Reflected Far-Red Light Effects on Chlorophyll and Light-Harvesting Chlorophyll Protein (LHC-II) Contents under Field Conditions

The influence of various colors of soil cover (mulch) on the farred/red (FR/R) ratio in upwardly reflected light and on concentrations of chlorophyll (Chl) and light-harvesting Chl protein (LHC-II) were measured under field conditions. The FR/R ratios above green surfaces were higher than over white surfaces. Even though plants (Gossyplum hirsutum L. cv PD-1) were grown in full sunlight, those that received higher FR/R ratios in upwardly reflected light were taller and had thinner leaves with higher concentrations of Chl and LHC-II. A controlled environment experiment showed FR/R control of Chl and LHC-II concentrations. The results illustrate the importance of spectral distribution of reflected light on plant growth and a potential means of altering the chemistry of leaf crops under field conditions.     

Effect of Mulch Surface Color on Root-knot of Tomato Grown in Simulated Planting Beds

The effect of different-colored polyethylene mulches on quantity and spectra of reflected light, plant morphology, and root-knot disease was studied in tomato (Lycopersicon esculentum) grown in simulated planting beds. Tomato plants were inoculated with Meloidogyne incognita at initial populations (Pi) of 0, 1,000, 10,000, or 50,000 eggs/plant, and grown in a greenhouse for 50 days over white, red, or black mulch. Soil temperature was kept constant among the mulch treatments by placing an insulation barrier between the colored mulch and the soil surface. Soil temperature varied less than 0.5 °C between soil chambers at solar noon. Tomatoes grown over white mulch received more reflected photosynthetic light and had greater shoot weights (27%), root weights (32%), and leaf area (20%) than plants grown over black mulch. Plants grown over red mulch received a higher far-red-to-red ratio in the reflected light. Mulch color altered the plant''s response to root-knot nematode infection by changing the distribution of mass in axillary shoots. At high Pi, axillary leaf area and leaf weight were greater in tomato grown over white mulch than when grown over red mulch. The root-gall index was lower for plants grown over white mulch than similar plants grown over red mulch.     

Early detection of neighbour plants by phytochrome perception of spectral changes in reflected sunlight

Abstract We have tested the hypothesis that a plant may detect the presence of a neighboug42r, before being shaded by it, through the perception of the spectral composition of reflected sunlight. Within seedling canopies the red: far-red ratio (R: FR) of the light received by a sensor with a geometry approximating that of a stem was significantly reduced by selective reflection. This effect was observed before any reduction in the amount of photosynthetic light energy received by an individual seedling could be detected. Small green fences of grass, east-west orientated, altered the spectral distribution of the light on the north (sunlit) side of them. Fully illuminated seedlings of Sinapis alba grown on the north side of these green fences produced longer internodes and had a lower leaf: stem dry weight ratio than those grown in front of fences of bleached grasses. A similar redistribution of growth was elicited in seedlings of Chenopodium album, Datura ferox and S. alba growing in full sunlight by exposing plants to additional small quantities of far-red reflected by selective mirrors. These results suggest that the change in the R: FR ratio serves as an early warning signal of oncoming competition.     

Colored Mulches Affect Yield of Fresh-market Tomato Infected with Meloidogyne incognita

The effects of different-colored polyethylene mulches on the quantity and spectra of reflected light, earliness of fruit set, fruit yield and quality, and root-knot disease were studied in field-grown, staked tomato (Lycopersicon esculentum). White mulch reflected more photosynthetic light and a lower far-red-to-red ratio than red mulch, whereas black mulch reflected less than 5 percent of any color. Soil temperatures and fruit yields were recorded for tomato plants inoculated with Meloidogyne incognita race 3 at initial populations of 0, 1,000, 10,000, 50,000, or 100,000 eggs/plant and grown over black, white, or red plastic mulch in both spring and fall. Soil temperatures were lower under white mulch than under red or black mulch. Tomato yields declined as inoculum level increased. Plants grown over red mulch in the spring and inoculated with 50,000 eggs of M. incognita had greater early marketable yields than similarly inoculated plants grown over black or white mulch. Tomato plants inoculated with 100,000 eggs and grown over white mulch or red mulch in the spring had greater total yields per plot than similar plants grown over black mulch (7.39 kg and 7.71 kg vs. 3.65 kg, respectively).     

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.     

Row orientation effects on FR/R light ratio, growth and development of field-grown bush bean

Bush bean ( Phaseolus vulgaris L. cv. Tenderette) plants were grown in north-south (N-S) and east-west (E-W) rows under field conditions to test effects of row orientation on reflected far-red (FR) light patterns and on shoot size and edible bean productivity. Soil water and nutrients were adequate. Plants in N-S rows received slightly higher ratios of FR relative to red (R) light, because of heliotropic movement of the leaves. Plants in N-S rows partitioned more dry matter to shoots and edible green beans than those in E-W rows. We conclude that row orientation of broadleaf plants can affect the FR/R light ratio and the phytochrome-mediated regulation of plant development under field conditions.     

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.     

Overexpression of oat phytochrome A gene differentially affects stem growth responses to red/far-red ratio signals characteristic of sparse or dense canopies

The effects of overexpression of oat phytochrome A on neighbour detection and on stem-growth responses to changes in red light (R), far-red light (FR) and blue light (B) simulating neighbours were investigated in transgenic tobacco seedlings grown under natural radiation. In wild-type (WT) seedlings, stem extension growth was promoted: (1) by lowering the R:FR by means of daytime supplementary FR, end-of-day FR, neighbours reflecting FR, or selective light filters placed around the base of the shoot to reduce R without affecting FR; and (2) by lowering phytochrome-absorbable radiation (R+FR) reaching the stem. Transgenic seedlings only responded to reductions in R:FR involving no significant changes in FR irra-diance, i.e. end-of-day FR and filters placed around the stem to reduce R. Neither daytime supplementary R nor selective filters placed around the stem to reduce B affected stem growth in any genotype. In growing canopies, WT seedlings responded to the reduction of R:FR caused by FR reflected in neighbour plants. Transgenic seedlings responded to plant density about a week later, when mutual plant shading reduced R and (to a lesser extent) FR below sunlight levels. Overexpression of phytochrome A impaired early neighbour detection.     

Phytochrome and internode elongation in Chenopodium polyspermum L. the light fluence rate during the day and the end-of-day effect

The elongation of the fourth internode of fully green Chenopodium polyspermum L. is strongly stimulated by far-red light (FR) given at the end of the day. The end-of-day effect is more important when the plants had been cultivated for several days with a main light period of 140 Wm^-2 than with a main light period of 85 Wm^-2. There exists a quantitative relationship between the FR end-of-day effect mediated by phytochrome and the value of the light fluence during the day.Abbreviations D darkness - FR far-red light - HWL white light at 140 Wm^-2 - LWL white light at 85 Wm^-2 - PAR photosynthetically active radiation - R red light - WL white light     

Phytochrome photoreceptors mediate plasticity to light quality in flowers of the Brassicaceae

The family of phytochrome photoreceptors mediates stem-elongation responses to ambient ratios of red?:?far-red light (R?:?FR). Although phytochrome genes are expressed in flowers in addition to vegetative parts, nothing is known about floral plasticity to R?:?FR or the pleiotropic effects of phytochrome genes on flowers. Here, the following floral morphologies were compared: (1) wild-type Arabidopsis thaliana and Brassica rapa plants experiencing high R?:?FR characteristic of sunlight vs. low R?:?FR typical of foliar shade and (2) wild-type and phytochrome-deficient A. thaliana plants. Wild-type A. thaliana exposed to low R?:?FR had diminished petal and pistil lengths but longer filaments for a given petal size than plants experiencing high R?:?FR. Brassica rapa plants had qualitatively similar responses. In comparison to wild-type A. thaliana, mutants lacking phytochrome A had smaller flowers (smaller petals, pistils, and filaments), whereas phytochrome B-deficient mutants exhibited longer filament lengths. These results provide the first evidence that R?:?FR and phytochromes affect floral phenotypes in addition to vegetative ones. Although the ecological relevance remains to be established, the observed plasticity of flowers to R?:?FR may be relevant to individual fitness in some species because stigma and filament positions can affect pollen removal and levels of self-pollination.     

Effect of light quality and 5-azacytidine on genomic methylation and stem elongation in two ecotypes of Stellaria longipes

Changes in cytosine methylation are known to occur in response to various environmental stimuli, therefore, we looked at methylation changes in relation to stem elongation. More specifically, we investigated the response of genomic cytosine methylation to irradiance-mediated plasticity of stem elongation in two ecotypes of Stellaria longipes . Ramets of S. longipes were grown under high and low ratios of red/far-red light (F/FR; 3.7 and 0.7, respectively). Stem elongation and methylated cytosine content were measured over a period of 7 days. Ramets of S. longipes demonstrated the highest level of demethylation after 4 days of long-day warm (LDW) treatment, which coincides with the first day of rapid stem elongation initiation. The extent of demethylation associated with day 4 depended upon the relative ratio of R/FR light. In particular, those plants treated with low R/FR light ratios showed a lower level of methylation, and were taller than the high R/FR light grown counterparts. In addition, prairie ecotype plants demonstrated lower day 4 methylation levels, as well as longer day 7 stem lengths, than the alpine ecotype plants within the same R/FR light treatments. To investigate if the degree of methylation was a crucial factor in controlling the stem elongation response, ramets of both alpine and prairie plants were grown in MS media supplemented with 5-azacytidine (5-AzaC), and grown for 14 days under a R/FR ratio of 3.7 and two different PAR values. 5-AzaC treatments demonstrated that the prairie ecotype plants required greater doses of 5-AzaC, and thus lower methylation levels, than the alpine ecotype plants in order to promote maximal stem elongation. These observations suggest that DNA demethylation is involved in the shade-avoidance response.     

Rhizome transition to storage organ is under phytochrome control in lotus <Emphasis Type="Italic">(Nelumbo nucifera)</Emphasis>

We examined photoperiodic response of lotus (Nelumbo nucifera) rhizome morphogenesis (its transition to a storage organ) by using seed-derived plants. Rhizome enlargement (increase in girth) was brought about under 8, 10 and 12 h photoperiods, whereas the rhizomes elongated under 13 and 14 h photoperiods. Rhizomes elongated under 14 h light regimes supplied as 8 h of natural light plus 6 h supplemental hours of white, yellow or red light, but similar treatments with supplemental blue, green or far red light, caused enlargement in girth of the rhizomes. A 2 h interruption of the night with white, yellow or red light, in plants entrained to 8 h photoperiod brought rhizome elongation, whereas 2 h-blue, green or far red light night breaks still resulted in rhizome increase in girth. The inhibitory effect of a red (R) light night break on rhizome increase in girth was reversed by a far-red (FR) light given immediately afterwards. Irradiation with R/FR/R inhibited the rhizome increase in girth. FR light irradiation following R/FR/R irradiation cancelled the effect of the last R light irradiation. It was demonstrated that the critical photoperiod for rhizome transition to storage organ is between 12 and 13 h photoperiod. It was also evident that the optimal light quality range for interruption of dark period (night break) is between yellow and red light and that a R/FR reversible reaction is observed. From these results, we propose that phytochrome plays an important role in photoperiodic response of rhizome increase in girth in lotus. This is the first report on phytochrome-dependent morphogenesis of storage organs in rhizomous plants.     

Morphological responses of Datura ferox L. seedlings to the presence of neighbours

Summary We studied the effects of density on the dynamics of seedling growth and canopy microclimate within experimental stands composed of Datura ferox L. seedlings grown in individual pots. Interception of photosynthetically active radiation (PAR) by seedlings was evaluated either indirectly, by measuring leaf area, proportion of leaf area shaded by neighbouring individuals and laminar orientation with respect to sunlight, or directly, by measuring PAR at individual leaves at their natural angle of display. An integrating cylinder, with a geometry approximating that of a stem, was used within the canopies to measure the red:far-red (R:FR) ratio of the light flux from all compass points parallel to the soil surface. Seedlings responded rapidly (i.e. 1–2 weeks) to increased density by producing longer internodes and partitioning more dry matter to stems relative to leaves. These responses were observed before either PAR interception of growth rate were reduced by the presence of neighbours. Conversely, morphogenetic adjustment was preceded by a consistent effect of plant density on the R:FR ratio of the light received by the integrating cylinder. Air and soil temperature were not affected by density in these experiments. Differences in wind velocity within the canopy associated with plant density were avoided by the experimental procedure. The results support the idea that the drop in R:FR ratio of the light flux parallel to the ground — e.g. reflected sunlight — is an early signal that allows rapid adjustment of plant form to changes in canopy structure.     

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.     

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     

Photoperiodism and photocontrol of stem elongation in two photomorphogenic mutants of Pisum sativum L.

The photomorphogenic mutation lv in the garden pea (Pisum sativum L.), which appears to reduce the response to light-stable phytochrome, has been isolated on a tall, late photoperiodic genetic background and its effects further characterised. Plants possessing lv have a reduced flowering response to photoperiod relative to wild-type plants, indicating that light-stable phytochrome may have a flower-inhibitory role in the flowering response of long-day plants to photoperiod. In general, lv plants are longer and have reduced leaf development relative to Lv plants. These differences are maximised under continuous light from fluorescent lamps (containing negligible far-red (FR) light), and decrease with addition of FR to the incident light. Enrichment of white light from fluorescent lamps with FR promotes stem elongation in the wild type but causes a reduction in elongation in the lv mutant. This “negative” shade-avoidance response appears to be the consequence of a strong inhibitory effect of light rich in FR, revealed in lv plants in the absence of a normal response to red (R) light. These results indicate that the wild-type response to the R: FR ratio may be comprised of two distinct photoresponses, one in which FR supplementation promotes elongation by reducing the inhibitory effect of R, and the other in which light rich in FR actively inhibits elongation. This hypothesis is discussed in relation to functional differentiation of phytochrome types in the light-grown plant. Gene lw has been reported previously to reduce internode length and the response to gibberellin A₁, and to delay flowering. The present study shows that the lw mutation confers an increased response to photoperiod. In all these responses the lw phenotype is superficially “opposite” to the lv phenotype. The possibility that the mutation might primarily affect light perception was therefore considered. The degree of dwarfing of lw plants was found to depend upon light quality and quantity. Dwarfing is more extreme in plants grown under continuous R light than in those grown in continuous FR or blue light or in darkness. Studies of the fluence-rate response show that the lw mutation imparts a lower fluence requirement for inhibition of elongation by white light from fluorescent lamps. Dark-grown lw plants are more strongly inhibited by a R pulse than are wild-type plants but, as in the wild type, this inhibition remains reversible by FR. Light-grown lw plants show an exaggerated elongation response to end-of-day FR light. Taken together, these findings indicate that the lw mutant may be hypersensitive to phytochrome action.     

Maternal and ambient environmental effects of light on germination in Plantago lanceolata: correlated responses to selection on leaf length

1. Seeds from artificial selection lines were exposed to different maternal and ambient conditions, simulating sunlight and vegetation shade.
2. Lines selected for longer leaves also produced larger seeds, indicating a positive genetic correlation between leaf length and seed size.
3. Light conditions during maturation had no large effect on seed size.
4. Seed germination was reduced by a low ratio of red to far-red light (R/FR ratio) in the ambient environment.
5. Seeds maturated under simulated vegetation shade germinated less readily and were more inhibited by a low ambient R/FR ratio than seeds maturated under full sunlight or R/FR-neutral shade. Thus, low R/FR-ratios in the maternal and ambient environment operated synergistically.
6. Large genotypic variation in the germination responses to both maternal and ambient light conditions was found among and within selection lines, indicating that such responses might have the potential to evolve in response to natural selection.
7. Artificial selection for leaf length had affected seed germination characteristics but correlated responses and thus genetic correlations largely depended on light conditions in the selective environment. Selection for longer leaves under a low R/FR ratio increased seed dormancy and plasticity of germination in response to the R/FR ratio. However, in the opposite selective environment selection for longer leaves reduced seed dormancy and plasticity to the R/FR ratio. It is argued that leaf length and seed germination characteristics are somehow linked by shared physiological mechanisms, which may facilitate concerted changes in shade avoidance responses.     

Photocontrol of stem elongation in plant neighbourhoods: effects of photon fluence rate under natural conditions of radiation

Abstract. We investigated the effects of photon fluence rate on internode elongation in fully de-etiolated plants growing under sunlight. Our goal was to find out whether perception by the stems of fluence rate changes related to canopy density may be involved in the modulation of internode growth in canopies formed by plants of similar stature (e.g. crop stands). Using Datura ferox L. and Sinapis alba L. seedlings growing under natural radiation, we found that internode elongation is promoted by localized shading. This effect was observed with internodes receiving light with a high (>0.9) or a low (0.3) red (R) to far-red (FR) photon ratio. Selective removal of the different wavebands from the light impinging on the internodes showed that part of the response to fluence rate is due to photons in the R + FR range. The blue (B) component, most likely acting through a specific photoreceptor, also inhibited elongation. However, changes in the fluence rate of B light did not have detectable effects on the response of the internodes to R:FR ratio. Fibre-optic studies and measurements with integrating-cylinder sensors in even-aged populations of seedlings showed that both the quality and quantity of radiation received by the stems are profoundly influenced by changes in canopy density. When density is very low (leaf area index = LAI ≥ 1) only the R:FR ratio is reduced, due to FR reflected from nearby leaves. In the LAI range of 1 to 2, though a large proportion of the leaf area is still receiving full sunlight, the photon fluence rate at the stem level drops dramatically. These results suggest that in even-aged populations of LAI > 1 elongation growth is promoted by the low R:FR ratio and the reduced fluence rate. Perception of these two factors at the stem level may elicit morphological adaptations in the canopy before the onset of severe competition among neighbours for the resource of light.     

Phytochrome A, phytochrome B and other phytochrome(s) regulate ATHB-2 gene expression in etiolated and green Arabidopsis plants

The expression of the Arabidopsis ATHB-2 gene is light-regulated both in seedlings and in adult plants. The gene is expressed at high levels in rapidly elongating etiolated seedlings and is down-regulated by a pulse of red light (R) through the action of a phytochrome other than phytochrome A or B, or by a pulse of far-red light (FR) through the action of phytochrome A. In green plants, the expression of the ATHB-2 gene is rapidly and strongly enhanced by lowering the R:FR ratio perceived by a phytochrome other than A or B. Returning the plant to a high R:FR ratio results in an equally rapid decrease of the ATHB-2 mRNA. Consistently, plants overproducing ATHB-2 show developmental phenotypes characteristic of plants grown in low R:FR: elongated petioles, reduced leaf area, early flowering, and reduced number of rosette leaves. Taken together, the data strongly suggest a direct involvement of ATHB-2 in light-regulated growth phenomena throughout Arabidopsis development.     

Tillering responses to red:far-red light ratio during different phenological stages in Eragrostis curvula

Tillering responses to light quality in different phenological stages of a perennial warm-season grass Eragrostis curvula were investigated in controlled environments. In vegetative plants, the tillering rate was greater (P<0.01) in the high (1.1–1.3) than in the low (0.59–0.70) red:far-red ratio (R/FR) light regime. Tillering rates were higher in the low R/FR treatment when the plants in the high R/FR regime reached the reproductive stage, while the plants in the low R/FR regime remained vegetative. After the reproductive tillers were removed by defoliation, more tillers were produced in the defoliated plants grown in the high R/FR regime. When the plants in both light treatments entered the reproductive stage, the tillering rate under the two light regimes became similar, suggesting a significant interaction between tillering and inflorescence development. The more advanced inflorescence development in the high R/FR regime may have reduced assimilate availability to tiller growth and overshadowed the effect of high R/FR on tillering. Both tillering and inflorescence development appeared to be controlled by R/FR ratio. The higher rate of aerial tiller production on the reproductive culms during the post-anthesis period in the high R/FR regime suggested that high R/FR ratio stimulated not only basal tillering, but also aerial tillering.