Interaction of red to far red light ratio and ethylene in regulating stem elongation of <Emphasis Type="Italic">Helianthus annuus</Emphasis>

Sunflower (Helianthus annuus L.) stems showed increased elongation under two types of vegetative shade: canopy shade (low red to far red [R/FR] ratio) and neighbouring proximity shade (FR enrichment). Hypocotyls also elongated more under narrow-band FR light than under narrow-band R light. Ethylene levels were determined in actively elongating 7-day-old hypocotyls and 17-day-old internodes under three R/FR ratios. Ethylene levels were lower in both sunflower hypocotyls and internodes when the R/FR ratio was reduced. Both FR enrichment of normal R/FR ratio and narrow-band FR light with very low light irradiance resulted in reduction in ethylene levels in 7-day-old hypocotyls. Further, in application experiments, sunflower stems grown under low R/FR ratio were more sensitive to ethephon and less sensitive to aminoethoxyvinylglycine (AVG) than stems grown under high R/FR ratio. Low R/FR ratio appears to initiate reduction in ethylene levels in sunflower seedlings, allowing maximum stem elongation. These results, and findings of other authors, suggest that various plant species may have developed different ways of regulating stem elongation and ethylene levels in response to low R/FR ratio.     

Temperature‐dependent shade avoidance involves the receptor‐like kinase ERECTA

Plants detect the presence of neighbouring vegetation by monitoring changes in the ratio of red (R) to far‐red (FR) wavelengths (R:FR) in ambient light. Reductions in R:FR are perceived by the phytochrome family of plant photoreceptors and initiate a suite of developmental responses termed the shade avoidance syndrome. These include increased elongation growth of stems and petioles, enabling plants to overtop competing vegetation. The majority of shade avoidance experiments are performed at standard laboratory growing temperatures (>20°C). In these conditions, elongation responses to low R:FR are often accompanied by reductions in leaf development and accumulation of plant biomass. Here we investigated shade avoidance responses at a cooler temperature (16°C). In these conditions, Arabidopsis thaliana displays considerable low R:FR‐mediated increases in leaf area, with reduced low R:FR‐mediated petiole elongation and leaf hyponasty responses. In Landsberg erecta, these strikingly different shade avoidance phenotypes are accompanied by increased leaf thickness, increased biomass and an altered metabolite profile. At 16°C, low R:FR treatment results in the accumulation of soluble sugars and metabolites associated with cold acclimation. Analyses of natural genetic variation in shade avoidance responses at 16°C have revealed a regulatory role for the receptor‐like kinase ERECTA.     

Growth and ethylene evolution by shade and sun ecotypes of Stellaria longipes in response to varied light quality and irradiance

Plants growing in the shade receive both low light irradiance and light enriched in far red (FR) (i.e., light with a low red (R) to FR ratio). In an attempt to uncouple the R/FR ratio effects from light irradiance effects, we utilized Stellaria longipes because this species has two distinct natural population ecotypes, alpine (dwarf) and prairie (tall). The alpine population occupies the open, sun habitat. By contrast, the prairie population grows in the shade of other plants. Both 'sun' and 'shade' ecotypes responded with increased stem elongation responses under low irradiance, relative to growth under 'normal' irradiance, and this increased growth was proportionally similar. However, only the shade ecotype had increased shoot elongation in response to a low R/FR ratio. By contrast, the sun ecotype showed increased stem elongation in response to increasing R/FR ratio. Varying the R/FR ratios had no significant effect on ethylene evolution in either sun or shade ecotype. Under low irradiance, only the sun ecotype showed a significantly changed (decreased) ethylene evolution. We conclude that R/FR ratio and irradiance both regulate growth, and that irradiance can also influence ethylene evolution of the sun ecotype. By contrast, R/FR ratio and irradiance, while having profound influences on growth of the shade ecotype, do not appear to regulate these growth changes via effects on ethylene production.     

Differential response of Scots pine seedlings to variable intensity and ratio of red and far‐red light

We investigated the response to increasing intensity of red (R) and far‐R (FR) light and to a decrease in R:FR ratio in Pinus sylvestris L. (Scots pine) seedling. The results showed that FR high‐irradiance response for hypocotyl elongation may be present in Scots pine and that this response is enhanced by increasing light intensity. However, both hypocotyl inhibition and pigment accumulation were more strongly affected by the R light compared with FR light. This is in contrast to previous reports in Arabidopsis thaliana (L.) Heynh. In the angiosperm, A. thaliana R light shows an overall milder effect on inhibition of hypocotyl elongation and on pigment biosynthesis compared with FR suggesting conifers and angiosperms respond very differently to the different light regimes. Scots pine shade avoidance syndrome with longer hypocotyls, shorter cotyledons and lower chlorophyll content in response to shade conditions resembles the response observed in A. thaliana. However, anthocyanin accumulation increased with shade in Scots pine, which again differs from what is known in angiosperms. Overall, the response of seedling development and physiology to R and FR light in Scots pine indicates that the regulatory mechanism for light response may differ between gymnosperms and angiosperms.     

Plasticity and environment-specific covariances: an investigation of floral-vegetative and within flower correlations

Floral traits are commonly thought to be more canalized than vegetative ones. In addition, floral and vegetative traits are hypothesized to be genetically decoupled, enabling vegetative structures to respond plastically to environmental heterogeneity, and to evolve in response to selection without disrupting the reproductive function of flowers. To test these hypotheses, we evaluate the genetic architecture of floral and vegetative traits in natural populations of Arabidopsis thaliana raised under variable light-quality environments. Plants were grown either under high or low ratios of red to far-red (R:FR) light, an aspect of light quality that varies with neighbor proximity and regulates competitive shade-avoidance responses. Across environments, we detected significant genetic variation for the average expression of all measured floral traits (petal length and width, stamen length, pistil length, stigma-anther separation, and exsertion of both the stamen and pistil beyond the corolla). Light quality significantly influenced the absolute size of several floral traits as well as the allometry (i.e., relative scaling) of all floral traits, and genotypes differed in the plasticity of floral traits to the light treatments. Exposure to low relative to high R:FR resulted in significantly greater elongation in the vegetative trait, petiole length, and genotypes again differed in the plasticity of this trait to R:FR. Consistent with prior studies, most floral traits were less plastic than the vegetative trait; herkogamy (i.e., stigma-anther separation) was the exception and expressed more variable trait values across environments than petiole length, apparently as a consequence of the independent responses of stamens and pistils. Flowers also showed strong phenotypic integration; genotypic correlations were significantly positive among floral traits within each light treatment. Although floral-vegetative correlations were not significant in the high R:FR light treatment, significant correlations were detected between petal traits, pistil length, and petiole length under low R:FR, in contrast to the widely held hypothesis that floral and vegetative traits are genetically independent. Finally, we detected selection for reduced herkogamy in the low R:FR light treatment. The observed correlation between functional trait groups suggest that vegetative plasticity may affect the expression of floral traits in some environments, and that environment-specific constraints may exist on the evolution of floral and vegetative traits.     

THE DEVELOPMENTAL RESPONSES OF PAPAYA LEAVES TO SIMULATED CANOPY SHADE

The developmental responses of plants to shade underneath foliage are influenced by reductions in irradiance and shifts in spectral quality (characterized by reductions in the quantum ratio of red to far-red wavelengths, R:FR). Previous research on the influence of shadelight on leaf development has neglected the reductions in R:FR characteristic of foliage shade, and these studies have almost certainly underestimated the extent and array of developmental responses to foliage shade. We have studied the effects of reduced irradiance and R:FR on the leaf development of papaya (Carica papaya L., Caricaceae). Using experimental shadehouses, replicates of plants grown in high light conditions (0.20 of sunlight and R:FR = 0.90) were compared to low light conditions (0.02 of sunlight) with either the spectral quality of sunlight (R:FR = 0.99) or of foliage shade (F:FR = 0.26). Although many characteristics, such as leaf thickness, specific leaf weight, stomatal density, palisade parenchyma cell shape, and the ratio of mesophyll air surface/leaf surface were affected by reductions in irradiance, reduced R:FR contributed to further changes. Some characters, such as reduced chlorophyll a/b ratios, reduced lobing, and greater internode length, were affected primarily by low R:FR. The reduced R:FR of foliage shade, presumably affecting phytochrome equilibrium, strongly influences the morphology and anatomy of papaya leaves.     

The interaction of light quality and irradiance with gibberellins, cytokinins and auxin in regulating growth of Helianthus annuus hypocotyls

A reduced red to far-red (R/FR) light ratio and low photosynthetically active radiation (PAR) irradiance are both strong signals for inducing etiolation growth of plant stems. Under natural field conditions, plants can be exposed to either a reduced R/FR ratio or lower PAR, or to a combination of both. We used Helianthus annuus L., the sunflower, to study the effect of reduced R/FR ratio, low PAR or their combination on hypocotyl elongation. To accomplish this, we attempted to uncouple light quality from light irradiance as factors controlling hypocotyl elongation. We measured alterations in the levels of endogenous gibberellins (GAs), cytokinins (CKs) and the auxin indole-3-acetic acid (IAA), and the effect of exogenous hormones on hypocotyl growth. As expected, both reduced R/FR ratio and lower PAR can significantly promote sunflower hypocotyl elongation when given separately. However, providing the reduced R/FR ratio at a low PAR resulted in the greatest hypocotyl growth, and this was accompanied by significantly higher levels of endogenous IAA, GA1, GA8, GA20 and of a wide range of CKs. Providing a reduced R/FR ratio under normal PAR also significantly increased growth and again gave significantly higher levels of endogenous IAA, GAs and CKs. However, only under the de-etiolating influence of a normal R/FR ratio did lowering PAR significantly increase levels of GA1, GA8 and GA20. We thus conclude that light quality (e.g. the R/FR ratio) is the most important component of shade for controlling hypocotyl growth and elevated growth hormone content.     

Light quality regulation of endogenous levels of auxin,abscisic acid and ethylene production in petioles and leaves of wild type and ACC deaminase transgenic <Emphasis Type="Italic">Brassica napus</Emphasis> seedlings

Brassica napus L. seedlings responded to low red to far-red (R/FR) ratio by elongating petioles and decreasing leaf expansion. These typical shade avoidance traits were correlated with significantly decreased endogenous indole-3-acetic acid (IAA) levels and significantly increased endogenous abscisic acid (ABA) levels and ethylene production. The transgenic (T) B. napus line bearing the bacterial ACC deaminase gene, did not respond to low R/FR ratio with altered petiole and leaf growth and less ethylene (especially by petioles) was produced. As with WT seedlings, T seedlings had significantly lower IAA levels in both petioles and leaves under low R/FR ratio. However, ABA levels of low R/FR ratio-grown T seedlings either increased (petioles) or were unaltered (leaves). Our results further suggest that low R/FR ratio regulates endogenous IAA levels independently of ethylene, but there may be an interaction between ABA and ethylene in leaf development.     

phyB-1 sorghum maintains responsiveness to simulated shade, irradiance and red light: far-red light

The sorghum [Sorghum bicolor (L.) Moench] phyB-1 mutant exhibits a constitutive shade-avoidance phenotype including excessive shoot elongation. It was previously shown that this mutant also overproduces ethylene. Although phytochrome B (phyB) is assumed to be the pigment most important in sensing and transducing shade signals, the sorghum phyB-1 mutant still responds to light signals characteristic of shade. Specifically, it was determined that the leaf blade : leaf sheath elongation of phyB-1 is responsive to red : far red (R : FR), but this response is opposite that of wild type (WT). Reducing the photosynthetic photon flux density (PPFD) strongly reduced the leaf blade : leaf sheath of WT but did not affect phyB-1, demonstrating a role for phyB in sensing PPFD. Using light-emitting diode (LED) lighting, it was found that WT ethylene production was increased with low R : FR while PPFD had no effect. Conversely, phyB-1 ethylene production increased only with high PPFD, high R : FR which was the treatment resulting in the least ethylene production by WT. Elevated ethylene production inhibits shoot elongation, but may contribute to shade avoidance by reducing leaf blade : leaf sheath elongation. Ethylene responses to light treatments designed to promote or reduce phytochrome A (phyA) activity, and the analysis of PHYA levels in the two cultivars suggests that phyA could be involved in transducing shade signals in light-grown sorghum. Responses potentially tranduced by phyA are elevated in phyB-1 which also over-expresses PHYA.     

Uncoupling light quality from light irradiance effects in Helianthus annuus shoots: putative roles for plant hormones in leaf and internode growth

An attempt has been made to uncouple the effects of the two primary components of shade light, a reduced red to far-red (R/FR) ratio and low photosynthetically active radiation (PAR), on the elongation of the youngest internode of sunflower (Helianthus annuus) seedlings. Maximal internode growth (length and biomass) was induced by a shade light having a reduced R/FR ratio (0.85) under the low PAR of 157 micromol m(-2) s(-1). Reducing the R/FR ratio under normal PAR (421 micromol m(-2) s(-1)) gave similar growth trends, albeit with a reduced magnitude of the response. Leaf area growth showed a rather different pattern, with maximal growth occurring at the higher (normal) PAR of 421 micromol m(-2) s(-1)), but with variable effects being seen with changes in light quality. Reducing the R/FR ratio (by enrichment with FR) gave significant increases in gibberellin A(1) (GA(1)) and indole-3-acetic acid (IAA) contents in both internodes and leaves. By contrast, a lower PAR irradiance had no significant effect on GA(1) and IAA levels in internodes or leaves, but did increase the levels of other GAs, including two precursors of GA(1). Interestingly, both leaf and internode hormone content (GAs, IAA) are positively and significantly correlated with growth of the internode, as are leaf levels of abscisic acid (ABA). However, changes in these three hormones bear little relationship to leaf growth. By implication, then, the leaf may be the major source of GAs and IAA, at least, for the rapidly elongating internode. Several other hormones were also assessed in leaves for plants grown under varying R/FR ratios and PARs. Leaf ethylene production was not influenced by changes in R/FR ratio, but was significantly reduced under the normal (higher) PAR, the irradiance treatment which increased leaf growth. Levels of the growth-active free base and riboside cytokinins were significantly increased in leaves under a reduced R/FR ratio, but only at the higher (normal) PAR irradiance; other light quality treatments evoked no significant changes. Taken in toto, these results indicate that both components of shade light can influence the levels of a wide range of endogenous hormones in internodes and leaves while evoking increased internode elongation and biomass accumulation. However, it is light quality changes (FR enrichment) which are most closely tied to increased hormone content, and especially with increased GA and IAA levels. Finally, the increases seen in internode and leaf GA content with a reduced R/FR ratio are consistent with FR enrichment inducing an overall increase in sunflower seedling GA biosynthesis.     

Responses of carbon acquisition traits to irradiance and light quality in Mercurialis annua (Euphorbiaceae): evidence for weak integration of plastic responses

It is often suggested that traits will be integrated, either because of pleiotropy or because natural selection may favor suites of integrated traits. Plant responses to different environments can provide evidence of such integration. We grew Mercurialis annua plants in high-density stands in high irradiance, in neutral shade, and in high red to far-red (R:FR) shade, resulting in environments of high irradiance, low R:FR; low irradiance, low R:FR; and low irradiance, high R:FR. We measured gas exchange, leaf morphology, stem elongation, and biomass traits and tested the prediction that traits within each functional group would show higher trait integration, as evidenced by high correlations among traits within environments, higher correlations of trait plasticity, and lower plasticity of trait correlations. Overall, we found evidence of only moderate integration for some groups of traits. Functionally related groups of traits, or pairs of traits, could be strongly integrated by one criterion but weakly integrated by another of the criteria. Stem elongation traits, though often observed to be strongly integrated in other taxa, showed little evidence of integration. Internode traits exhibited a novel pattern of responses to low R:FR, with increased elongation of the hypocotyl, decreased elongation of the first internode, and no change in the second internode. We propose that these responses to light are more likely to be the result of natural selection than the consequence of constraints imposed by pleiotropy.     

Light from below matters: Quantifying the consequences of responses to far‐red light reflected upwards for plant performance in heterogeneous canopies

In vegetation stands, plants receive red to far‐red ratio (R:FR) signals of varying strength from all directions. However, plant responses to variations in R:FR reflected from below have been largely ignored despite their potential consequences for plant performance. Using a heterogeneous rose canopy, which consists of bent shoots down in the canopy and vertically growing upright shoots, we quantified upward far‐red reflection by bent shoots and its consequences for upright shoot architecture. With a three‐dimensional plant model, we assessed consequences of responses to R:FR from below for plant photosynthesis. Bent shoots reflected substantially more far‐red than red light, causing reduced R:FR in light reflected upwards. Leaf inclination angles increased in upright shoots which received low R:FR reflected from below. The increased leaf angle led to an increase in simulated plant photosynthesis only when this low R:FR was reflected off their own bent shoots and not when it reflected off neighbour bent shoots. We conclude that plant response to R:FR from below is an under‐explored phenomenon which may have contrasting consequences for plant performance depending on the type of vegetation or crop system. The responses are beneficial for performance only when R:FR is reflected by lower foliage of the same plants.     

Variation in growth form in relation to spectral light quality (red/far-red ratio) in Plantago lanceolata L. in sun and shade populations

Plants from a sun and shade population were grown in two environments differing in the ratio of red to far-red light (R/FR ratio). A low R/FR ratio, simulating vegetation shade, promoted the formation of long, upright-growing leaves and allocation towards shoot growth, whereas a high R/FR ratio had the opposite effects. The increase in plant height under the low R/FR ratio was accompanied by a reduction in the number of leaves. Population differences in growth form resembled the differences between plants grown in different light environments: plants from the shade population had rosettes with long erect leaves, whereas plants from the sun population formed prostrate rosettes with short leaves. Plants from the shade population were more responsive to the R/FR ratio than plants from the sun population: the increases in leaf length, plant height, and leaf area ratio under a low R/FR ratio were larger in the shade population. However, differences in plasticity were small compared to the population difference in growth form itself. We argue that plants do not respond optimally to shading and that developmental constraints might have limited the evolution of an optimal response. Received: 8 December 1996 / Accepted: 31 March 1997     

Regulation of the photosynthetic capacity of primary bean leaves by the red:far-red ratio and photosynthetic photon flux density of incident light

The main objective of the present work was to examine the effects of the red:far-red ratio (R:FR) prevailing during leaf development on the photosynthetic capacity of mature leaves. Plants of Phaseolus vulgaris L. cv. Balin de Albenga were grown from time of emergence in a controlled environment room, 25 ± 3°C, 12-h photoperiod, with different light treatments:a) high photosynthetic photon flux density (PPFD) = 800 μmol m⁻¹ s⁻¹+ high R:FR= 1.3;b) low PPFD= 300 μmol m⁻² s⁻¹+ high R:FR= 1.3; c) high PPFD=800 μmol m⁻² s⁻¹+ low R:FR= 0.7; d) low PPFD= 300 μmol m⁻²s⁻¹+ low R:FR=0.7. With an R:FR ratio of 1.3, a decrease in irradiance during leaf growth reduced photosynthesis when measured at moderate to high PPFD; but when measured at low PPFD, leaves expanded under low irradiance actually had photosynthesis rates higher than those of leaves grown in high irradiance. A low R:FR ratio during development reduced the photosynthetic capacity of the leaves. In leaves expanded under R:FR = 0.7 and high irradiance photosynthesis was reduced by 42 to 89%, depending on the PPFD at which measurements were made, whereas for leaves developed at R:FR = 0.7 and low irradiance photosynthesis decreased by 21 to 24%, compared to leaves under R:FR = 1.3 and similar irradiance. The reduced photosynthetic capacity under R:FR = 0.7 and high irradiance. In natural environments, leaves may experience low R:FR conditions temporarily during their development, and this may affect their future photosynthetic capacity in full sunlight.     

Species-specific variation in the importance of the spectral quality gradient in canopies as a signal for photosynthetic resource partitioning

BACKGROUND AND AIMS: Plants adjust the distribution of photosynthetic capacity and chlorophyll to canopy density. The importance of the gradient in the red : far-red ratio (R : FR) relative to the irradiance gradient was studied for its perception with respect to this partitioning of photosynthetic resources. Whether the relative importance of these two signals varied between six species of different growth habit (Phaseolus vulgaris, Lysimachia vulgaris, Hedera helix, Ficus benjamina, Carex acutiformis and Brachypodium pinnatum) was investigated further. METHODS: Single leaves of plants were shaded in daylight by a spectrally neutral filter or a leaf. In another experiment, leaves were treated with supplemental FR. In most cases, treatment effects were evaluated after 2 weeks. KEY RESULTS: Nitrogen and photosynthetic capacity (Amax) per leaf area, parameters pertaining to between-leaf resource partitioning, were strongly reduced in neutral shade but not additionally by spectral leaf shade. Supplemental FR reduced these parameters also, except in Carex. Acceleration of induction of senescence was observed in spectral leaf shade in primary bean leaves. Amax per unit chlorophyll, a parameter pertaining to within-leaf resource partitioning, was reduced in neutral shade, but not in spectral leaf shade or supplemental FR. CONCLUSIONS: Signalling mechanisms associated with perception of the R : FR gradient in canopies were less important than those associated with the irradiance gradient for between-leaf and within-leaf partitioning of photosynthetic resources. The relative importance of the signals differed between species because Carex was the only species for which no indications were found for an involvement of the spectral gradient in perception of canopy density.     

Canopy studies on ethylene-insensitive tobacco identify ethylene as a novel element in blue light and plant-plant signalling

Plants growing at high densities express shade avoidance traits as a response to the presence of neighbours. Enhanced shoot elongation is one of the best researched shade avoidance components and increases light capture in dense stands. We show here that also leaf movements, leading to a more vertical leaf orientation (hyponasty), may be crucial in the early phase of competition. The initiation of shade avoidance responses is classically attributed to the action of phytochrome photoreceptors that sense red:far-red (R:FR) ratios in light reflected by neighbours, but also other signals may be involved. It was recently shown that ethylene-insensitive, transgenic (Tetr) tobacco plants, which are insensitive to the gaseous plant hormone ethylene, have reduced shade avoidance responses to neighbours. Here, we report that this is not related to a reduced response to low R:FR ratio, but that Tetr tobacco plants are unresponsive to a reduced photon fluence rate of blue light, which normally suppresses growth inhibition in wild-type (WT) plants. In addition to these light signals, ethylene levels in the canopy atmosphere increased to concentrations that could induce shade avoidance responses in WT plants. Together, these data show that neighbour detection signals other than the R:FR ratio are more important than previously anticipated and argue for a particularly important role for ethylene in determining plant responses to neighbours.     

藤本植物Potentilla reptans叶对风和遮阴的响应

生长在一个密集植物群中的植株由于相互遮蔽而不可能对风的影响做出反应,因为这样的环境条件(有限的光资源)对由风导致的矮小表型植株的生长是不利的。为弄清在密集植物群体中生长的植株对风的响应,利用藤本植物Potentilla reptans的10种基因型做实验材料,在温室条件下(光照强度为日光照的50%,红光/远红光=1.2)模拟冠层遮阴(相当于15%的日光照,红光/远红光=0.3),研究了藤本植物叶对风的响应。结果表明,Potentilla reptans的10种基因型植株在冠层遮阴下(低的红光/远红光)都表现出典型的避阴生长响应:较少的叶(叶生物量少),长而细但硬度系数高(higher Young's modulus)的叶柄;而受风影响的植株,无论遮阴或不遮阴,其植株的叶相对较多,叶柄短、粗且柔韧性强(lower Young's modulus),说明Potentilla reptans叶对风的响应并未因遮阴而被压抑,其可塑性变化不过是对复杂生境做出的一种生长权衡:尽可能增强抗风能力(矮壮)和获取最大光能(足够高而避免被遮光),即保证在存活下去的前提下获取最大的生长效率。     

Selected Components of the Shade-Avoidance Syndrome Are Displayed in a Normal Manner in Mutants of Arabidopsis thaliana and Brassica rapa Deficient in Phytochrome B

Several growth parameters associated with the phytochrome-mediated shade avoidance syndrome have been measured in seedlings and mature plants of a wild-type and a hy3 mutant of Arabidopsis thaliana deficient in phytochrome B. Growth parameters were compared in plants grown in either white light (high red:far-red [R:FR] ratio) or white light plus added far-red (FR) light (low R:FR ratio). Wild-type Arabidopsis exhibited increased hypocotyl and petiole extension under a low, compared with a high, R:FR ratio. The hy3 mutant did not respond to low R:FR ratio by increase in hypocotyl or petiole length. Extension growth of wild-type plants was stimulated by brief end-of-day FR pulses, but similar treatment had no effect on extension growth of hy3 mutant plants. However, some responses to low R:FR ratio seen in the wild-type plants were also evident in the hy3 mutants. The number of days to bolting, the developmental stage at bolting, the leaf area, and the specific stem weight (weight per unit of length) all decreased in the wild-type and hy3 seedlings in response to low R:FR ratio. Low R:FR ratio caused a larger decrease in leaf area and specific stem weight in the mutant seedlings than in wild-type seedlings. The effects of low R:FR ratio on leaf area and specific stem weight were opposite to those of the hy3 lesion, which resulted in increased leaf area and specific stem weight in comparison with the wild type. Both leaf area and specific stem weight responses to low R:FR ratio also were unchanged in the ein mutant of Brassica rapa, known to be deficient in phytochrome B. These responses represent components of the shade-avoidance syndrome, and, consequently, the results indicate that phytochrome B cannot be solely responsible for the perception of R:FR ratio and the induction of shade-avoidance responses. The hypothesis is proposed that different phytochromes may be responsible for the regulation of extension growth and the regulation of lateral or radial expansion.     

Control of ascorbic acid synthesis and accumulation and glutathione by the incident light red/far red ratio in Phaseolus vulgaris leaves

The effects of red/far red (R/FR) ratios on leaf ascorbate (AA) and glutathione (GSH) accumulation were examined in common bean (Phaseolus vulgaris L.). Growth under low R/FR ratios resulted in a “shade” phenotype and much lower leaf AA and GSH contents than high (R/FR) ratios. Photosynthesis rates were unaffected by changes in the R/FR ratio but leaf respiration rates, pyridine nucleotide pools and antioxidant enzyme activities were decreased under the low R/FR regime. The GSH pool changed slowly in response to altered R/FR ratios but leaf ascorbate acclimated over a single photoperiod. We conclude that light quality signals, particularly R/FR ratios, are important regulators of antioxidant synthesis and accumulation. These acclimatory changes are an early response to changing light environment.