁶-benzyladenine beforetransfer to a 48-h inductive dark period, after which they weregrown under continuous white light. A second red irradiationpromoted flowering when given at the 5 and 24th hour of theinductive dark period but inhibited flowering at the 10 and15th hour. Far-red light inhibited flowering when given at anytime during the first 24 h of the dark period. Red/far-red reversibilitywas clearly observed at the 0, 5, 10 and 24th hour, but notat the 15th hour when both red and far-red lights completelyinhibited flowering. The action spectrum for the inhibition of flowering at the 15thhour of the inductive dark period had a sharply defined peakat 660 nm and closely resembled the absorption spectrum of theP_R form of phytochrome. The photoreceptors involved in thesephotoreactions are discussed. (Received June 10, 1983; Accepted July 6, 1983)     

Flowering in Pisum: the Effect of Light Quality on the Genotype If e Sn Hr

Far-red light, when given as a 16 h photoperiod extension, iamore effective than red light in reducing the flowering nodeof genotype Pisum. In contrast, when a 16 h dark period is interruptedby a 2 h light break red light is more effective than far-redlight. In addition, the stimulatory effect of a red interruptionis partially reversed by a subsequent period of far-red. However,a light interruption is not effective until over 12 h have elapsedsince the start of the previous photoperiod, regardless of whetherthe photoperiod was of 4 or 8 h duration. The results suggest that there are two light-dependent reactionscontrolling flowering in peas, one operating through the phytochromesystem with high levels of Pfr suppressing production of flowerinhibitor by the sn gene and a second requiring continuous illuminationwith wavelengths above 700 nm. The role of time measurementin the photoperiod response in peas is suggested to be filledby the proportion of time the Sn gene is effectively producinginhibitor. The photoperiod response in peas is not independentof temperature or plant age since the activity of gene Sn isalso varied by these factors.     

Flowering of the long-day plant, Lemna gibba, under short-day schedules composed of red and far-red light

Flowering in Lemna gibba, a long-day duckweed, can be inducedunder a short-day condition when the photoperiodic regimes areR₇FR₃ (7 hr red followed by 3 hr far-red), R₅FR₅ and R₃FR₇.This indicates the necessity of a proper balance between redand far-red effects for flowering. The flowering induced bythese regimes is inhibited by a brief exposure to red givenat the start of darkness and this inhibition is reversed bysubsequent exposure to far-red. Thus, the red/far-red reversibleeffect is found only at the beginning of darkness for floweringof L. gibba. However, flowering of L. gibba is promoted by a red light breakgiven near the middle of a 14 hr dark period. The promotiveeffect is not reversed by subsequent exposure to far-red, i.e.,the effect of the red break converts from inhibition to promotionas when given later in the dark period, which suggests the involvementof a timing mechanism. (Received July 21, 1973; )     

TWO LIGHT REACTIONS IN THE PHOTOPERIODIC CONTROL OF FLOWERING OF LEMNA PERPUSILLA AND L. GIBBA

The effects of light quality on the photoperiodic control inthe flowering of a SD duckweed, Lemna perpusilla strain 6746,and a LD duckweed, L. gibba strain G3, were investigated withspecial reference to the interaction between R and B or FR lights. In the diurnal alternation of R or G light and dark periods,L. perpusilla responded as a SDP, but in that of B or FR lightit was almost daylength-indifferent. On the other hand, L. gibbaresponded as a LDP under B, R or FR light, although the criticallight length was altered by the light quality. In the diurnal alternation of R and B or FR light periods containingno dark period, L. perpusilla flowered with the shortening ofthe optimal and critical R light lengths, compared with theplant exposed to that of R light and dark period. The floweringresponse of L. gibba to the R light length showed double peaks,that is, the first peak at the R duration less than 9 hours,and the second at the R duration longer than 9 hours. The firstpeak corresponds to the optimal R light length in L. perpusilla. Under the CL with a mixture of R and B or FR lights, the floweringand frond production were influenced by the intensity ratioof two light given. In both plants, the optimal ratio of B toR or FR to R for the flowering was always greater than thatfor the frond production. It is suggested that the B or FR light interacts with the Rlight in the photoperiodic process in the plants and this interactionbetween the R and B or FR lights should be of importance forobtaining a better understanding of photoperiodism. (Received August 28, 1965; )     

Photomorphogenesis in Pteris vittata II. Recovery from blue-light-induced inhibition of spore germination

1. 1. In the fern Pteris vittata, low-energy blue-light-inducedinhibition of phytochrome-dependent spore germination and darkrecovery from this inhibition were repeatedly observed severaltimes at intervals of 3 days at 26. The same amount of incidentenergy of blue light was required for inhibition in each successivetreatment.
2. 2. The recovery from blue-light-induced inhibitionof germinationwas markedly accelerated by continuous illuminationwith redlight, and this red light effect was not affected bythe presenceof CMU.
3. 3. The recovery process was not influencedby a single exposureto redlight, but was definitely promotedby brief red irradiationsgiven intermittently, at least 2 times,at equal intervals duringthe first 8 hr after blue light treatment.The effect of intermittentlygiven red light was annulled wheneach red exposure was followedby brief far-red irradiation.These facts suggest that phytochromemay be involved in therestoration of the ability of sporesto germinate (in responseto red light) which had been lostby blue irradiation.

¹Present address: Botany Department, Faculty of Science, Universityof Tokyo, Hongo, Tokyo 113.     

EXPERIMENTAL CONTROL OF FLOWERING IN LEMNA I. GENERAL METHODS. PHOTOPERIODISM IN L. PEPUSILLA 6746

Hillman , William S. (Yale U., New Haven, Conn.) Experimental control of flowering in Lemna. I. General methods. Photoperiodism in L. perpusilla 6746. Amer. Jour. Bot. 46(6): 466–473. Illus. 1959.—Lemna perpusilla strain 6746 flowers as a typical short-day plant when grown aseptically in Hutner's medium (containing ethylenediaminetetraacetic acid, [EDTA]) at 26–28°C. A method is described for quantitatively assaying the degree of flowering in a culture. Maximal flowering takes place under photoperiods of 6–11 hr., and none under photoperiods exceeding 15 hr. The flower-promoting effects of long nights are inhibited by brief interruptions with red light, such interruptions being most effective in the middle of the dark period. A single long night will cause the subsequent production of flowering fronds, but vegetative growth in the culture is resumed after a time. Only frond primordia at a very early stage of development appear to be sensitive to induction. Quantitative flowering experiments lasting a week or less can easily be performed with this plant; it is ideally suited for studies of the effects of light, darkness, temperature, organic compounds and other factors under highly controlled conditions.     

The Effects of Temperature, Photoperiod and Light Integral on the Time to Flowering of Pansy cv. Universal Violet (ViolaxwittrockianaGams.)

The effects of temperature, photoperiod and light integral onthe time to first flowering of pansy (ViolaxwittrockianaGams)were investigated. Plants were grown at six temperatures (meansbetween 14.8 and 26.1 °C), combined with four photoperiods(8, 11, 14 and 17 h). The rate of progress to flowering increasedlinearly with temperature (up to an optimum of 21.7 °C)and with increase in photoperiod (r²=0.91, 19 d.f.), the latterindicating that pansies are quantitative long day plants (LDPs).In a second experiment, plants were sown on five dates betweenJuly and December 1992 and grown in glasshouse compartmentsunder natural day lengths at six temperatures (means between9.4 and 26.3 °C). The optimum temperature for time to floweringdecreased linearly (from 21.3 °C) with declining light integralfrom 3.4 MJ m^-2d^-1(total solar radiation). Data from both experimentswere used to construct a photo-thermal model of flowering inpansy. This assumed that the rate of progress to flowering increasedas an additive linear function of light integral, temperatureand photoperiod. Independent data from plants sown on threedates, and grown at five temperatures (means between 9.8 and23.6 °C) were used to validate this model which gave a goodfit to the data (r²=0.88, 15 d.f.). Possible confounding ofthe effects of photoperiod and light integral are discussed. Pansy;Violaxwittrockiana; flowering; photo-thermal model; temperature; photoperiod; light integral