Red:far-red light conditions affect the emission of volatile organic compounds from barley (Hordeum vulgare), leading to altered biomass allocation in neighbouring plants

Background and Aims Volatile organic compounds (VOCs) play various roles in plant–plant interactions, and constitutively produced VOCs might act as a cue to sense neighbouring plants. Previous studies have shown that VOCs emitted from the barley (Hordeum vulgare) cultivar ‘Alva’ cause changes in biomass allocation in plants of the cultivar ‘Kara’. Other studies have shown that shading and the low red:far-red (R:FR) conditions that prevail at high plant densities can reduce the quantity and alter the composition of the VOCs emitted by Arabidopsis thaliana, but whether this affects plant–plant signalling remains unknown. This study therefore examines the effects of far-red light enrichment on VOC emissions and plant–plant signalling between ‘Alva’ and ‘Kara’.Methods The proximity of neighbouring plants was mimicked by supplemental far-red light treatment of VOC emitter plants of barley grown in growth chambers. Volatiles emitted by ‘Alva’ under control and far-red light-enriched conditions were analysed using gas chromatography–mass spectrometry (GC-MS). ‘Kara’ plants were exposed to the VOC blend emitted by the ‘Alva’ plants that were subjected to either of the light treatments. Dry matter partitioning, leaf area, stem and total root length were determined for ‘Kara’ plants exposed to ‘Alva’ VOCs, and also for ‘Alva’ plants exposed to either control or far-red-enriched light treatments.Key Results Total VOC emissions by ‘Alva’ were reduced under low R:FR conditions compared with control light conditions, although individual volatile compounds were found to be either suppressed, induced or not affected by R:FR. The altered composition of the VOC blend emitted by ‘Alva’ plants exposed to low R:FR was found to affect carbon allocation in receiver plants of ‘Kara’.Conclusions The results indicate that changes in R:FR light conditions influence the emissions of VOCs in barley, and that these altered emissions affect VOC-mediated plant–plant interactions.     

Green light: a signal to slow down or stop

Light has a profound effect on plant growth and development. Red and blue light best drive photosynthetic metabolism, so it is no surprise that these light qualities are particularly efficient in advancing the developmental characteristics associated with autotrophic growth habits. Photosynthetically inefficient light qualities also impart important environmental information to a developing plant. For example, far-red light reverses the effect of phytochromes, leading to changes in gene expression, plant architecture, and reproductive responses. Recent evidence shows that green light also has discrete effects on plant biology, and the mechanisms that sense this light quality are now being elucidated. Green light has been shown to affect plant processes via cryptochrome-dependent and cryptochrome-independent means. Generally, the effects of green light oppose those directed by red and blue wavebands. This review examines the literature where green light has been implicated in physiological or developmental outcomes, many not easily attributable to known sensory systems. Here roles of green light in the regulation of vegetative development, photoperiodic flowering, stomatal opening, stem growth modulation, chloroplast gene expression and plant stature are discussed, drawing from data gathered over the last 50 years of plant photobiological research. Together these reports support a conclusion that green light sensory systems adjust development and growth in orchestration with red and blue sensors.     

Genome-wide analysis of light-dependent transcript accumulation patterns during early stages of Arabidopsis seedling deetiolation

Light is among the most important exogenous factors that regulate plant development. To sense light quality, intensity, direction, and duration, plants have evolved multiple photoreceptors that enable the detection of photons from the ultraviolet B (UV-B) to the far-red spectrum. To study the effect of different light qualities on early gene expression, dark-grown Arabidopsis (Arabidopsis thaliana) seedlings were either irradiated with continuous far-red, red, or blue light or received pulses of red, UV-A, or UV-A/B light. The expression profiles of seedlings harvested at 45 min and 4 h were determined on a full genome level and compared with the profiles of dark controls. Data were used to identify light-regulated genes and to group these genes according to their light responses. While most of the genes were regulated by more than one light quality, a considerable number of UV-B-specific gene expression responses were obtained. An extraordinarily high similarity in gene expression patterns was obtained for samples that perceived continuous irradiation with either far-red or blue light for 4 h. Mutant analyses hint that this coincidence is caused by a convergence of the signaling cascades that regulate gene expression downstream of cryptochrome blue light photoreceptors and phytochrome A. Whereas many early light-regulated genes exhibited uniform responses to all applied light treatments, highly divergent expression patterns developed at 4 h. These data clearly indicate that light signaling during early deetiolation undergoes a switch from a rapid, but unspecific, response mode to regulatory systems that measure the spectral composition and duration of incident light.     

Red-light and blue-light photoreceptors controlling chlorophyll a synthesis in the red alga Porphyra umbilicalis and in the green alga Ulva rigida

Chlorophyll synthesis is stimulated by red light in the green alga Ulva rigida C. Ag. and in the red alga Porphyra umbilicalis (L.) Kützing. Because the effect of red light showed some far-red reversibility in successive red and far-red light treatments, the involvement of phytochrome or a phytochrome-like photoreceptor is suggested. The extent of the response is dependent on exposure and photon fluence rate of red-light pulses. In addition to the effect of red light, a strong stimulation of chlorophyll synthesis by blue light was only observed in Ulva rigida. The effect of blue light shows also some far-red reversibility. In the green alga the accumulated chlorophyll is higher after blue light pulses than after red light pulses. In Porphyra umbilicalis , however, the contrary is observed. In Ulva rigida the involvement of a blue light photoreceptor in addition to phytochrome or a phytochrome-like photoreceptor is proposed. The different responses to red and blue light in both algae are explained in terms of their adaptation to the natural light environment.     

Diatom Phytochromes Reveal the Existence of Far-Red-Light-Based Sensing in the Ocean

The absorption of visible light in aquatic environments has led to the common assumption that aquatic organisms sense and adapt to penetrative blue/green light wavelengths but show little or no response to the more attenuated red/far-red wavelengths. Here, we show that two marine diatom species, Phaeodactylum tricornutum and Thalassiosira pseudonana, possess a bona fide red/far-red light sensing phytochrome (DPH) that uses biliverdin as a chromophore and displays accentuated red-shifted absorbance peaks compared with other characterized plant and algal phytochromes. Exposure to both red and far-red light causes changes in gene expression in P. tricornutum, and the responses to far-red light disappear in DPH knockout cells, demonstrating that P. tricornutum DPH mediates far-red light signaling. The identification of DPH genes in diverse diatom species widely distributed along the water column further emphasizes the ecological significance of far-red light sensing, raising questions about the sources of far-red light. Our analyses indicate that, although far-red wavelengths from sunlight are only detectable at the ocean surface, chlorophyll fluorescence and Raman scattering can generate red/far-red photons in deeper layers. This study opens up novel perspectives on phytochrome-mediated far-red light signaling in the ocean and on the light sensing and adaptive capabilities of marine phototrophs.     

Light-stimulated respiration in the green algaDunaliella tertiolecta: Involvement of the ultraviolet/blue-light photoreceptor(s) and phytochrome?

Starch breakdown and respiratory O₂ uptake in the green algaDunaliella tertiolecta (Butcher) are stimulated not only by blue, but also by red light. In the present study, attempts are described to identify the photoreceptor(s) involved. Fluence rate-response curves with different slopes in the ultraviolet (UV)/blue and in the red spectral region as well as differences in the kinetics and in the unfluence of dark pre-incubation on the stimulation of respiratory O₂ uptake by blue and red light strongly indicate the action of two photoreceptors. Since the effect of red light shows some far-red reversibility, and since simultaneous irradiation with red and far-red light decreases the effectiveness of red light, the involvement of phytochrome — in addition to the UV/blue photoreceptor(s) — is suggested in the light-stimulated respiration inDunaliella.Abbreviation UV ultraviolet     

Photocontrol of Spirodela intermedia flavonoids

Clone 115 of Spirodela intermedia W. Koch grown in Hutner's medium with sucrose produces the glycoflavones vitexin and orientin in darkness or in light of various wavelengths. The anthocyanin cyanidin-3-monoglucoside was present only after prolonged illumination of the plants with white or blue light. No cyanidin-glucoside was formed under constant red light. The substitution of red, blue, or far-red light for the last 24 hours of culture under constant white light reduced each flavonoid over those maintained in white light or given 24 hours of darkness. Reducing the light intensity from 900 to 400 ft-c of constant cool-white fluorescent light had no appreciable influence on vitexin (4′-hydroxyl) but markedly reduced orientin and cyanidin-glucoside (both 3′4′-hydroxyl). Substituting alternate 12-hour periods of light and darkness for continuous light reduced the glycoflavones approximately 50% while cyanidin-glucoside was reduced about 85%. Most responses to red, blue, or far-red light are consistent with a phytochrome-controlled promotion of vitexin synthesis.

The evidence suggests that in S. intermedia: A) Environmental conditions which elicit cyanidin-glucoside and glycoflavone synthesis are different since a prolonged illumination with white light is required for the former but not the latter. B) The availability of a 3′4′-hydroxyl precursor for orientin and anthocyanin probably limits their synthesis in low intensity light. Since vitexin is essentially unaltered under these conditions this also suggests that acetate or malonate units for the A-ring and the deamination products of aromatic amino acids for the B-ring and carbons of the C-ring are not limiting factors. C) Light controls the biosynthesis of flavonols in the same manner as glycoflavones; under all experimental conditions the synthesis of kaempferol paralleled vitexin while quercetin responded in the same manner as crientin.

     

Loss of short-day requirement for dormancy breaking of bulbils from gibberellic acid-treated plants in Sedum bulbiferum

Sedum bulbiferum forms bulbils at superterranean nodes under long-day conditions, and the detached bulbils sprout after exposure to short-days [1]. When gibberellic acid was applied to the mother plant at the start of the long-day induction period, the number of bulbils formed increased slightly and these bulbils sprouted on incubation in the dark but not under short days of continuous light. However, when gibberellic acid was applied directly to detached bulbils during incubation, the short-day requirement for sprouting was conserved. Gibberellic acid application to the mother plants enhanced sprouting ability of detached bulbils when incubated under illumination with blue, green or far-red light. However, presence of gibberellic acid during bulbil exposure to light did not induced marked enhancement in sprouting under blue, green of far-red light. Thus, gibberellic acid application to the mother plant modified light and photoperiodic requirements for the sprouting of detached bulbils of S. bulbiferum.     

Flowering responses to light-breaks in photomorphogenic mutants of Arabidopsis thaliana, a long-day plant

Flowering response and plant form of photomorphogenic mutants (hy1, hy2, hy3, hy4 and hy5) of Arabidopsis thaliana (L.), a long-day plant, were examined in long and short days. There were only slight differences among genotypes including Landsberg wild type with respect to the flowering time under long days. The effect of 1 h light-(night)-breaks of far-red, red, blue and white light given in the middle of the dark period of plants grown under short days, was studied. Effects of far-red light applied at the end or the beginning of the main photoperiod on flowering and plant form were also examined. The light-breaks with all the above mentioned light qualities promoted floral initiation of all the genotypes including the wild type in terms of both the flowering time and the number of rosette leaves. In general, far-red light was most effective. It is possible to classify the hy-mutants into 3 groups by their responses to light-breaks under short day conditions: (a) Mutants hy2 and hy3, which have a reduced number of rosette leaves, and flower early. Red light is as effective as far-red light. The wavelength of light-breaks is relatively unimportant for flowering response. (b) Mutants hy4, hy5 and Landsberg wild type, which have a greater number of rosette leaves, and flower relatively late. The effectiveness of light-breaks is in the following order, far-red, blue, and red light, which is in reverse order to the transformation of phytochrome to the P_fr form. (c) Mutant hy1, which behaves anomalously with respect to relations between flowering time and number of rosette leaves; late flowering with reduced number of rosette leaves. Red, blue and far-red light are effective, but white light is ineffective for reducing the number of rosette leaves. When far-red light was given in the middle of the night or at the end of the main photoperiod, it markedly reduced the number of rosette leaves compared to those grown under short days for all the genotypes, while when applied at the beginning of the main photoperiod far-red light did not affect the number of rosette leaves. Different effects on the plant form dependent on the time of treatment with far-red light-breaks are also discussed.     

Perception of solar UV radiation by plants: photoreceptors and mechanisms

About 95% of the ultraviolet (UV) photons reaching the Earth’s surface are UV-A (315–400 nm) photons. Plant responses to UV-A radiation have been less frequently studied than those to UV-B (280–315 nm) radiation. Most previous studies on UV-A radiation have used an unrealistic balance between UV-A, UV-B, and photosynthetically active radiation (PAR). Consequently, results from these studies are difficult to interpret from an ecological perspective, leaving an important gap in our understanding of the perception of solar UV radiation by plants. Previously, it was assumed UV-A/blue photoreceptors, cryptochromes and phototropins mediated photomorphogenic responses to UV-A radiation and “UV-B photoreceptor” UV RESISTANCE LOCUS 8 (UVR8) to UV-B radiation. However, our understanding of how UV-A radiation is perceived by plants has recently improved. Experiments using a realistic balance between UV-B, UV-A, and PAR have demonstrated that UVR8 can play a major role in the perception of both UV-B and short-wavelength UV-A (UV-A_sw, 315 to ∼350 nm) radiation. These experiments also showed that UVR8 and cryptochromes jointly regulate gene expression through interactions that alter the relative sensitivity to UV-B, UV-A, and blue wavelengths. Negative feedback loops on the action of these photoreceptors can arise from gene expression, signaling crosstalk, and absorption of UV photons by phenolic metabolites. These interactions explain why exposure to blue light modulates photomorphogenic responses to UV-B and UV-A_sw radiation. Future studies will need to distinguish between short and long wavelengths of UV-A radiation and to consider UVR8’s role as a UV-B/UV-A_sw photoreceptor in sunlight.

In sunlight, UVR8 mediates the perception of both UV-B and short-wavelength UV-A radiation with its sensitivity moderated by blue light perceived through cryptochromes.     

Effects of light quality on the circadian rhythm of leaf movement of a short-day-plant

Studies were made of the effects of blue, green, red and far-red (FR) light on the circadian rhythm of leaf movement of Coleus blumei × C. frederici, a short day plant. Under continuous illumination with blue light, there was a significant lengthening of the period of the rhythm to about 24.0 hr, as compared to 22.5 hr in continuous darkness. Under continuous red light, the period length was significantly shortened to 20.5 hr. Under continuous green or FR, the period length was not significantly different from the dark control. It was observed that under continuous FR illumination, the leaves tended to oscillate in a more downward position. Eight-hr red light signals were effective in advancing the phase of the rhythm as compared to a control under continuous green light. Blue light signals were effective in delaying the phase of the rhythm. FR light signals were ineffective in producing either delay or advance phase shifts. Far-red light did not reverse the effects of either red or blue light signals. On the basis of these results it is suggested, that pigments which absorb blue or red light, rather than phytochrome, mediate the effect of light on the circadian rhythm of leaf movement.     

Die Wirkung von „Störlicht” auf die Blütenbildung von Sinapis alba L.

Summary The induction of flowering in mustard (Sinapis alba L.) was studied by means of night-breaks (Störlicht). The plants were cultivated under fully controlled conditions: 8000 Lux white light (mixed fluorescent and incandescent) 18°C, 80% relative humidity. Raised under our conditions in short days (8 hours of white light) mustard behaved as a quantitative long-day plant (Fig. 2). Flowering can be promoted by long-day treatment (Fig. 3). The long day (16 hours of white light) can be replaced by a short day plus a night-break. The highest effectiveness of the night-break is found near the middle of the dark period (Figs. 4, 5). —The spectral dependence of flower induction was studied with blue, green, yellow, red (Fig. 1) and far-red light using a 2-hour break near the middle of the dark period. The dose response curves (Fig. 6) and the action spectrum (Fig. 7) indicate a very strong effectiveness in the blue part of the spectrum, a small response in red and yellow light and no response at all in green and far-red light. The participation of phytochrome is indicated (Table 1), but no far-red reversibility could be detected (Table 2). Simultaneous irradiation with red and far-red light yielded significant enhancement effects (Fig. 8). In view of the strong shadowing in the leaves (Figs. 9, 10) these data are interpretable on the basis of phytochrome.     

The effects of blue and far red light on rhythmic leaflet movements in samanea and albizzia

The opening of excised Samanea saman pulvini is promoted by prolonged blue or far-red irradiation. Far-red effects are attributed partially but not completely to lowering of the Pfr level. Two hours of continuous or pulsed blue light or pulsed far-red light (total dosage = 2.2 × 10¹⁸ quanta per square centimeter in all cases) also phase shifts the rhythm in Samanea while two hours of continuous blue light phase shifts the rhythm in the related plant Albizzia julibrissin. The same pigments appear to regulate opening and rhythmic phase shifting. The blue light-induced phase response curve has smaller advance and delay peaks and differs in shape from the curve induced by brief red light pulses absorbed by phytochrome. The blue absorbing pigment has not been identified, but it does not appear to be phytochrome acting in a photoreversible mode.     

Far-Red Light Reflection from Green Leaves and Effects on Phytochrome-Mediated Assimilate Partitioning under Field Conditions

The influence of plant spacing and row orientation on spectral distribution of light received by growing soybean (Gylcine max [L.] Merr.) plants was measured under field conditions. Light absorption, reflection and transmission of individual leaves showed that most of the blue and red was absorbed while most of the far-red was either reflected or transmitted. Plants growing in the field received different ratios of far-red relative to red, depending on nearness and/or orientation of other vegetation. Plants grown in close-spaced rows, or high population densities, received higher far-red/red ratios than did those grown in wide rows, or sparse populations. Heliotropic movements of the leaves also contributed to the far-red reflection patterns associated with row orientation. Under field conditions, differences in far-red/red ratios associated with nearness of competing vegetation became more pronounced with low solar angle near the end of the day. Plants exposed to far-red for 5 minutes at the end of each day in controlled environments, and those grown in close-spaced rows in the field, developed longer internodes and fewer branches. Red, far-red photoreversibility in the controlled environment study indicated involvement of phytochrome. Dry matter partitioning among plant components in the field was related to far-red/red light ratio received during growth and development.     

Light quantity and quality interactions in the control of elongation growth in light-grown Chenopodium rubrum L. seedlings

The spectral control of hypocotyl elongation in light-grown Chenopodium rubrum L. seedlings has been studied. The results showed that although the seedlings responded to changes in the quantity of combined red and far-red radiation, they were also very sensitive to changes in the quantity of blue radiation reaching the plant. Altering the proportion of red: far-red radiation in broad waveband white light caused marked differences in hypocotyl extension. Comparison of the responses of green and chlorophyll-free seedlings indicated no qualitative difference in the response to any of the light sources used, although photosynthetically incompetent plants were more sensitive to all wavelengths. Blue light was found to act primarily of a photoreceptor which is different from phytochrome. It is concluded that hypocotyl extension rate in vegetation shade is photoregulated by the quantity of blue light and the proportion of red: far-red radiation. In neutral shade, such as that caused by stones or overlying soil, hypocotyl extension appears to be regulated primarily by the quantity of light in the blue waveband and secondarily by the quantity of light in the red and far-red wavebands.Abbreviations B blue - FR far-red - k ₁, k ₂ rate constants for photoconverison of Pr to Pfr and Pfr to Pr, respective - k ₁/k ₁ +k ₂= phytochrome photoequilibrium - k ₁ +k ₂= phytochrome cycling rate - Pr=R absorbing form of phytochrome - Pfr=FR absorbing form of phytochrome - P_tot Pr+Pfr - PAR photosynthetically active radiation = 400–700 nm - R red - WL white light     

The regulation of cell wall extensibility during shade avoidance: a study using two contrasting ecotypes of Stellaria longipes

Shade avoidance in plants involves rapid shoot elongation to grow toward the light. Cell wall-modifying mechanisms are vital regulatory points for control of these elongation responses. Two protein families involved in cell wall modification are expansins and xyloglucan endotransglucosylase/hydrolases. We used an alpine and a prairie ecotype of Stellaria longipes differing in their response to shade to study the regulation of cell wall extensibility in response to low red to far-red ratio (R/FR), an early neighbor detection signal, and dense canopy shade (green shade: low R/FR, blue, and total light intensity). Alpine plants were nonresponsive to low R/FR, while prairie plants elongated rapidly. These responses reflect adaptation to the dense vegetation of the prairie habitat, unlike the alpine plants, which almost never encounter shade. Under green shade, both ecotypes rapidly elongate, showing that alpine plants can react only to a deep shade treatment. Xyloglucan endotransglucosylase/hydrolase activity was strongly regulated by green shade and low blue light conditions but not by low R/FR. Expansin activity, expressed as acid-induced extension, correlated with growth responses to all light changes. Expansin genes cloned from the internodes of the two ecotypes showed differential regulation in response to the light manipulations. This regulation was ecotype and light signal specific and correlated with the growth responses. Our results imply that elongation responses to shade require the regulation of cell wall extensibility via the control of expansin gene expression. Ecotypic differences demonstrate how responses to environmental stimuli are differently regulated to survive a particular habitat.     

Integration of light signaling with photoperiodic flowering and circadian rhythm

PLANT DE-ETIOLATION IS TRIGGERED BY LIGHT SIGNALS Light is arguably the most important resource for plants, and plants have evolved an array of photosensory pig- ments enabling them to develop optimally in a broad range of ambient light conditions. The ph…     

Light-quality and irradiance effects on pigments, light-harvesting proteins and Rubisco activity in a chlorophyll- and light- harvesting-deficient soybean mutant

Soybean ( Glycine max [L.] Merrill) plants, normal green (Clark L1) and mutant yellow (Clark y₉y₉), were grown in (1) full-spectrum solar irradiation; (2) either red plus far-red or blue plus far-red; (3) either red or blue light with no far-red light. Young leaves harvested from first (1TF) trifoliolate or fifth (5TF) trifoliolate stages of development showed that the mutant plants express pigment and protein deficiencies as a direct function of irradiance. Response of the mutant to light quality indicates that blue light slightly enhances expression of the mutation at higher irradiances. Direct response of light-harvesting proteins of photosystem 2 (LHCP2) and light-harvesting protein of photosystem 1 (LHCP1) to light quality increases the ratio of LHCP1/LHCP2 in blue light compared to that in red or red/far-red light. Rubisco proteins and Rubisco activity (leaf area basis) are directly related to irradiance level but are enhanced in blue light over equal irradiance red. This enhancement is not shown in the presence of far-red light.