Are two photoreceptors involved in the flowering of a long-day plant?

The effect of daylength extension with narrow spectral bands on the flowering of a long-day plant, Brassica campestris L. cv. Ceres, was investigated to obtain clues to the identity of the photoreceptor involved. Extension of a 9 h photoperiod with 5 h of light pulses at various wavelengths resulted in maximal flowering occurring after irradiation at 710 nm, less at 730 nm, and none at 550, 660 and 750 nm. Flowering at 710 and 730 nm was negated by simultaneous exposures at 550 nm, but not at 660 nm. A short preirradiation at 660 nm enabled a following irradiation at 750 nm to induce flowering. This latter induction was prevented by 550 nm irradiation.
Short flashes of light at 710 nm induced flowering that was negated by a following flash at 550 nm but not at 660 nm. The negation by 550 nm radiation was prevented by subsequent flashes at 710 nm, indicating photoreversibility. A flash at 660 nm enabled subsequent light flashes at 750 nm to initiate flowering that was reversed by a following 550 nm flash.
From the results showing the necessity of red and far-red lights, it is proposed that flowering in this long-day plant is due to two photoreceptors - one is phytochrome and the other an unknown pigment with far-red, green photoreversible properties. By using fluence response data, it is deduced that the unidentified photoreceptor has weak absorption bands in the far-red, but has a strong absorption band in the green. Flowering is induced when effects of red light absorbed by phytochrome interact with effects of far-red light absorbed by the unidentified photoreceptor.     

Effects of blue and red light on unrolling of rice leaves

Unrolling of the second leaf of 8-day-old rice (Oryza sativa L.) seedlings was promoted by weak blue light (B), but not by red light (R). The effect of B was counteracted by irradiation with R just before or after the B. The counteracting effect of R was reversed by subsequent irradiation with far-red light but not by B, even if B was applied for 10 h. The B was effective when the region 0.5–2 cm from the tip of the leaf was irradiated. These results indicate that in rice photoreceptors for blue light located in the region 0.5–2 cm from the tip of the leaf play a key role in leaf unrolling and that a B-absorbing pigment and phytochrome participate in leaf unrolling in a closely related manner.Abbreviations B blue light - R red light - FR far-red light - W white light - D dark This work was presented at the Annual Meeting of the Japanese Society of Plant Physiologists on April 4, 1978, in Hiroshima     

The influence of blue light on dark-germinating seeds of Nemophila insignis

Summary After inhibition of Nemophila insignis seeds by far-red (FR) light, a short exposure to blue (Bl) will not induce germination again but stimulation by red (R), with reversion by FR, can be observed. Germination is inhibited by long exposures to Bl (maxima at 455 and 475 nm). These radiations are absorbed either directly by phytochrome or through intermediary pigments such as flavoproteins.Abbreviations Bl blue - FR far-red - R red     

Light Actions in the Germination of Cocklebur Seeds: I. DIFFERENCES IN THE LIGHT RESPONSES OF THE UPPER AND LOWER SEEDS

Germination responses to light were studied in the upper andlower seeds of cocklebur (Xanthium pennsylvanicum Wallr.). Thelower seed was dark-germinating and negatively photoblastic;the upper one had a red-light (R) requirement and was positivelyphotoblastic. Germination of the lower seeds was inhibited bya prolonged single irradiation with R, blue (B) or far-red (FR)light applied during imbibition. The maximal inhibitory effectof a single irradiation occurred 9 h and 13 h after the startof soaking at 33 °C and 23 °C, respectively. However,the inhibitory effect of R differed from that of B and FR, byonly delaying germination. A single exposure to B or FR lightcould be replaced by intermittent B or FR irradiation, and theireffects were repeatedly reversible by the following R irradiation.If the upper seeds were not exposed to R during imbibition,they failed to germinate even at 33 °C which was optimalfor germination, and the promotive effect of R increased withdelay of its application time. The photoperceptive locus incocklebur seeds was the axial tissue for all B, R and FR. Lightreceived by the cotyledonary tissue had little effect. Germinationdimorphism in response to light is discussed with respect tothe phytochrome content and the ageing of axial tissues. Key words: Blue light, Dimorphism, Far red light, Germination, Red light, Xanthium seed     

Kinetics of the Unrolling of the Second Leaves Detached from Rice Seedlings by Blue Light

Unrolling due to blue light (B) irradiation of the second leavesdetached from 8-day-old rice (Oryza saliva L.) seedlings wassimilar to that reported previously for nondetached leaves.The effect of B was counteracted by irradiation with red light(R). The counteracting effect of R was reversed by subsequentirradiation with far-red light (FR). When the detached leaf was irradiated with B passed througha 1-mm-wide slit 5, 8, 10, 12 or 15 mm from the leaf tip, irradiation10 mm from the leaf tip was the most effective. The effect of a 1 mm-wide-B irradiation 10 mm from the leaftip was counteracted by a 1 mm-wide-R irradiation at the sameposition, but not by irradiations at the other points. The counteractingeffect of R was reversed by a 1 mm-wide-FR irradiation at thesame position. This suggests that the excitation or the reactionof the B photoreceptor(s) is affected directly by the PFR formof phytochrome. The dose-response curve for the unrolling caused by B showeda simple Bunsen-Roscoe relation without two peaks, which differsfrom that for the phototropism in Avena caused by B. (Received August 21, 1980; Accepted December 20, 1980)     

不同光质的LED对蝴蝶兰组织培养增殖及生根的影响

以蝴蝶兰(Phalaenopsis ssp.)栽培种绿熊(Green Bear)和大辣椒(Big Chilli)为培养材料, 研究不同光质的发光二极管(LED)组合对组织培养过程中增殖及生根的影响。结果表明, 红光更有利于蝴蝶兰单芽增殖、干重、鲜重以及株高的增加, 但不利于叶片叶绿素的积累; 蓝光有利于叶片叶绿素的积累, 并能提高根系活力; 远红光则对根长和根系活力的增加作用更显著。增殖扩繁阶段的最适LED为暖白, 2种蝴蝶兰芽增殖系数分别比白色荧光灯(对照)高出53.17%和46.37%。生根诱导阶段的最适LED组合为红:蓝:远红=3:6:1, 根长及根系活力均较对照显著增加。该研究结果为LED光源在蝴蝶兰组织培养中的应用奠定了基础。     

Action of Near UV and Blue Light on the Photocontrol of Phycobiliprotein Formation; A Complementary Chromatic Adaptation

Action of near UV to blue light on photocontrol of phycoerythrin(PE) and phycocyanin (PC) formation was investigated with non-photobleachedTolypothrix tenuis and Fremyella diplosiphon; this study wasdone to evaluate the proposition of Haury and Bogorad [(1977)Plant Physiol., 60: 835] that near UV to blue light is as effectiveas green and red light for photocontrol of PE and PC formationin blue-green algae and that lack of the blue effect in previousexperiments was due to destruction of blue-absorbing pigment(s)by the photobleaching treatment involved in the experimentalmethod. In our present work, light effect was measured in heterotrophiccultures incubated in darkness following brief exposure to differentwavelengths of light. Results indicated that (1) near UV to blue light was not effectivefor induction of PE formation either in T. tenuis or in F. diplosiphon,and (2) PC formation was induced by near UV light at 360 nmbut not by blue light at 460 nm. These features are identicalwith those previously reported for photobleached cells but notwith those reported by Haury and Bogorad for non-photobleachedcells. We conclude that photobleaching treatment does not haveany influence on the action of near UV to blue light. Actionat 390 and 460 nm observed by Haury and Bogorad probably resultedfrom light effects other than photocontrol, e.g., the actionof photosynthesis. (Received December 18, 1981; Accepted April 8, 1982)     

Photocontrol of germination of cucumis anguria l

Germination ofCucumis anguria was inhibited by white, blue (B), and far-red (FR) irradiation and promoted by darkness and red (R) irradiation. The effect of white light was greater when supplied after rather than before the dark period. Darkness was more effective in reversing the effect of FR than FR in reversing the effect of darkness. FR was also more effective than B. When darkness followed B pretreatments, final germination percentage was higher than with FR pretreatment. R fully reversed the inhibitory effect of FR.     

Elimination of the Far-Red-Induced Lag Phase of Chlorophyll a Accumulation by a Brief Light in Pharbitis nil Cotyledons

In dark-grown Pharbitis nil seedlings, far-red light (FR) irradiationof 48 h or less promotes Chl a accumulation in the first 2-hof a subsequent white light (WL) period, without a lag phaseof Chl a accumulation. However, continuous FR irradiation of72 h or more, causes the so-called FR-induced lag phase. A 5-minWL given 4 h before the onset of the continuous WL promotesChl a accumulation irrespective of the length of the precedingFR irradiation period, if a 4-h dark period is inserted betweenthe 5-min WL and continuous WL. This suggests that the effectof the brief WL is independent of and additive to the effectof the preceding FR irradiation, although the effect of theFR irradiation changes from promotive to inhibitory with anincrease in the irradiation period. Red light (R) is more active than blue light (B) in this brieflight effect. The R effect is reversed by subsequent exposureto FR when the period of the preceding FR irradiation is 24h, but not when it is 72 h. The relative effectiveness of Bto R increases after prolonged FR irradiation. (Received August 6, 1986; Accepted March 12, 1987)     

Action spectra for the inhibition of growth in radish hypocotyls

In etiolated seedlings of Raphanus sativus L. the inhibition of hypocotyl elongation by continuous light showed a major bimodal peak of action in the red and far-red, and two minor peaks in the blue regions of the spectrum. It is argued that, under conditions of prolonged irradiation, phytochrome is the pigment controlling the inhibition of hypocotyl elongation by red and far-red light, but that its mode of action in far-red is different from that in red. A distinct pigment is postulated for blue light.Abbreviations B blue - FR far red - G green - R red - HIR high irradiance reaction - P_r and P_fr red and far red absorbing forms of phytochrome - R red     

Phytochrome-Mediated Photoorientation of Chloroplasts in Protonemal Cells of the Fern Adiantum Can be Induced by Brief Irradiation with Red Light

Measuring the ratio of the number of photooriented chloroplaststo the total number of chloroplasts, we found that photoorientationof chloroplasts in protonemata of the fern Adiantum capillus-veneriscould be induced by brief irradiation with polarized red light.After irradiation with red light (R) of 3 or 10 min, orientationalmovement was detected as early as 10 min after the irradiation;it continued during the subsequent dark period for 30–60min, after which chloroplasts gradually dispersed again. WhenR-treated protonemata were irradiated briefly with a second10-min pulse of R, 60 min after the onset of the first irradiation,the orientational response of chloroplasts was again observed.Typical red/far-red photoreversibility was apparent in the response,indicating the involvement of phytochrome. By contrast, irradiationwith polarized blue light for 10 min was ineffective, whileirradiation with blue light (B) at the same fluence for a longerperiod of time clearly induced the photoorientation of chloroplasts.It is likely that longterm irradiation is necessary for theresponse mediated by a blue-light receptor. When protonemata were irradiated with far-red light (FR) immediatelyafter R or after a subsequent dark period of 10 min, the magnitudeof the orientational response was smaller and chloroplasts dispersedmore quickly than those exposed to R alone. When FR was appliedat 50 min, when the response to R had reached the maximum level,chloroplasts again dispersed rapidly to their dark positions.These results indicate that P_FR not only induces the photoorientationmovement of chloroplasts but also fixes the chloroplasts atthe sites to which they have moved as a result of photoorientation. (Received June 2, 1993; Accepted January 11, 1994)     

Spectral Requirements for Flower Initiation in Two Longday Plants, Rape (Brassica campestris cv. Ceres), and Spring Wheat (Triticum×aestivum)

Action spectra for the promotion of flowering by long periods of irradiation in the red and far-red regions of the spectrum have been determined by the use of interference filters. The percentage floral initiation was greatest at 710–720 um for both wheat and rape (about 1 k erg · cm^?2· s^?1). Lowering the temperature from 35 to 15°C did not shift the most effective wavelength, suggesting that the effect of high levels of radiation at 710 nm was not related to a balance between photoactivation of phytochrome and its destruction. In both wheat and rape, an increase in temperature promoted the flower-initiating action of broad-band far-red and 710 nm radiation more than the action of broadband red or 660 nm radiation. The flower-promoting effect of broad-band red and 600 nm radiation was particularly depressed by temperatures above 25°C. It is concluded that the promotion of flowering by long periods of high energy at 710 nm is caused by the activity of a high-energy pigment, rather than through a special effect of phytochrome.     

红光与远红光比值对切花菊‘神马’发育和外观品质的影响

为了探讨不同R/FR值对温室切花菊发育进程和品质的影响,以切花菊品种‘神马’(Chrysanthemum morifoliumcv.‘Jingba’)为试材,设计不同红光(R:660nm!10nm)与远红光(FR:730nm!10nm)比值(R/FR)为0.5、2.5、4.5、6.5的LED灯照射处理试验,以自然光为对照(CK),观测不同处理的菊花发育阶段和品质指标。结果表明:与CK相比,R/FR=2.5处理显著加快菊花的发育进程(P<0.05),R/FR=2.5处理下短日照处理到现蕾、现蕾到破蕾及破蕾到收获3个发育阶段分别比CK提前4d、8d和5d,R/FR=0.5处理发育速度最慢,3个阶段分别比CK晚4d、2d和2d;不同R/FR值处理下菊花株高、单株叶龄、单株叶面积、茎粗、花径和花梗长度均随温光效应的增加呈增加的趋势,并在R/FR=2.5时取得最大值;收获时切花菊达到A级和B级产品等级的比例分别以R/FR=2.5和4.5处理最高。本研究发现,R/FR为2.5能够显著促进菊花发育进程和提高菊花的外观品质和A级切花的比例。     

Phytochrome- and blue-light-absorbing pigment-mediated directional movement of chloroplasts in dark-adapted prothallial cells of fernAdiantum as analyzed by microbeam irradiation

When prothalli ofAdiantum capillus-veneris L. were kept for 2 d in the dark, chloroplasts gathered along the anticlinal walls (Kagawa and Wada, 1994, J Plant Res 107: 389–398). In these dark-adapted prothallial cells, irradiation with a microbeam (10 gm in diameter) of red (R) or blue light (B) for 60 s moved the chloroplasts towards the irradiated locus during a subsequent dark period. Chloroplasts located less than 20 gm from the center of the R microbeam (18 J·m^–2) moved towards the irradiated locus. The higher the fluence of the light, the greater the distance from which chloroplasts could be attracted. The B microbeam was less effective than the R microbeam. Chloroplasts started to move anytime up to 20 min after the R stimulus, but with the B microbeam the effect of the stimulus was usually apparent within 10 min after irradiation. The velocity of chloroplast migration was independent of light-fluence in both R and B and was about - 0.3 m·min^–1 between 15 min and 30 min after irradiation. Whole-cell irradiation with far-red light immediately after R- and B-microbeam irradiations demonstrated that these responses were mediated by phytochrome and a blue-light-absorbing pigment, respectively. Sequential treatment with R and B microbeams, whose fluence rates were less than the threshold values when applied separately, resulted in an additive effect and induced chloroplast movement, strongly suggesting that signals from phytochrome and the blue-light-absorbing pigment could interact at some point before the induction of chloroplast movement.Abbreviations B blue light - FR far-red light - IR infrared light - R red light     

Growth control by phytochrome of de-etiolated bean hypocotyls mediated by chlorophyll fluorescence

The elongation of hypocotyls excised from de-etiolated seedlings of beans (Phaseolus vulgaris L. cv. British Wax) is inhibited by light, blue and red irradiations being equally effective. Conditions which decrease chlorophyll fluorescence, such as CO₂-free air, abolish the inhibitory effect of blue irradiation and enhance the inhibition by red light. Conversely, conditions which increase chlorophyll fluorescence, such as a N₂ atmosphere or irradiation through a chlorophyll filter, abolish the inhibitory effect of red light and enhance the inhibition by blue irradiation. The inhibitory effect of blue light is reversible by red irradiation under increased fluorescence as well as by far red. We propose that the chlorophyll fluorescence excited by blue and red irradiations in λ_F > 660 nm and λ_F > 720 nm, respectively, is responsible for the inhibitory effect of blue light and the reduction of the inhibitory effect of non fluorescing red light. Both red and blue wavelengths seem, therefore, to control hypocotyl elongation through phytochrome.     

Effects of light quality on leaf morphogenesis of a heterophyllous amphibious plant, Rotala hippuris

Background and Aims

For heterophyllous amphibious plants that experience fluctuating water levels, it is critical to control leaf development precisely in response to environmental cues that can serve as a quantitative index of water depth. Light quality can serve as such a cue because the ratio of red light relative to far-red light (R/FR) increases and blue-light intensity decreases with increasing water depth. Growth experiments were conducted to examine how R/FR and blue-light intensity alter leaf morphology of a heterophyllous amphibious plant, Rotala hippuris.

Methods

Using combinations of far red (730 nm), red (660 nm) and blue (470 nm) light-emitting diodes (LEDs), growth experiments were used to quantitatively evaluate the effects of the R/FR ratio and blue-light intensity on leaf morphology.

Key Results

Under the natural light regime in an outside growth garden, R. hippuris produced distinct leaves under submerged and aerial conditions. R/FR and blue-light intensity were found to markedly affect heterophyllous leaf formation. Higher and lower R/FR caused leaf characters more typical of submerged and aerial leaves, respectively, in both aerial and submerged conditions, in accordance with natural distribution of leaf types and light under water. High blue light caused a shift of trait values toward those of typical aerial leaves, and the response was most prominent under conditions of R/FR that were expected near the water surface.

Conclusions

R/FR and blue-light intensity provides quantitative cues for R. hippuris to detect water depth and determine the developmental fates of leaves, especially near the water surface. The utilization of these quantitative cues is expected to be important in habitats where plants experience water-level fluctuation.     

Action Spectra for Light-induced Elongation in Fern Protonemata

The action spectrum for promotion of elongation of protonemata of Onoclea sensibilis has peaks at 400–420, 580–600 and 640–660 nm. The largest growth increments at saturating light doses are produced by yellow and far-red light. Elongation induced by yellow and far-red irradiation persists in old as well as young filaments, while red-light promotion is found only in young filaments. The growth promotion caused by yellow light is partially reversed by red light down to the level of growth produced by red irradiation alone. Elongation of rhizoids is under reversible red, far-red control, while yellow light is inactive. A model is proposed and discussed in which the light-sensitive elongation of filaments is accounted for by the presence of three distinct photoreceptors: phytochrome; a pigment absorbing yellow light. P₅₈₀; and a pigment absorbing blue light, P₄₂₀.     

Developmental and photobiological factors affecting photoperiodic induction in Arabidopsis thaliana Heynh. Landsberg erecta

We have tested whether the promotion of flowering by long days(LD) in Arabidopsis thaliana is a consequence of photoperiodicinduction. To achieve this, the flowering responses of Arabidopsisthaliana (L.) Heynh. Landsberg erecta (Ler) and the long-hypocotylmutants hy2, hy3 and hy4 were determined with respect to age,daylength and light quality. Ler was capable of distinguishingbetween short days (SD) and long days (LD) from about 4 d aftersowing at 20 C, the time at which cotyledons were expandingand greening. At this stage, the critical daylength was between8 h and 10 h. At 7 d, seedlings required five LD for inductionand, as the seedlings aged, they became more sensitive so thatby day 20, one LD was fully inductive. The response to SD innewly germinated seedlings was to delay flowering without alteringleaf number, but after about 10 d, delay of flowering by SDwas accompanied by extra leaves. In light quality experiments,blue light (B) was inductive for 5-d-old plants and in all subsequenttreatments, far-red (FR) caused induction in treatments at 12d and 18 d and low pressure sodium, equivalent to red, was notinductive at 5 d and 12 d, but partially inductive at day 18.Hence, both a specific blue-light photoreceptor and phytochromeA in High Irradiance Response mode promote floral induction.In daylength transfer experiments all three hy mutants respondedto LD by earlier flowering. Both hy2 and hy3 produced substantiallyfewer leaves than Ler in SD and hy3 flowered slightly earlierthan Ler. The hy4 mutants flowered later than Ler in SD andhad a higher leaf number. A scheme is proposed in which photoperiodicinduction depends on the ability of the plant to sense photoperiod,the stage of development and the photobiological input. We alsopropose that phytochrome A and the blue photoreceptor promoteflowering whereas phytochrome B promotes vegetative development. Key words: Arabidopsis thaliana, blue-absorbing photoreceptor, flowering, photoperiodic induction, phytochrome     

Expansion of Vaucheria Cell Apex Caused by Blue or Red Light

Vaucheria geminata shows typical tip-growth. Symmetrical irradiationof the cell apex with strong blue or red light caused expansionat the very apex. The apical expansion required not only a largedosage, but also a long duration (>5 min), of irradiation.The expansion took place ca. 6 min after the onset of lighttreatment. The action spectrum for the response revealed thatblue light was the most effective: red light was two ordersof magnitude less effective. DCMU completely cancelled the expansioneffect of red light, yet the drug totally failed to inhibitthe effect of blue light. The expansion is thus controlled byboth the blue light-absorbing pigment system and photosynthesis.The expansion is a three-dimensional growth promotion event,i.e., it is accompanied by simultaneous longitudinal growth.Blue light caused another longitudinal growth promotion, a positivelight-growth response. The light-growth response is the firstfound in such a tip-growing cell. (Received January 17, 1981; Accepted February 27, 1981)     

The impact of red and blue light-emitting diode illumination on radish physiological indices

The objective was to evaluate the effect of different combinations of red (638 nm) and blue (455 nm) light produced by solid-state light-emitting diodes (LEDs) on physiological indices (net assimilation rate, hypocotyl-to-leaf ratio, leaf area, leaf dry weight, hypocotyl length and diameter, plant length, developed leaves), variation of photosynthetic pigments and non-structural carbohydrates in radish (Raphanus sativus L., var. ‘Faraon’). Lighting experiments were performed under controlled conditions (total PPFD - 200 μmol m⁻² s⁻¹; 16 h photoperiod; 14/18°C night/day temperature). The LED conditions: 638 nm; 638 + 5% 455 nm; 638 + 10% 455 nm; 638 + 10% 455 + 731 nm; 638 + 10% 455 + 731 + 669 nm. Our results showed that radishes grown under red (638 nm) alone were elongated, and the formation of hypocotyl was weak. The net assimilation rate, hypocotyl-to-leaf ratio, and leaf dry weight also were low due to the low accumulation of photosynthetic pigments and non-structural carbohydrates in leaves. The supplemented blue (455 nm) light was necessary for the non-structural carbohydrates distribution between radish storage organs and leaves which resulted in hypocotyl thickening. Red alone (638 nm) or in combination with far-red (731 nm), or red669 for radish generative development was required.