The loci of perception for phytochrome control of internode growth in light-grown mustard: Promotion by low phytochrome photoequilibria in the internode is enhanced by blue light perceived by the leaves

Under continuous white light (WL), extension growth of the first internode in Sinapis alba L. was promoted by low red (R): far-red (FR) ratios reaching the stem and-or the leaves. Conversely, the growth promotion by end-of-day light treatments was only triggered by FR perceived by the leaves and cotyledons, while FR given to the growning internode alone was tatally ineffective. Continuous WL+FR given to the internode was also in-effective if the rest of the shoot remained in darkness. Both the background stem growth, and the growth promotion caused by either an end-of-day FR pulse or continuous WL+FR given to the internode, increased with increasing fluence rates of WL given to the rest of the shoot. The increase by WL of the growth-stimulatory effect of low phytochrome photoequilibria in the internode appears to be mediated by a specific blue-light-absorbing photoreceptor, as blue-deficient light from sodium-discharge lamps, or from filtered fluorescent tubes, promoted background stem growth similarly to WL but did not amplify the response to the R:FR ratio in the internode. Supplementing the blue-deficient light (94 mol·m^-2·s^-1) with low fluence rates of blue (<9 mol·m^-2·s^-1) restored the promotive effect of low R:FR reaching the internode.Abbreviations BL blue light - FR far-red light - PAR photosynthetically active radiation - Pfr/P ratio between the FR-absorbing form and total phytochrome - R red light - SOX low-pressure sodium lamp - WL white light Supported by the Consejo Nacional de Investigaciones Cientificas y Técnicas (República Argentina) and the ORS scheme (UK)     

Floral initiation in Lolium temulentum L.: the role of phytochrome in the responses to red and far-red light

Summary The possibility that phytochrome is involved in the promotion of flowering by far-red light was investigated. The addition of far-red (FR) to a day extension with red (R) light promotes inflorescence initiation in Lolium. A 2-hour interruption with darkness also promoted flowering compared with the uninterrupted red light control; apex length was further increased by a 10-minute FR irradiation given before the 2-hour dark interruption and was decreased by 10-minutes of R light given in the middle: both FR promotion and R inhibition were reversed by R and FR respectively. Apex length increased approximately linearly with increasing duration of dark interruption up to at least 2 1/2 hours. When varying ratios of R:FR light were substituted for a 2-hour dark period, apex length was increasingly depressed as the % R was increased above 25%; no difference between 25% R/75% FR and 100% FR could be detected. Apex length was inversely linearly related to the calculated [Pfr]/[P] ratios above about 40% Pfr.FR promoted flowering when given during a 5-hour interruption of a day extension with R light but, between 0.25 and 0.90 J m² s^-1, there was no effect of intensity of FR; at 0.11 J m^-2 s^-1 apex length was shorter than at 0.25 J m^-2 s^-1 but longer than in darkness. When the duration of FR (from the beginning of a dark interruption of a day extension with R) was varied, apex length increased with increasing duration of FR up to 1 1/4 to 2 hours but further increasing the duration of FR did not promote flowering more.The results implicate phytochrome in the promotion of flowering by FR light. It has been demonstrated that a low [Pfr]/[P] ratio (less than present in 25% R/75% FR) is needed over a relatively long period of time: this explains why a relatively high proportion of FR light must be added to R for several hours in order to give maximum promotion of flowering. It is concluded that, in Lolium, the increased flowering response to FR light is brought about by a reduction of [Pfr]/[P] ratio at the appropriate time, although the possibility that another effect of far-red is also involved has not been rigorously excluded.     

Phytochrome and internode elongation in Chenopodium polyspermum L. sites of photoreception

The elongation of the fourth internode of fully green Chenopodium polyspermum L. is modulated by far-red light (FR) given in addition to the main light period. Two different types of organs are responsible for the photoreception of FR producing the end-of-day effect; the stem and the leaves situated just above and below the reacting internode. Photoreversibility can be obtained within certain limits in the two organs. Evidence is presented which shows that in the fully green plant there is an interorgan reaction whose primary reaction is the photoconversion of phytochrome.Abbreviations and Symbols D darkness - FR far red light - R red light - P phytochrome - P_FR phytochrome in the FR absorbing form - 9+15 D (or light treatment) photoperiod of a 9 h main light period followed by 15 h of D (or light treatment)     

Light inhibition of internode elongation in green plants

The clongation of the first internode of fully greenVigna sinensis L. is inhibited by white light (W). This inhibition is fluence-rate dependent between 0 and 70 Wm^–2. The kinetics of elongation rate in the light after darkness were investigated with linear displacement transducers. The internode elongation rate does not exhibit any endogenous rhythm. A rapid inhibition occurs during the first 2 or 3 h after the onset of light, and a second type of inhibition (slow reaction) increases from the beginning to the 8th hour of light. The rapid inhibition is not fluence-rate dependent between 20 and 70 Wm^–2, but the slow reaction is. There is no rapid inhibition in a low fluence rate white light to high fluence rate white light transition, only the slow reaction is observed. The responses to different wavebands, i.e., blue light (B), yellow and green light (YG), and red light (R), are the same for the two inhibition reactions. Each waveband used separately does not reproduce the full effect observed in W. Results show a stimulation with B, a greater inhibition activity with YG than with R, and a synergistic action of B and R which when given together lead to an inhibition similar to that obtained in W. Plants returned from the light to darkness progressively recover a high elongation rate without any latent period. The W light regulating internode elongation rate is mainly perceived by the growing internode itself.Abbreviations B blue light - D darkness - F far-red light - HW high fluence rate white light - LW low fluence rate white light - R red light - W white light - YG yellow and green light     

Photomorphogenesis in Chenopodium album. Effects of supplementary far-red light on the kinetics of stem extension

The effects of far-red light given against a background of white light on the stem-extension kinetics of three-week-old, light-grown Chenopodium album seedlings were investigated. Under white light alone, the stems (cotyledon-to-apex) extended almost exactly logarithmically with time. Under these conditions the increase in log [stem length in mm] per hour was approx. 3.7·10^-3, equivalent to about 1% per h during both skoto-and photoperiods. Supplementary far-red given throughout each photoperiod massively stimulated extension. The calculated logarithmic extension rate, however, slowly returned to that of the controls, following an initial large increase. This is predicted by a model in which far-red light linearly increases the extension rate of individual internodes which arise at an exponentially increasing rate. The behaviour of the model is also consistent with critical experiments in which far-red was given as a pre-treatment or transiently, as well as with other published data. Far-red stimulation of logarithmic extension rate in successive photoperiods was closely and linearly correlated with calculated phytochrome photoequilibrium. Daily short periods of supplementary far-red were especially potent in accelerating extension; the plants seemed least responsive at the end of the photoperiod.Abbreviations FR supplementary far-red light - I stem length (mm) - LSER logarithmic stem extension rate - Pfr far-red absorbing form of phytochrome - R:FR red:far-red fluence rate ratio - WL white light - _c calculated phytochrome photoequilibrium     

Light-grown plants of transgenic tobacco expressing an introduced oat phytochrome A gene under the control of a constitutive viral promoter exhibit persistent growth inhibition by far-red light

A comparison of the photoregulation of development has been made for etiolated and light-grown plants of wild-type (WT) tobacco (Nicotiana tabacun L.) and an isogenic transgenic line which expresses an introduced oat phytochrome gene (phyA) under the control of a constitutive viral promoter. Etiolated seedlings of both the WT and transgenic line showed irradiance-dependent inhibition of hypocotyl growth under continuous far-red (FR) light; transgenic seedlings showed a greater level of inhibition under a given fluence rate and this is considered to be the result of the heterologous phytochrome protein (PhyA) functioning in a compatible manner with the native etiolated phytochrome. Deetiolation of WT seedlings resulted in a loss of responsiveness to prolonged FR. Light-grown transgenic seedlings, however, continued to respond in an irradiance-dependent manner to prolonged FR and it is proposed that this is a specific function of the constitutive PhyA. Mature green plants of the WT and transgenic lines showed a qualitatively similar growth promotion to a brief end-of-day FR-treatment but this response was abolished in the transgenic plants under prolonged irradiation by this same FR source. Growth inhibition (McCormac et al. 1991, Planta 185, 162–170) and enhanced levels of nitrate-reductase activity under irradiance of low red:far-red ratio, as achieved by the FR-supplementation of white light, emphasised that the introduced PhyA was eliciting an aberrant mode of photoresponse compared with the normal phytochrome population of light-grown plants. Total levels of the oat-encoded phytochrome in the etiolated transgenic tobacco were shown to be influenced by the wavelength of continuous irradiation in a manner which was qualitatively similar to that seen for the native, etiolated tobacco phytochrome, and distinct from that seen in etiolated oat tissues. These results are discussed in terms of the proposal that the constitutive oat-PhyA pool in the transgenic plants leads to a persistence of a mode of response normally restricted to the situation in etiolated plants.Abbreviations FR far-red light - R red light - WL white light - WL + FR white light supplemented with FR - HIR high-irradiance response - PAR photosynthetically active radiation - Pr, Pfr R- and FR-absorbing forms of phytochrome - Ptot total phytochrome - phyA (PhyA) gene (encoded protein) for phytochrome - WT wild type This work was supported by an Agricultural and Food Research Council research grant to H.S. and A.M.; J.R. Cherry and R.D. Vierstra, (Department of Horticulture, University of Wisconsin-Madison, USA) are thanked for the provision of the transgenic tobacco line.     

Light requirement,phytochrome and photoperiodic induction of flowering of Pharbitis nil Chois

For dark-grown seedlings of Pharbitis nil capacity to flower in response to a single inductive dark period was established by 24 h white, far-red (FR) or ruby-red (BCJ) light and by a skeleton photoperiod of 10 min red (R)-24 h dark-10 min R. FR alone was ineffective without a brief terminal (R) irradiation, confirming that the form of phytochrome immediately prior to darkness is a crucial factor for flowering in Pharbitis. The magnitude of the flowering response was significantly greater after 24 h FR or white light (WL) (at 18° C and 27° C) than after two brief skeleton R irradiations, but the increased flowering response was not attributable to photosynthetic CO₂ uptake because this could not be detected in seedlings exposed to 24 h WL at 18° C. Photophosphorylation could have contributed to the increased flowering response as photosystem I fluorescence was detectable in plants exposed to FR, BCJ, or WL, but there were large differences between flowering response and photosystem I capacity as indicated by fluorescence. We conclude that phytochrome plays a major role in photoresponses regulating flowering. There was no simple correlation between developmental changes, such as cotyledon expansion and chlorophyll formation during the 24-h irradiation period, and the capacity to flower in response to a following inductive dark period. Changes in plastid ultrastructure were considerable in light from fluorescent lamps and there was complete breakdown of the prolamellar body with or without lamellar stacking at 27 or 18° C, respectively, but plastid reorganization was minimal in FR-irradiated seedlings.Abbreviations BCJ irradiation from photographic ruby-red lamps - FR far-red light - P_fr far-red-absorbing from of phytochrome - P total phytochrome content - R red light - WL white light from fluorescent lamps     

Is the far-red-absorbing form of Avena phytochrome A that is present at the end of the day able to sustain stem-growth inhibition during the night in transgenic tobacco and tomato seedlings?

Avena phytochrome A (phyA) overexpressed in tobacco (Nicotiana tabacum L.) and tomato (Lycopersicon sculentum Mill) was functionally characterised by comparing wild-type (WT) and transgenic seedlings. Different proportions of phytochrome in its far-red-absorbing form (Pfr/P) were provided by end-of-day (EOD) light pulses. Stem-length responses occurred largely in the range of low Pfr/P (3–61%) for WT seedlings and in the range of high Pfr/P (61–87%) for transgenic seedlings. A similar shift was observed when the photoperiod was interrupted by short light pulses providing different Pfr/P ratios and followed by 1 h dark incubation. In other experiments, Avena phyA was allowed to re-accumulate in darkness and subsequently phototransformed to Pfr but no extra inhibition of stem extension growth was observed. In transgenic tomato seedlings the response to EOD far-red light was faster and the response to a far-red light pulse delayed into darkness was larger than in the WT. Avena phyA Pfr remaining at the end of the photoperiod appears intrinsically unable to sustain growth inhibition in subsequent darkness. Avena phyA modifies the sensitivity and the kinetics of EOD responses mediated by native phytochrome.Abbreviations EOD end-of-day - FR far-red light - Pfr/P pro-portion of phytochrome in its FR-absorbing form - phyA phyto-chrome A - phyB phytochrome B - R red light - RFR R to FR ratio - WT wild type We thank Dr Brian Thomas for providing the antibodies used in this work, and Federico Guerendiain for his excellent technical assistance. This work was financially supported by grants UBA AG 040 and Fundacion Antorchas A-12830/1-19 (both to J.J.C.), PID-CONICET (to R.A.S. and J.J.C.), United States Department of Energy DE-FG02-88ER13968 (to R.D.V.).     

Sensitivity to far-red radiation in stomata of Phaseolus vulgaris L.: Rhythmic effects on conductance and photosynthesis

The influence of far-red (FR; 700–800 nm) radiation on steady-state stomatal conductance and net photosynthesis in P. vulgaris has been studied. Whereas FR radiation alone was relatively ineffective, addition of FR to a background of white light (WL; predominantly 400–700 nm) resulted in increased stomatal conductance. Stomata exhibited a marked diurnal sensitivity to FR. The action maximum for enhancing stomatal conductance was near 714 nm. A combination of FR and infra-red (IR; >800 nm) enhanced net photosynthesis when added to a background of WL. When IR alone was added to WL, there was a net decrease in photosynthesis, indicating that it is the FR waveband which is responsible for the observed photosynthetic effects. Naturally occurring levels of FR radiation (235 mol·m^-2·s^-1) in vegetation-canopy shade enhanced net photosynthetic CO₂ gain by 28% when added to a background of 55 mol·m^-2·s^-1 WL.Abbreviations BL blue - FR far-red - IR infra-red - PAR photosynthetically active radiation - R red - WL white light     

Floral initiation in sorghum hastened by gibberellic acid and far-red light

Combinations of far-red light (FR) (4 min) and gibberellic acid (GA₃), given at the beginning of a daily 12-h dark period in a growth room, were used to study floral induction in four maturity genotypes of the milo group of sorghum (Sorghum bicolor (L.) Moench). The 12-h dark period without GA₃ application or FR induced flowering in only the early genotype; FR hastened initiation in the early genotype, while GA₃ hastened floral initiation in the two intermidiate-flowering genotypes. GA₃ and FR together had a strong synergistic effect, hastening floral initiation by 30 to more than 80 d in the early and intermediate genotypes. Red light (R) did not hasten flowering; FR preceded by R gave the same effect as FR alone. GA₃ promoted stem elongation equally whether floral initiation occurred or not; thus, its effect on stem elongation was independent of floral initiation. The capacity of GA₃ to induce flowering in sorghum, a short-day plant, seems to be enhanced by phytochrome being in the P_R form at the beginning of the night when GA₃ was applied.Abbreviations FR far-red light - GA(s) gibberellin(s) - GA₃ gibberellic acid - R red light     

Rapid photomodulation of stem extension in light-grownSinapis alba L.

Treatment of the whole of aSinapis alba plant with supplementary far-red light (FR), in back-ground white light (WL), induces a rapid increase in stem extension rate. This rapid increase is regulated by the light environment of the stem itself. Supplementary FR to the stem increases extension rate after a lag period of 10–15 min. A lag period of 3–4 h follows FR irradiation of the leaf, before an increase in extension rate is detectable. When the stem is given supplementary FR, the change in extension rate which is induced increases with increasing FR fluence rate, and with decreasing phytochrome photoequilibrium. There is no difference between the effects of supplementary FR _max 719 nm and supplementary FR _max 739 nm for these relationships. The increase in extension rate induced by supplementary FR is reversed by an increase in the fluence rate of red light (R). These data indicate that the response is controlled by phytochrome photoequilibrium.Abbreviations B blue light - FR far-red light - R red light - WL white light - Pfr far-red absorbing form of phytochrome - Pr red absorbing form of phytochrome - P_tot total phytochrome level (=Pr+Pfr); -Pfr/Ptot, measured - ER difference in stem extension rate, before and after treatment     

Light effects on in vitro rooting of pear cultivars of different rhizogenic ability

The effect of varying light regimes on in vitro rooting of microcuttings of two pear (Pyrus communis L.) cultivars was investigated. Cultures of the easy to-root Conference and the difficult-to-root Doyenne d'Hiver were incubated for 21 days with or without indole-3-butyric acid (IBA) in the medium in darkness or under continuous far-red (8 µmol m^–2 s^–1), blue, white or red (15 or 36 µmol m^–2 s^–1) light. Conference rooted without IBA when exposed to red, blue or white light while no rooting was observed under far-red light and in darkness. The high rooting efficiency under red and, by contrast, the inhibition under far-red light and darkness suggest the involvement of the phytochrome system in rhizogenesis. The addition of IBA to the culture medium enhanced root production under all light regimes in both cultivars. Red light, especially at the lower photon fluence rate, had a positive effect by increasing root extension (number × length of roots) and stimulating secondary root formation.Abbreviations IBA Indole-3-butyric acid - R red light - B blue light - FR far-red light - W white light - D darkness - P_fr active (far-red light absorbing) form of phytochrome - P_tot total phytochrome - BA benzyl-adenine     

Far-red light inhibits growth but promotes carotenoid accumulation in the green microalga Dunaliella bardawil

It has been demonstrated that far-red light reduces growth of marine phytoplankton and that light quality controls growth and photosynthetic metabolism in algae. The green halotolerant microalga, Dunaliella bardawil, accumulates high amounts of β-carotene (up to 10% of its dry weight) under conditions of high light or nutrient limitation. The influence of increasing irradiance and of far-red light in D. bardawil was studied. Continuous irradiance was provided by white fluorescent lamps alone (WL) or supplemented with far-red Linestra lamps (WL+FR). For both types of light, cultures were acclimatized at increasing irradiances (50-300 µmol m^?2 s^?1), and cell density, photosynthetic activity and pigment content were determined. Cell density increased with the photon irradiance, and was higher in WL than in WL+FR under the same irradiance, but the reverse occurred in respect of cell volume. Growth rate was higher under WL+FR. Far-red light induced faster growth but reduced the maximal cell density of the cultures. Chlorophyll a concentration was higher in white light, but total carotenoid content increased dramatically in both far-red light treatments (about 50% on a per cell basis) and with the increase of irradiance. Our results show that far-red light has a significant influence on growth and photosynthesis of D. bardawil, inducing a decrease in cell density, photosynthetic activity and chlorophyll concentration, and an increase in growth rate, cell volume and carotenoid content.     

Far-red light-insensitive, phytochrome A-deficient mutants of tomato

We have selected two recessive mutants of tomato with slightly longer hypocotyls than the wild type, one under low fluence rate (3 mol/m²/s) red light (R) and the other under low fluence rate blue light. These two mutants were shown to be allelic and further analysis revealed that hypocotyl growth was totally insensitive to far-red light (FR). We propose the gene symbol fri (far-red light insensitive) for this locus and have mapped it on chromosome 10. Immunochemically detectable phytochrome A polypeptide is essentially absent in the fri mutants as is the bulk spectrophotometrically detectable labile phytochrome pool in etiolated seedlings. A phytochrome B-like polypeptide is present in normal amounts and a small stable phytochrome pool can be readily detected by spectrophotometry in the fri mutants. Inhibition of hypocotyl growth by a R pulse given every 4 h is quantitatively similar in the fri mutants and wild type and the effect is to a large extent reversible if R pulses are followed immediately by a FR pulse. After 7 days in darkness, both fri mutants and the wild type become green on transfer to white light, but after 7 days in FR, the wild-type seedlings that have expanded their cotyledons lose their capacity to green in white light, while the fri mutants de-etiolate. Adult plants of the fri mutants show retarded growth and are prone to wilting, but exhibit a normal elongation response to FR given at the end of the daily photoperiod. The inhibition of seed germination by continuous FR exhibited by the wild type is normal in the fri mutants. It is proposed that these fri mutants are putative phytochrome A mutants which have normal pools of other phytochromes.     

Fluence rate and wavelength dependence of photobleaching in the cyanobacterium Anabaena variabilis

The fluence rate dependence of the photobleaching in the cyanobacterium Anabaena variabilis was studied under physiological conditions. According to the in-vivo absorption spectra measured every day during the 5 d exposition the phycobiliproteins are more sensitive to high fluence rates than chlorophyll a. The carotenoids are least sensitive, so that a relative, but not an absolute increase in the carotenoid content occurred. At very high fluence rates exceeding about 50 Wm^-2 white light the organisms were photokilled after 5 d of irradiation. Measurements of the nitrate concentrations during the experiments have shown that nitrate was not the limiting factor in these experiments. Analysis of the photobleaching kinetics at 13.5 Wm^-2 white light revealed that after about 8 d the contents of all the pigments studied have reached a new, constant level. After exposure of the photobleached cyanobacteria to low irradiances repigmentation occurred. Thus, photobleaching is a light adaptation process and not simply a photodamage phenomenon. Studying the wavelength dependence of photobleaching at a constant photon fluence rate of 4·10^-8 mol cm^-2 s^-1 we found that the photobleaching of both phycobiliproteins and chlorophyll a was exclusively caused by wavelengths absorbed by the phycobiliproteins, mainly phycoerythrocaynin, and red light absorbed by short wavelength chlorophyll. Wavelengths <520 nm were ineffective.     

Phototaxis in the flagellates,Euglena gracilis and Ochromonas danica

Oriented movement with respect to laterally impinging white light of the flagellates Euglena gracilis and Ochromonas danica has been analyzed in an individual cell study with a microvideographic technique. Using the deviation of track segments (in given time intervals of 1 s) from the light direction as raw data allowed a computer based analysis of the direction distribution. A number of statistical methods employed to test the significance of the obtained results demonstrated an obvious phototactic orientation in Ochromonas which was positive (toward the light source) in low illuminance (1.25 lx=5.3×10^-3 Wm^-2) and negative in higher illuminance (>12.5 lx=5.3×10^-2 Wm^-2). Since in this flagellate the threshold for negative phototaxis is much lower than that for the step-up photophobic response, the hypothesis that negative phototaxis may be brought about by repetitive step-up phobic responses can be rejected for at least this organism. In Euglena positive phototaxis was observed in 50 lx (=0.21 Wm^-2), while an illuminance of 500 lx (=2.1 Wm^-2) caused a negative phototaxis.The experiments were carried out in this laboratory     

Phytochrome-mediated effects of near-ultraviolet radiation in the induction of flowering in etiolated Lemna paucicostata T-101, a short-day plant

Induction of flowering of etiolated Lemna paucicostata Hegelm. T-101, a short-day plant, was inhibited by far-red (FR) or blue light (BL) applied at the beginning of a 72-h inductive dark period which was followed by two short days. In either case the inhibition was reversed by a subsequent exposure of the plants to near-ultraviolet radiation (NUV), with a peak of effectiveness near 380 nm. Inhibition by BL or FR and its reversion by NUV are repeatable, i.e., NUV is acting in these photoresponses like red light although with much lower effectiveness. Thus, it is considered that NUV acts through phytochrome and no specific BL and NUV photoreceptor is involved in photocontrol of floral induction on this plant.Abbreviations BL blue light - FR far-red light - NUV near ultraviolet radiation - P red-absorbing form of phytochrome - P_fr far-red absorbing form of phytochrome - R red light     

Effect of washing on the plasma membrane and on stress reactions of cultured rose cells

The effects of plant growth regulators, light intensity, and end-of-day (EOD) light quality treatments on node and microtuber induction (% of cultures with microtubers) and development (fresh weight of microtubers) in yam (Dioscorea alata L. cv. Oriental) cultures were investigated. Nodal segments were excised from plantlets cultured on tuberization medium containing growth regulators and exposed to various light treatments. Absciscic acid (1 M) stimulated and cytokinins (2.5 M) inhibited microtuber development from yam nodal segments cultured on Mantell's and Hugo's full-strength tuberization medium under 8-h photoperiods. EOD far-red (FR) light inhibited microtuber induction and development and enhanced node formation. EOD FR light effects were nullified by immediately following the FR treatment with red light. This suggested the involvement of phytochrome in these processes. The lowest light intensity evaluated (12 mol m^–2 s^–1) inhibited microtuber, root and shoot production as compared to light intensities of 42, 72 and 102 mol m^–2 s^–1. Kinetin (2.5 m) in half-strength tuberization medium inhibited microtuber induction and development but did not affect node production in the light intensity evaluation.Abbreviations ABA abscisic acid - BA 6-benzylaminopurine - 2iP 6-(c,c-dimethylallylamino)-purine - NAA napthaleneacetic acid - R light red light - FR light far-red light - EOD light end-of-day light     

Gibberellins and phytochrome regulation of stem elongation in pea

In garden pea (Pisum sativum L.) neither etiolation nor the phytochrome B (phyB)-response mutation lv substantially alters the level of the major active endogenous gibberellin, GA₁ in the apical portion of young seedlings. The phyB-controlled responses to continuous red light and end-of-day far-red light are retained even in a GA-overproducing mutant (sln). Comparison of the effects of the lv mutation and GA₁ application on seedling development shows important differences in rate of node development, cell extension and division, and leaf development. These results suggest that in pea the control of stem elongation by light in general and phyB in particular is not mediated by changes in GA₁ content. Instead, the increased elongation of dark-grown and lv plants appears to result from increased responsiveness of the plant to its endogenous levels of GA₁. Three GA₁-deficient mutants, na, ls and le have been used to investigate these changes in responsiveness, and study of these and the double mutants na lv, ls lv and le lv has demonstrated that the relative magnitude of the change in responsiveness is dependent on GA₁ level. The difference in pleiotropic effects of GA₁ application and the lv mutation suggest that light and GA₁ interact late in their respective transduction pathways. A model for the relationship between light, GA₁ level and elongation in pea is presented and discussed.Abbreviations B blue light - cv cultivar - EOD-FR end-of-day far-red light - FR far-red light - GA_n Gibberellin A_n - GC-SIM gas chromatography-selected ion monitoring - HIR high irradiance response - W white light We thank Prof. L.N. Mander for provision of deuterated internal standards, Peter Bobbi, Noel Davies, Omar Hasan, and Katherine McPherson for technical assistance, Stephen Swain for discussion and provision of GA-level data, and the Australian Research Council for financial assistance. J.L.W. is in receipt of an Australian Postgraduate Research scholarship.     

The role of calcium ions in phytochrome-controlled swelling of etiolated wheat (Triticum aestivum L.) protoplasts

Protoplasts from dark-grown wheat (Triticum aestivum L.) maintained at a constant osmotic potential at 22°C, were found to swell upon red irradiation (R) and the effect was negated by subsequent far-red light (FR), indicating phytochrome involvement. Swelling only occurred when Ca²⁺ ions were present in the surrounding medium, or were added within 10 min after R. Furthermore, Mg²⁺, Ba²⁺ or K⁺ could not replace this requirement for Ca²⁺. The presence of K⁺ did not enhance the Ca²⁺-dependent swelling response. When the Ca²⁺-ionophore A 23187 was added to the medium, protoplasts swelled in the dark to the same extent as after R. Both the Ca²⁺-channelblocker Verapamil and La³⁺ inhibited R-induced swelling. It is proposed that R causes the opening of Ca²⁺-channels in the plasma membrane. Boyle-van't Hoff analyses of protoplast volume after R and FR are consistent with the conclusion that R irradiation causes changes in membrane properties.Abbreviations EDTA ethylenediaminetetraacetic acid - FR far-red light - nov non-osmotic-volume - Pfr FR-absorbing form of phytochrome - Pr R-absorbing form of phytochrome - R red light