Vascular tissue in the stem and roots of woody plants can conduct light

The role of vascular tissue in conducting light was analysed in 21 species of woody plants. Vessels, fibres (both xylem and phloem fibres) and tracheids in woody plants are shown to conduct light efficiently along the axial direction of both stems and roots, via their lumina (vessels) or cell walls (fibres and tracheids). Other components, such as sieve tubes and parenchyma cells, are not efficient axial light conductors. Investigation of the spectral properties of the conducted light indicated that far-red light was conducted most efficiently by vascular tissue. Light gradients in the axial direction were also investigated and revealed that conducted light leaked out of the light-conducting structures to the surrounding living tissues. These properties of the conducted light suggest a close relationship with metabolic activities mediated by phytochromes. The results therefore indicate not only that signals from the external light environment can enter the interior of stems above ground and are conducted by vascular tissue towards roots under ground, but also that the light conducted probably contributes directly to photomorphogenic activities within them.     

Photosynthate partitioning and nodule formation in soybean plants that received red or far-red light at the end of the photosynthetic period

The influence of row orientation on spectral distribution of light received by growing soybean [ Glycine max (L.) Merr. (cv. Coker 338)] plants was measured under field conditions, and light spectrum effects on photosynthate partitioning were studied under controlled environments. Light received by leaves under field conditions differed among those grown in north-south vs east-west oriented rows. In morning and late afternoon, the far-red/ red ratio received by leaves at the surface of the canopy differed about 3-fold from the east to west sides of north-south rows, but only 1.3-fold from the south to north sides of east-west rows.
In controlled environments, brief exposures to red or far-red light at the end of the photosynthetic period influenced partitioning of photosynthate among leaves, stems and roots. The top/root ratios differed significantly between the red and far-red treated plants. Red treated plants partitioned less photosynthate to stems and more to roots than did those treated with far-red. Also, plants with larger root systems developed more nodules. Phytochrome effects on photosynthate partitioning between tops and roots may influence yield of soybean plants grown in soils with low water-holding capacity.     

The nodulation and nitrogen fixation of isolated roots of Phaseolus vulgaris L.

Summary The exposure of isolated bean roots to white light immediately before inoculation with nodule bacteria stimulated nodulation during a subsequent 7-day dark period. Strong white light applied to inoculated roots for 3 days or longer suppressed nodulation except when the light was applied immediately after inoculation and the duration of the light period did not exceed 3 days. Alternating periods of 12 hours light and 12 hours darkness were almost as inhibitory to nodulation as a continuous exposure to light. A combination of blue and far-red light inhibited nodule formation more than a combination of red and far-red light. Light treatments that inhibited nodulation also suppressed the production of lateral roots, but to a lesser extent. The effect of light on nodulation and lateral root formation was not influenced by the presence or absence of an attached hypocotyl segment in the explant.     

Light-mediated bioregulation of tillering and photosynthate partitioning in wheat

The influence of plant population density on spectral distribution of light received by wheat ( Triticum aestivum L. cv. Coker 797) seedlings was measured under field conditions, and effects of red and far-red light on tillering and photosynthate partitioning were studied in controlled environments. Spectral distribution of light was measured in sunflecks at soil level in close-, intermediate-, and wide-spaced field populations during the tillering stage. Close-spaced seedlings received higher far-red/red light ratios than wide-spaced plants because of the larger amount of far-red reflected from green leaves of the more numerous nearby plants. The far-red/red light ratios in all population densities were higher in late afternoon than at noon. Close-spaced plants developed fewer tillers, less roots and longer leaves than wide-spaced seedlings under field conditions. In controlled environments, a higher far-red/red ratio during photosynthetic periods resulted in fewer tillers and longer leaves; whereas, brief red or far-red exposures at the end of each day had a more pronounced effect. Wheat seedlings that received 5-min exposures to far-red light at the end of the photosynthetic period each day for 20 consecutive days developed fewer tillers, longer leaves, less roots, and a higher shoot/root biomass ratio. The effects of far-red light were reversed by red light. The light spectral shifts associated with field plant population densities and the responses to red and far-red treatments under controlled environments suggest that phytochrome serves as a sensing mechanism that detects the amount of competition from other plants, and regulates the development of tillers and the partitioning of photosynthate between shoots and roots.     

Anatomical features of pepper plants (Capsicum annuum L.) grown under red light-emitting diodes supplemented with blue or far-red light

Pepper plants (Capsicum annuum L. cv., Hungarian Wax) were grown under metal halide (MH) lamps or light-emitting diode (LED) arrays with different spectra to determine the effects of light quality on plant anatomy of leaves and stems. One LED (660) array supplied 90% red light at 660 nm (25nm band-width at half-peak height) and 1% far-red light between 700-800nm. A second LED (660/735) array supplied 83% red light at 660nm and 17% far-red light at 735nm (25nm band-width at half-peak height). A third LED (660/blue) array supplied 98% red light at 660nm, 1% blue light between 350-550nm, and 1% far-red light between 700-800nm. Control plants were grown under broad spectrum metal halide lamps. Plants were gron at a mean photon flux (300-800nm) of 330 micromol m-2 s-1 under a 12 h day-night photoperiod. Significant anatomical changes in stem and leaf morphologies were observed in plants grown under the LED arrays compared to plants grown under the broad-spectrum MH lamp. Cross-sectional areas of pepper stems, thickness of secondary xylem, numbers of intraxylary phloem bundles in the periphery of stem pith tissues, leaf thickness, numbers of choloplasts per palisade mesophyll cell, and thickness of palisade and spongy mesophyll tissues were greatest in peppers grown under MH lamps, intermediate in plants grown under the 660/blue LED array, and lowest in peppers grown under the 660 or 660/735 LED arrays. Most anatomical features of pepper stems and leaves were similar among plants grown under 660 or 660/735 LED arrays. The effects of spectral quality on anatomical changes in stem and leaf tissues of peppers generally correlate to the amount of blue light present in the primary light source.     

Functional relationship between chlorophyll content and leaf reflectance, and light-capturing efficiency of Japanese forest species

We examined the functional relationship between chlorophyll concentrations and light spectral absorption in 16 species of woody, vine and herbaceous plants in northern Japan. Leaves of each species from under forest shade and in more open sites were measured for chlorophyll, specific leaf area (SLA) and spectral absorption. In all species, SLA increased and the Chl a : b ratio declined in shade- vs open-grown leaves indicating an adaptive adjustment to forest shade in these leaf characters. However, the expected increase in the ratio of 680 to 700 nm absorption in shade leaves did not occur in all species. Light absorption at 680 relative to 700 nm was lower in the shade leaves of Acer japonicum. Kalopanax pictus, Panax japonicus and Petasites japonicus even with a reduced Chl a : b , a commonly accepted indicator of shade adaptation. Therefore, spectral measurements in these species failed to support Chl concentrations that were expected to confer an improvement in the absorption of red light (<680nm) deficient relative to far-red light (>700 nm) in the forest shade. Compared with other species, the absorption pattern of these four 'non-conforming' species is associated with a higher ratio of shade:open leaves in reflectance spectra in the 600–750 nm range. This suggests an increased reflectance in shade leaves caused by changes in leaf surface properties which are not immediately apparent. We conclude that adaptive spectral absorption cannot always be inferred from changes in specific leaf area and chlorophyll a and b concentrations.     

Non-photosynthetic effects of red and far-red light on root-nodule formation by leguminous plants

Summary Nodulation of pea and broad bean plants grown in the light was found to be reduced when the roots were exposed to far-red light for 5–15 minutes daily during 5 consecutive days following inoculation with nodule bacteria. Similar results were obtained following a single exposure to far-red light during a period of 15 minutes at the 3rd or 4th day after inoculation. When the roots were exposed to far-red light either before inoculation or during the first two days afterwards there were either no effects or only slight effects on nodulation The inhibitory effect of far-red light on nodulation was partly reduced by subsequent exposure to red light, provided that the same part of the plant was exposed to both red and far-red light,viz either the root or the shoot. When different parts of the plant were exposed to red and far-red light respectively, there was no interaction between the two kinds of light on nodulation. Plants whose roots were exposed to far-red light did not subsequently show stem elongation.Nodules were found to develop on the roots of pea plants grown in the dark, provided that the plants were kept at or below 22°C. At 25°C nodulation was almost absent. Nodulation was decreased by addition of kinetin and IAA. In contrast to plants grown in the light pea plants grown in the dark, inoculated with either an effective or ineffective strain of Rhizobium, developed equal numbers of nodules. Exposure to red light slightly increased the percentage of nodulated plants but decreased the number of nodules per plant. Exposure to far-red light slightly decreased both the percentage of nodulated plants and the number of nodules per plant. The effect of far-red light was counteracted by red light andvice versa.     

<Emphasis Type="Italic">In vitro</Emphasis> culture of <Emphasis Type="Italic">Chrysanthemum</Emphasis> plantlets using light-emitting diodes

Effects of illumination spectrum on the morphogenesis of chrysanthemum plantlets (Chrysanthemum morifolium Ramat. ‘Ellen’) grown in vitro were studied using an illumination system consisting of four groups of light-emitting diodes (LEDs) in the following spectral regions: blue (450nm), red (640nm), red (660nm), and far-red (735nm). Taking into account all differences in shoot height, root length, and fresh and dry weight (FW and DW, respectively), observed while changing the total photon flux density (PFD), the optimal total PFD for growth of chrysanthemum plantlets in vitro was estimated. For 16 h photoperiod and typical fractions of the spectral components (14%, 50%, 28%, and 8%, respectively), the optimal total PFD was found to be 40 μmol m⁻² s⁻¹. Our study shows that the blue component in the illumination spectrum inhibits the plantlet extension and formation of roots and simultaneously increases the DW to FW ratio and content of photosynthetic pigments. We demonstrate photomorphogenetic effects in the blue region and its interaction with the fractional PFD of the far-red spectral component. Under constant fractional PFD of the blue component, the root number, length of roots and stems, and fresh weight of the plantlets have a correlated nonmonotonous dependence on the fractional PFD of the far-red component.     

Green light signaling and adaptive response

To a plant, the sun’s light is not exclusively energy for photosynthesis, it also provides a package of data about time and prevailing conditions. The plant’s surroundings may dampen or filter solar energies, altering spectral profiles of their light environment. Plants use this information to adjust form and physiology, tailoring gene expression to best match ambient conditions. Extensive literature exists on how blue, red and far-red light contribute to plant adaptive responses. A growing body of work identifies discrete effects of green light (500–565 nm) that also shape plant biology. Green light responses are known to be either mediated through, or independent of, the cryptochrome blue light receptors. Responses to green light share a general tendency to oppose blue- or red-light-induced responses, including stem growth rate inhibition, anthocyanin accumulation or chloroplast gene expression. Recent evidence demonstrates a role for green light in sensing a shaded environment, independent from far-red shade responses.     

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

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

The characteristics of light in floral induction in vitro of Cichorium intybus. The possible role of phytochrome

The action of light in the initiation of floral buds in vitro has been studied using monochromatic light qualities on root explants of a long day plant, Cichorium intybus L. cv. Witloof. Red light (660 nm, 0.30 W m^-2) promotes flowering, while far-red (730 nm, 0.31 W m^-2) and irradiation with combined red + far-red (0.20 + 0.41 W m^-2) have no effect. In short day conditions floral response can be obtained in two ways: 1) by interrupting the dark period with 5 brief irradiations of red light (0.45 W m^-2, 12 min) at regular intervals, although these are counteracted by far-red irradiations of equal intensity and duration; 2) by interrupting the long night with 5 h red light applied during the second third of the night, while at the beginning or at the end it is ineffective. Red light efficiency appears to depend on the photosynthetic activity of the tissues, so that flowering increases with increasing intensity of white light and is suppressed if no white light is supplied. The reproductive development is determined by the coordination of proper irradiation conditions with sufficient sensitivity of the perceiving meristematic cells. The period of highest sensitivity to environmental light conditions in the life cycle of a Cichorium root explant occurs between the 8th and the 16th day after the start of the culture. The data strongly suggest that phytochrome is involved in flower induction of Cichorium in vitro.     

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.     

Accumulation and fractionation of rare earth elements (REEs) in the naturally grown Phytolacca americana L. in southern China

The widespread use of rare earth elements (REEs) has resulted in problems for soil and human health. Phytolacca americana L. is a herbaceous plant widely distributed in Dingnan county of Jiangxi province, China, which is a REE mining region (ion absorption rare earth mine) and the soil has high levels of REEs. An investigation of REE content of P. americana growing naturally in Dingnan county was conducted. REE concentrations in the roots, stems, and leaves of P. americana and in their rhizospheric soils were determined. Results showed that plant REEs concentrations varied among the sampling sites and can reach 1040 mg/kg in the leaves. Plant REEs concentrations decreased in the order of leaf > root > stem and all tissues were characterized by a light REE enrichment and a heavy REE depletion. However, P. americana exhibited preferential accumulation of light REEs during the absorption process (from soil to root) and preferential accumulation of heavy REEs during the translocation process (from stem to leaf). The ability of P. americana to accumulate high REEs in the shoot makes it a potential candidate for understanding the absorption mechanisms of REEs and for the phytoremediation of REEs contaminated soil.     

A phytochrome-mediated alteration from stem to rosette growth on chicory root explants in vitro

Chicory root explants (Cichorium intybus L. var. foliosum) of two cultivars, taken before and after hydroponic forcing, were cultured in vitro in complete darkness supplemented with red and far-red light treatments. Using 5 min red light per day, the strong stem elongation occurring in complete darkness was converted to rosette formation. This reaction was reversed to stem elongation (accompanied by leaf formation) adding 15 min far-red light after the red light. Fifteen min far-red light per day alone caused the same reaction as 5 min red/15 min far-red light. Far-red light followed by red light caused rosette formation. In stems, formed under complete darkness in vitro, the presence of phytochrome was shown. No phytochrome was detected in the root fragment itself.Abbreviations R red light - FR far-red light - GA gibberellinic acid - A absorbance - FW fresh weight     

Spatial-specific regulation of root development by phytochromes in Arabidopsis thaliana

Distinct tissues and organs of plants exhibit dissimilar responses to light exposure – cotyledon growth is promoted by light, whereas hypocotyl growth is inhibited by light. Light can have different impacts on root development, including impacting root elongation, morphology, lateral root proliferation and root tropisms. In many cases, light inhibits root elongation. There has been much attention given to whether roots themselves are the sites of photoperception for light that impacts light-dependent growth and development of roots. A number of approaches including photoreceptor localization in planta, localized irradiation and exposure of dissected roots to light have been used to explore the site(s) of light perception for the photoregulation of root development. Such approaches have led to the observation that photoreceptors are localized to roots in many plant species, and that roots are capable of light absorption that can alter morphology and/or gene expression. Our recent results show that localized depletion of phytochrome photoreceptors in Arabidopsis thaliana disrupts root development and root responsiveness to the plant hormone jasmonic acid. Thus, root-localized light perception appears central to organ-specific, photoregulation of growth and development in roots.     

Measurement of light gradients and spectral regime in plant tissue with a fiber optic probe

A method is described in which light gradients and spectral regime can be measured within plant tissue using fiber optics. A fiber optic probe was made by modifying a single optical fiber (200 μm diameter) so that it had a light harvesting end that was a truncated tip 20–70 μm in diameter. The probe was a directional sensor with a half-band acceptance angle of 17–20°. Light measurements were made as the fiber optic probe was driven through plant tissue by a motorized micromanipulator, and the light that entered the fiber tip was piped to a spectroradiometer. By irradiating green leaf tissue of the succulent Crassula falcata L. with collimated light and inserting the probe from different directions, it was possible to measure light quality and quantity at different depths. Collimated light was scattered completely by the initial 1.0 mm of leaf tissue, which also greatly attenuated all light except the green and far-red. Light scatter contributed significantly to light quantity and had a pronounced spectral structure. Immediately beneath the irradiated surface the amount of light at 550 nm was 1.2 times that of the incident light. The light gradient declined rapidly to 0.5 times incident light at 1.4 mm depth. In contrast, the amount of light at 750 nm increased during the initial 0.5 mm to 2.9 times incident light and then declined linearly to 0.5 times incident light at the dark side of the leaf (4.5 mm). The implications of the magnitude of the contribution of light scatter to the light gradient is also discussed.     

Two homologous ATP-binding cassette transporter proteins, AtMDR1 and AtPGP1, regulate Arabidopsis photomorphogenesis and root development by mediating polar auxin transport

Light and auxin control many aspects of plant growth and development in an overlapping manner. We report here functional characterization of two closely related ABC (ATP-binding cassette) transporter genes, AtMDR1 and AtPGP1, in light and auxin responses. We showed that loss-of-function atmdr1 and atpgp1 mutants display hypersensitivity to far-red, red, and blue-light inhibition of hypocotyl elongation, reduced chlorophyll and anthocyanin accumulation, and abnormal expression of several light-responsive genes, including CAB3, RBCS, CHS, and PORA, under both darkness and far-red light conditions. In addition, we showed that the atmdr1-100 and atmdr1-100/atpgp1-100 mutants are defective in multiple aspects of root development, including increased root-growth sensitivity to 1-naphthalene acetic acid (1-NAA), and decreased sensitivity to naphthylphthalamic acid (NPA)-mediated inhibition of root elongation. Consistent with the proposed role of AtMDR1 in basipetal auxin transport, we found that expression of the auxin responsive DR5::GUS reporter gene in the central elongation zone is significantly reduced in the atmdr1-100 mutant roots treated with 1-NAA at the root tips, compared to similarly treated wild-type plants. Moreover, atmdr1-100, atpgp1-100, and their double mutants produced fewer lateral roots, in the presence or absence of 1-NAA or NPA. The atmdr1-100 and atmdr1-100/atpgp1-100 mutants also displayed enhanced root gravitropism. Genetic-epistasis analysis revealed that mutations in phyA largely suppress the randomized-hypocotyl growth and the short-hypocotyl phenotype of the atmdr1-100 mutants under far-red light, suggesting that phyA acts downstream of AtMDR1. Together, our results suggest that AtMDR1 and AtPGP1 regulate Arabidopsis (Arabidopsis thaliana) photomorphogenesis and multiple aspects of root development by mediating polar auxin transport.     

Phytochrome Controlled Adhesion of Mung Bean Root Tips to Glass: A Detailed characterization of the Phenomenon

Various parameters of the Tanada effect (Proc. Natl. Acad. Sci. U.S. 59: 376–380. 1968) have been defined. This phenomenon, in which root tips of Phaseolus aureus L. adhere to a negatively charged glass surface when they are irradiated with 660 nm (red) light and release under 730 nm (far-red) light, has been characterized as follows. Secondary roots, whether etiolated or light grown exhibit photoreversible adhesion. Primary roots do not. Tips from 6–8 mm secondary roots exhibit the best response to red light, whereas tips from 3 mm roots respond best to far-red light. Red light saturetes the adhesion system at about 50 μ W/cm²xnm and far-red light, release system at about 150 ü W/cm² xnm. The adhesion effect begins to show escape from far-red reversibility within 60–90 seconds, an observation quite different from other “typical” long term de- etiolation effects. In addition, root tips irradiated with red light begin to release spontaneously in the dark after 10 min, and have nearly completed release after 50 min. Tips irradiated with continuous red light show gradual release after 15 minutes of exposure. Whether these data indicate an extremely rapid dark reversion of P_fr to P_r or decay of P_fr under continuous red light is not known at this time. In order to study tip adhesion and release, the glass beaker surface may be negatively charged with thiocyanate (SCN^-), nitrate (NO₃^-), sulfate (SO₄^2-), chloride (Cl^-), phosphate (PO₄^3-), citrate (C₆H₅O₇^3-), oxalate (C₂O₄^2-) or glutamine (C₅H₈NO₄^-). Benzoate (C₇H₅O₂^-) and acetate (CH₃COO^-) were found to be relatively ineffective for red light adhesion, however when citrate and oxalate were used release was inhibited. This was apparently due to a chelation of Ca²⁺since release began immediately as excess Ca⁺² was added to the bathing solution. Substitution of GTP, ITP, UTP, or CTP for ATP resulted in only 20 to 40% adhesion and release for GTP, ITP and UTP, CTP showed normal adhesion kinetics under red light but very slow release kinetics under far-red light. The effects of red and far-red light in the numbers of secondary roots are that red light inhibits root initiation while far-red light partially reverses the red light effect.     

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.