Influence of spectral quality on the growth response of intact bean plants to brassinosteroid, a growth-promoting steroidal lactone. I. Stem elongation and morphogenesis

The growth response of bean ( Phaseolus vulgaris L. 'Pinto') plants treated with 5 μg of brassinosteroid (BR) in the bean second-internode assay was measured in a controlled environment under 3 radiation sources: cool white fluorescent (CWF), far-red (FR) fluorescent and incandescent (INC) lamps. Growth comparisons were made under equal levels (90 μmol s^-1 m^-2) of photosynthetic photon flux density provided by CWF or INC lamps and equal levels of far-red (28 W m^-2, 700–800 nm) radiation provided by the same INC or FR lamps. Treatment of the second internode with BR produced a sequential increase in elongation, curvature, and swelling under normal bioassay conditions (CWF lamps), as observed previously with brassins. In addition, BR induced marked splitting of the treated internode provided that ample photosynthate was available. Spectral quality had a differential effect on internodal elongation. Under CWF lamps, internodes, 6 days after BR-treatment, were 2–3 times longer than those of controls; under INC or FR lamps they were 30–60% shorter than those of controls. In all cases, BR-treatment greatly stimulated accumulation of photosynthate in the treated internode, as indicated by fresh and dry weights and stem diameter measurements. This suggests a possible mobilization role for BR in the intact plant. Brassinosteroid also partially overcame the natural inhibitory effects of CWF radiation on stem elongation.     

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)     

The semidian rhythm in flowering response of Pharbitis nil in relation to dark period time measurement and to a circadian rhythm

When seedlings of Pharbitis nil Choisy, cv. Violet, are exposed to a single inductive dark period at 27°C, brief interruptions with red light (R) can be promotive after 2–3 h of darkness but increasingly inhibitory to flowering up to the 8–9th h of darkness. This rhythmic response to R interruptions can be advanced in phase by > 1 h when the preceding light period is interrupted with far-red (FR) 2 h before darkness (FR -2 h) or with FR – 15 h, whereas FR –8 h or FR–22 h retard the rhythm. These shifts in the R interruption rhythm are paralleled by equal shifts in the length of the dark period required for flowering. Brief FR interruptions of darkness displayed a similar rhythm which was also advanced by FR –2 h and retarded by FR –8 h. We conclude therefore that the semidian rhythm in the light, which we have previously described, continues through at least the first 12 h of darkness, is manifested in the R interruption rhythm, and determines the critical night length. A circadian rhythm with a marked effect on flowering was also identified, but several lines of evidence suggest that the circadian and semidian rhythms have independent additive effects on flowering and do not appear to show phase interaction.     

Changes in photosensitive stem growth in intact peas following irradiation

Etiolated pea seedlings given a short red-light pretreatment followed by 30 hr of darkness no longer showed a typical red-light inhibition of internode elongation. The induction of phytochrome-insensitive growth was itself mediated by phytochrome, since far-red light reversed the effect of the short red-light pretreatment. Peas grown in white light showed a similar insensitivity to red light. However, in this instance the phytochrome system exerted some control over internode elongation since far-red light promoted growth slightly, and this effect was red-reversible.     

Requirement for far-red light to maintain secondary needle extension growth in northern but not southern populations of Pinus sylvestris (Scots pine)

Extension growth of secondary needles is under photoperiodic control in Pinus sylvestris . To test for the effects of far-red light on maintaining this extension growth, seedlings of six populations originating from latitudes between 57° and 67°N were raised for 11 weeks in continuous incandescent (metal halogen) light at 300 µmol m⁻² s⁻¹ and 20°C and then transferred at the same temperature to a daily regime of 8 h incandescent light (230 µmol m⁻² s⁻¹) followed by a 16 h day extension with cool white fluorescent light (40 µmol m⁻² s⁻¹, R/FR ratio 7.5) or with incandescent lamps (20 µmol m⁻² s⁻¹, R/FR ratio 2.0). For the seedlings from the three populations north of 64°, needle extension growth over 42 days in the FR-poor day extension treatment was lower by up to 40% than in the FR-rich day extension treatment, whereas for the seedlings from the three southern populations the needle extension growth was similar in both day extension treatments. The requirement for FR in day extensions is characteristic of 'light-dominant' photoperiodic control mechanisms. It appears that P. sylvestris changes from dark-dominant night timekeeping to light-dominant day timekeeping with increasing latitude, as with the photoperiodic control of budset in Picea abies .     

Impaired phytochrome-mediated shade-avoidance responses in the aurea mutant of tomato

Internode extension-growth responses to neighbouring plants and to red to far-red ratios (R:FR) were investigated in wild-type (WT) and aurea (au)-mutant seedlings of tomato grown under natural radiation. The genomic location of the au mutant is not known, but one of its consequences is the reduced phytochrome level. In WT seedlings, internode growth was promoted by the presence of non-shading neighbours reflecting far-red light (FR), the shade of a tall canopy, FR provided as a supplement during the photoperiod, and FR pulses either provided at the end of the day or delayed into the dark period. Supplementary FR during the photoperiod also promoted growth in herbicide-treated partially bleached WT seedlings. The au mutant showed higher background extension-growth rates, but only responded to the most severe treatments: deep shade light and very low R:FR at the end of the day, i.e. au-mutant seedlings were less sensitive than WT seedlings to R:FR signals. Wild-type seedlings were transferred from the glasshouse to a growth room and exposed to white light with two levels of phytochrome-absorbable radiation but similar phytochrome photoequilibria and radiation for photosynthesis. The plants exposed to the lowest level showed a transient increase of internode extension growth rate and a simultaneous reduction of response to FR pulses, i.e. reproduced some of the features of au-mutant seedlings. Phytochrome itself could set the degree of response to Pfr during neighbour detection.     

Time to flowering of temperate pulses in vivo and generation turnover in vivo–in vitro of narrow-leaf lupin accelerated by low red to far-red ratio and high intensity in the far-red region

Understanding the role light quality plays on floral initiation is key to a range of pre-breeding tools, such as accelerated single-seed-descent. We have elucidated the effect of light quality on early flowering onset in cool-season grain legumes and developed predictive models for time to flowering under the optimised light conditions. Early and late flowering genotypes of pea, chickpea, faba bean, lentil and lupin were grown in controlled environments under different light spectra (blue and far red-enriched LED lights and metal halide). All species and genotypes showed a positive response to a decreasing red to far-red ratio (R:FR). In general, ratios above 3.5 resulted in the longest time to flowering. In environments with R:FR below 3.5, light with the highest intensity in the FR region was the most inductive. We demonstrate the importance of considering both relative (R:FR) and absolute (FR photons) light values for flower induction in grain legumes. Greater response to light spectra was observed in the later flowering genotypes, enabling a drastic compression of time to flowering between phenologically diverse genotypes. A novel protocol for robust in vitro germination of immature seeds was developed for lupin, a species known for its recalcitrance to in vitro manipulation. We show how combining this protocol with growth under conditions optimized for early flowering drastically speeds generation turnover. The improved understanding of the effect of light on flowering regulation and the development of robust in vitro culture protocols will assist the development and exploitation of biotechnological tools for legume breeding.     

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.     

Phytochrome D acts in the shade-avoidance syndrome in Arabidopsis by controlling elongation growth and flowering time

Shade avoidance in higher plants is regulated by the action of multiple phytochrome (phy) species that detect changes in the red/far-red ratio (R/FR) of incident light and initiate a redirection of growth and an acceleration of flowering. The phyB mutant of Arabidopsis is constitutively elongated and early flowering and displays attenuated responses to both reduced R/FR and end-of-day far-red light, conditions that induce strong shade-avoidance reactions in wild-type plants. This indicates that phyB plays an important role in the control of shade avoidance. In Arabidopsis phyB and phyD are the products of a recently duplicated gene and share approximately 80% identity. We investigated the role played by phyD in shade avoidance by analyzing the responses of phyD-deficient mutants. Compared with the monogenic phyB mutant, the phyB-phyD double mutant flowers early and has a smaller leaf area, phenotypes that are characteristic of shade avoidance. Furthermore, compared with the monogenic phyB mutant, the phyB-phyD double mutant shows a more attenuated response to a reduced R/FR for these responses. Compared with the phyA-phyB double mutant, the phyA-phyB-phyD triple mutant has elongated petioles and displays an enhanced elongation of internodes in response to end-of-day far-red light. These characteristics indicate that phyD acts in the shade-avoidance syndrome by controlling flowering time and leaf area and that phyC and/or phyE also play a role.     

The nature of floral signals in Arabidopsis. I. Photosynthesis and a far-red photoresponse independently regulate flowering by increasing expression of FLOWERING LOCUS T (FT)

Arabidopsis flowers in long day (LD) in response to signals transported from the photoinduced leaf to the shoot apex. These LD signals may include protein of the gene FLOWERING LOCUS T (FT) while in short day (SD) with its slower flowering, signalling may involve sucrose and gibberellin. Here, it is shown that after 5 weeks growth in SD, a single LD up-regulated leaf blade expression of FT and CONSTANS (CO) within 4-8 h, and flowers were visible within 2-3 weeks. Plants kept in SDs were still vegetative 7 weeks later. This LD response was blocked in ft-1 and a co mutant. Exposure to different LD light intensities and spectral qualities showed that two LD photoresponses are important for up-regulation of FT and for flowering. Phytochrome is effective at a low intensity from far-red (FR)-rich incandescent lamps. Independently, photosynthesis is active in an LD at a high intensity from red (R)-rich fluorescent lamps. The photosynthetic role of a single high light LD is demonstrated here by the blocking of the flowering and FT increase on removal of atmospheric CO(2) or by decreasing the LD light intensity by 10-fold. These conditions also reduced leaf blade sucrose content and photosynthetic gene expression. An SD light integral matching that in a single LD was not effective for flowering, although there was reasonable FT-independent flowering after 12 SD at high light. While a single photosynthetic LD strongly amplified FT expression, the ability to respond to the LD required an additional but unidentified photoresponse. The implications of these findings for studies with mutants and for flowering in natural conditions are discussed.     

Light Quality and Vernalization Interact in Controlling Late Flowering in Arabidopsis Ecotypes and Mutants

The late flowering ecotypes of Arabidopsis thaliana L. (Heyn.)Eifel, Pitztal and Innsbruck responded to 10 d vernalization(cold treatment) by flowering earlier with less with less thanhalf the number of leaves of non-induced plants. The vernalizationresponse was cumulative: increased numbers of days of vernalizationinduced earlier flowering up to an apparent saturation in responseafter 30 to 40 d. The ratio of red:far-red (R:FR) light alsoaffected non-vernalized time-to-flower. When grown under fluorescentplus incandescent lamps (R:FR = 1·0), time-to-flowerwas approximately half that required by plants grown under fluorescentlamps (R:FR = 5·8) at the same photon flux density andphotoperiod. Leaf production rate was unaffected by either vernalizationor light quality changes and time-to-flower and leaf numberwere highly correlated (r² = 0·973). The late flowering mutants of Landsberg erecta were grown underlighting which displayed a gradient of R:FR. Some mutants likeco, flowered at the same time in all R:FR treatment, while otherlike fca took nearly twice as long to flower, with double thenumber of leaves at R:FR ratio of 5·8 compared with theR:FR = 1 treatment. The ranking of the response from least tomost responsive was co, fe, gi, WT, fd, fwa, ft, fha, fpa, fy,fve and fca. Vernalization of these Landsberg mutants always resulted inearlier flowering, although only fca, fve, fy and fpa were significantlymore sensitive to thermoinduction than the wild type parent.There was a high correlation (r² = 0·89 between the responseto thermoinduction and to R:FR ratio. Vernalization of fca for24 d largely eliminated the R:FR time-to-flower response. Vernalizationand photoinduction similarly affect late flowering and can substitutefor each another.Copyright 1993, 1999 Academic Press Light quality, vernalization, flowering, Arabidopsis thaliana, phytochrome, thermoinduction, photoperiod, photoinduction, growth conditions, photon flux density, daylength, spectral quality, far-red light     

Influence of spectral quality on the growth response of intact bean plants to brassinosteroid, a growth-promoting steroidal lactone. II. Chlorophyll content and partitioning of assimilate

The chlorophyll content and partitioning of assimilate of bean ( Phaseolus vulgaris L. 'Pinto') plants were determined 6 days after treatment of the second internode (I₂ with 5 μg of brassinosteroid (BR), a growth-promoting steroidal lactone. Plants were grown for 6 days under equal levels (90 μmol s^-1 m^-2) of photosynthetic photon flux density (PPFD) provided by cool white fluorescent (CWF) or incandescent (INC) lamps and equal levels of far-red (28 W m^-2, 700–800 nm) radiation provided by the same INC or far-red (FR) fluorescent lamps. Brassinosteroid treatment had no appreciable effect on total biomass production but caused a decrease of 15–20% dry matter distribution in the upper portion of the shoot, a small (4%) but constant increase in dry matter in l₂ and a large (11–16%) increase in dry matter in the lower portion of the shoot (especially I₁). Treatment with BR increased assimilate accumulation in the primary leaves, especially under INC and FR lamps, and reduced dry matter in the trifoliate leaves. BR also caused a 16–21% reduction in total leaf area and even a greater reduction in area of the trifoliate leaves, but significantly increased specific leaf weight of the primary leaves and the first trifoliate leaf and the amount of dry matter in the lateral shoots under all radiation sources. In comparison to controls, BR treatment increased dry matter accumulation in the treated internode 3.3x under CWF and 1.6x under INC or FR. BR treatment also increased chlorophyll content in the primary leaves under all radiation sources and in the trifoliate leaves under CWF and INC lamps. These findings suggest a possible mobilization role of BR and establish the importance of adequate PPFD (and photosynthate) for maximum swelling and splitting response to brassinosteroid.     

Metabolic responses to red/far-red ratio and ontogeny show poor correlation with the growth rate of sunflower stems

In sparse canopies, low red to far-red (R/FR) ratios reach only vertically-oriented stems, which respond with faster rates of extension. It is shown here that this signal also promotes stem dry matter accumulation in sunflower (Helianthus annuus) but not in mustard (Sinapis alba L.). Physically blocking internode extension growth also blocked internode recovery of labelled carbon fed to the leaves, indicating that increased carbon accumulation is partially a consequence of increased extension growth in sunflower. However, low R/FR also promoted carbon accumulation in the lower section of the internode, where extension growth was unaffected. Although the levels of many soluble metabolites and of cell-wall carbohydrates increased in the stem in response to low R/FR, allowing conservation of their concentration, sucrose was present at a lower concentration under low R/FR. This change is anticipated to favour carbon unloading from the stem phloem. Low R/FR also reduced the levels of selected fatty acids, fatty acid alcohols, and sterols. Compared with the lower section, the upper section of the internode showed higher levels of organic acids, amino acids, fatty acids, and sterols. It is concluded that the promotion of stem extension growth by low R/FR ratios causes increased dry matter gain in sunflower internodes by a mechanism that is largely independent of changes in metabolism, since, whilst both low R/FR and ontogeny alter the metabolic profile, the changes do not correlate with the observed growth responses.     

Modelling photo-modulated internode elongation in growing glasshouse cucumber canopies

? Growing glasshouse plant canopies are exposed to natural fluctuations in light quantity, and the dynamically changing canopy architecture induces local variations in light quality. This modelling study aimed to analyse the importance of both light signals for an accurate prediction of individual internode lengths. ? We conceptualized two model approaches for estimating final internode lengths (FILs). The first one is only photosynthetically active radiation (PAR)-sensitive and ignores canopy architecture, whereas the second approach uses a functional-structural growth model for considering variations in both PAR and red : far-red (R : FR) ratio (L-Cucumber). Internode lengths measured in three experiments were used for model parameterization and evaluation. ? The overall trends for the simulated FILs using the exclusively PAR-sensitive model approach were already in line with the measured FILs, but they underestimated FILs at higher ranks. L-Cucumber provided considerably better FIL predictions under various light conditions and canopy architectures. ? Both light signals are needed for an accurate estimation of the FILs, and only L-Cucumber is able to consider R : FR signals from the growing canopy. Yet this study highlights the significance of the PAR signal for predicting FILs as neighbour effects increase, which indicates a potential role of photosynthate signalling in internode elongation.     

Photoresponses of Light-Grown phyA Mutants of Arabidopsis (Phytochrome A Is Required for the Perception of Daylength Extensions)

Several aspects of the photophysiology of wild-type Arabidopsis thaliana seedlings were compared with those of a phytochrome A null mutant, phyA-1, and a mutant, fhy1, that is putatively involved in the transduction of light signals from phytochrome A. Although phyA seedlings display a near wild-type phenotype when grown in white light (W), they nevertheless display several photomorphogenic abnormalities. Thus, whereas the germination of wild-type and fhy1 seeds is almost fully promoted by a pulse of red light (R) or by continuous far-red light (FR), phyA seed germination is responsive only to R. Following growth under day/night cycles, but not under continuous W, the hypocotyls of light-grown phyA and fhy1 seedlings are more elongated than those of wild-type seedlings. For seedlings grown under low red/far-red (R/FR) ratio light conditions, phyA and fhy1 seedlings display a more marked promotion of hypocotyl elongation than wild-type seedlings. Similarly, seedlings that are doubly null for phytochrome A and phytochrome B(phyA phyB) also have more elongated hypocotyls under low R/FR ratio conditions than phyB seedlings. This indicates that phytochrome A action in light-grown seedlings is antagonistic to the action of phytochrome B. Although wild-type, fhy1, and phyA seedlings flower at essentially the same time under both short-day and long-day conditions, an obvious consequence of phytochrome A deficiency is a pronounced late flowering under conditions where a short day of 8 h of fluorescent W is extended by 8 h of low-fluence-rate incandescent light. The evidence thus indicates that phytochrome A plays a role in seed germination, in the control of elongation growth of light-grown seedlings, and in the perception of daylength.     

Shoot/root assimilate allocation and nodulation ofVigna unguiculata seedlings as influenced by shoot light environment

Spectral balance of light received by southern pea [Vigna unguiculata (L.) Walp.] seedling shoots affected photoassimilate allocation among leaves, stems and roots. A higher ratio of far-red (FR) relative to red (R) light resulted in longer stems, higher shoot/root biomass ratio, less massive roots and fewer nodules. The same response pattern to FR/R ratio was obtained in a controlled environment with artificial light sources, or in sunlight where the FR/R ratio was modified by reflection from different colored soil surfaces or by FR reflected from competing plants. The importance of early shoot/root photoassimilate allocation and nodulation may differ according to soil nitrogen availability and moisture content.     

Control of Growth and Reproduction in Cultivated and Wild Rice by Light Quality and Dark Period

One cultivated and two wild rice varieties have been subjectedto variation in photoperiod and light quality by daily exposureof the seedlings at the four-leaf stage to 8 h of natural daylightfollowed by white incandescent, red, green or blue light for2,4 or 8 h in a temperature and humidity-controlled growth chamber.In some cases far-red irradiation was applied after white orred for 1 and 2 h. The treatments caused marked differencesin growth and reproduction between the cultivated and wild rices.The cultivar Dudkalmi showed extensive tillering after far-redexposure. Earliest flowering was observed with a 16-h dark periodboth in the cultivated and wild rices. Failure of floweringwith and 8-h day and 8-h artificial light of different wavelengthscould be overcome by red or far-red of 1-h duration. The lightquality interacted differently with the dark period in the accelerationof flowering in the three varieties. In another experiment theeffects of interruption of the dark period by a light periodof 2 h after from 4–12 h of darkness in a 24-h cycle werestudied in the two wild rice varieties. Light of different wavelengthsinterposed in the dark period caused variation in tiller numberand stem length in comparison to an uninterrupted dark periodof 16 h. The effect at the beginning of the dark period wasearlier flowering; flowering was delayed by interruption at4 h and inhibited after 8 h but accelerated after a 10- to 12-hdark period. The results are discussed in the light of the significanceof the dark period and light quality in regulating hormone balanceand phytochrome reactions.     

Photomodulation of growth and specific changes in fatty acid amounts in internodes of green Vigna sinensis L.

First internode growth of green Vigna sinensis L. can be widely modified by light or dark treatments. In all the treatments used there is a good correlation between the internode growth and the rate of C_18-1 accumulation. None of the other fatty acids show such a correlation.Abbreviations C_16-0 palmitoic acid - C₁₇ heptadecanoic acid - C_18-0 stearic acid - C_18-1 octadecenoic acid - C_18-2 linoleic acid - C_18-3 linolenic acid - D darkness - DW dry weight - FR far-red light - FW fresh weight - P_FR phytochrome in the FR absorbing form - R red light - W white light     

Photomorphogenic and Chlorophyll Studies in the Bryophyte Marchantia polymorpha

Besides the standard rod (R) and far-red (FK) irradiations, a graded series of different R/FR ratios were tested as 10 min terminal exposures at the end of the daily 8-hour photoperiod of white fluorescent light. Water filtered incandescent light of 3780 lux during 10 min caused A rather weak hut reproducible effect. A superposition to the water layer of different filter combinations shifting the initial transmittance more towards the FR region, and thus gradually lowering the R/FB ratio, resulted in a parallel increase in orthotropic growth and a decrease in chlorophyll content. Our data show growth similarities with the results of other authors on light grown seedlings of higher plants. Rather high levels of the P_FR form of phytochrome seem to he required to maintain horizontal growth and optimal chlorophyll content in Marchantia thalli.     

Interaction between the light quality and flowering time pathways in Arabidopsis

Flowering in Arabidopsis is accelerated by a reduced ratio of red light to far-red light (R/FR), which indicates the proximity of competitive vegetation. By exploiting the natural genetic variation in flowering time responses to low R/FR, we obtained further insight into the complex pathways that fine-tune the transition to flowering in Arabidopsis. The Bla-6 ecotype does not flower significantly earlier in response to low R/FR, but is still able to display other features of shade avoidance, suggesting branching of low R/FR signalling. Here we show that the muted flowering response of Bla-6 is due to high levels of the floral repressor FLOWERING LOCUS C (FLC), conferred by a combination of functional FLC and FRIGIDA ( FRI ) alleles with a 'weak' FY allele. The Bla-6 FY allele encodes a protein with a corrupted WW binding domain, and we provide evidence that this locus plays a key role in the natural variation in light quality-induced flowering in Arabidopsis. In Bla-6, FLC blocks promotion to flowering by reduced R/FR by inhibiting expression of the floral integrator FLOWERING LOCUS T ( FT ) in a dose-dependent manner. Reduction of FLC removes this obstruction, and Bla6 plants then exhibit strong induction of FT and flower early in response to a low R/FR signal. This paper illustrates the intricate interaction of environmental signals and genetic factors to regulate flowering in Arabidopsis.