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.     

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.     

Chloroplast Structure and Starch Grain Accumulation in Leaves That Received Different Red and Far-Red Levels during Development

An important step in understanding influence of growth environment on carbon metabolism in plants is to gain a better understanding of effects of light quality on the photosynthetic system. Electron microscopy was used to study chloroplast ultrastructure in developing and fully expanded leaves of tobacco (Nicotiana tabacum L. cv Burley 21). Brief exposures to red or far-red light at the end of each day during growth under controlled environments influenced granum size, granum number and starch grain accumulation in chloroplasts, and the concentration of sugars in leaf lamina. Far-red-treated leaves had chloroplasts with more but smaller grana than did red-treated leaves. Red light at the end of the photosynthetic period resulted in more and larger starch grains in the chloroplasts and a lower concentration of sugars in leaves. Chloroplast ultrastructure and starch grain accumulation patterns that were initiated in the expanding leaves were also evident in the fully expanded leaves that received the treatment during development. It appears that the phytochrome system in the developing leaves sensed the light environment and initiated events which influenced chloroplast development and partitioning of photosynthate to adapt the plant for better survival under those environmental conditions.     

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.     

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.     

Tillering Responses to the Light Environment and to Defoliation in Populations of Perennial Ryegrass (Lolium perenneL.) Selected for Contrasting Leaf Length

Tillering responses to the light environment and to defoliationwere studied in two populations of perennial ryegrass (LoliumperenneL.) selected for contrasting leaf lengths. The objectiveof this study was to determine whether differences in tilleringresponses between populations, as a result of management practices,affected their competitive ability. Young seedlings were exposed,under controlled conditions, to defoliation, neutral shading(decreased photosynthetic photon flux), low red:far-red ratioand/or decreased blue light. Selection for longer leaves reducedthe tillering rate. After defoliation, this difference betweenshort- and long-leaved populations was magnified. Defoliationdecreased both site filling and phyllochron of the long-leavedpopulation but had no effect on the short-leaved population.Lowering the photosynthetic photon flux reduced the phyllochronin both population. Decreasing the red:far-red ratio reducedtillering rate by reducing site filling, whereas decreasingblue light had no significant effects on tillering. Tilleringresponses to photosynthetic photon flux and to red:far-red ratiowere similar in the two populations selected for contrastingleaf length. The implication of these tillering responses indefining the competitive ability of the grass plants is discussedin relation to their management.Copyright 1999 Annals of BotanyCompany Blue light, defoliation, far-red, irradiance, leaf size,Lolium perenne(L.), perennial ryegrass, photomorphogenesis, phyllochron, shading, site filling, tiller production.     

Light quality effects on the appearance of tillers of different order in wheat (Triticum aestivum)

Plants of Triticum aestivum cvs Norkin Pan 70, La Paz Inta, Buck Cencerro and Buck Cimarron were grown outdoors in individual pots with two sowing dates and irradiated at the end of the day either with red or far-red light. In red-treated plants the number of tillers was close to the potential calculated from the number of leaves on the main shoot but far-red treated plants produced less tillers. The magnitude of the effect was larger for secondary and tertiary than primary tillers. Thus, the proportion of primary to secondary and tertiary tillers was larger in FR-treated plants. Leaf sheath and lamina were longer under FR in the second sowing date, possibly due to warmer nights. These responses were greatly affected by the genotype. Significant differences in ear number were not found but in one of the sowing dates red-light-treated plants showed a lower grain number and yield than those treated with far-red.     

Action and interaction of red and far-red radiation on lipoxidase metabolism of squash seedlings

Lipoxidase, in the cotyledons of squash (Cucurbita moscata) seedlings grown in the dark, reached its peak activity on the fifth day and then declined to its lowest activity on the eighth day. Under continuous irradiation, the rate of enzyme disappearance was accelerated by red (655 mμ) and was retarded by far-red (735 mμ) radiation. Acceleration of enzyme disappearance caused by red light was reversed repeatedly by far-red light in seedlings that received an initial exposure to red radiation. These responses were independent of the duration of irradiation at each of the alternating wavebands. No change was observed when the white light was administered either 24 hours before or 24 hours after the red, far-red treatment.

The lipoxidase system of the seedlings given an initial exposure to far-red radiation also responded reversibly to alternating far-red, red extended exposures, but it failed to respond reversibly when short exposures were employed. Similarly, no change occurred in these seedlings when either pre- or post-treatment with the white light was applied.

These results demonstrate that the capacity of lipoxidase to act reversibly depends primarily on the duration of exposure and on the kind of light (red or far-red) to which the seedlings were exposed initially. In spite of these variations, lipoxidase metabolism can be considered an additional biochemical manifestation of red, far-red reaction that operates in the photomorphogenesis of plants.

     

Morphogenetic Changes Induced by a Low Red: Far-Red Ratio and their Growth Consequences in Water Hyacinth (Eichhornia crassipes)

Plants of water hyacinth (Eichhornia crassipes) were grown undertwo red/far-red ratios (Z) to investigate the effects on morphologyand growth of the light quality component of canopy shade. Experimentswere conducted in diffuse sunlight in the presence or absenceof far-red radiation. Under low Z conditions, fewer new ramets were produced and theirstolons were shorter. The number of leaves per ramet was unchanged,but petiole length and blade area were increased. This changein biomass partitioning resulted in a lower investment in themain resource-acquiring organs (leaf blades and roots). Thelower allocation of biomass to the leaf blades was compensatedfor by a higher net assimilation rate, such that relative growthrate remained unchanged. Key words: Biomass partitioning, Eichhornia crassipes, growth, morphogenesis, red: far-red ratio     

Variations in Tiller Dynamics and Morphology in Lolium multiflorum Lam.Vegetative and Reproductive Plants as affected by Differences in Red/Far-Red Irradiation

Lolium multiflorum Lam, plants were grown in a growth room undertwo light sources with red/far-red ratios of 1·62 and0·84 but similar photosynthetically active radiation.In both situations the capacity to produce new tillers and thelight available per tiller decreased with canopy growth. Tilleringwas further reduced by the low red/far-red ratio while lightinterception and plant dry weight were unaffected by this treatment.In both reproductive and vegetative plants under the lower red/far-redratio the time between leaf expansion and the appearance ofa tiller in its axil was increased and the proportion of ‘maturebuds’ that developed was reduced. Irradiation with lowred/far-red advanced the reproductive development and increasedthe number of fertile tillers per plant. It also caused longerleaf sheaths, blades and reproductive shoots. The results suggestthat as canopy density increases the lower light interceptionper tiller and the photomorphogenic effect of low red/far-redratios may reduce the capacity to produce new tillers. Lolium multiflorum, Lam., annual ryegrass, tillering, tiller growth, leaf growth, flowering, light quality.     

Localization of expression of KNAT3, a class 2 knotted1-like gene

KNAT3 is a class 2 kn1 -like gene in Arabidopsis thaliana . The RNA expression patterns of KNAT3 were characterized through the use of promoter-GUS fusion analysis and in situ hybridization. KNAT3 is expressed in several tissues and at several times during development. There are three main expression patterns: (1) during early organ development in young leaves, buds and pedicels; (2) at and near the junction between two organs at specific times during development, including the hypocotyl-root boundary in young seedlings, the anther-filament junction in mature flowers, and the ovule-funiculus and peduncle-silique boundaries in elongating siliques; and (3) in maturing tissues such as the style of elongating siliques, the petioles of maturing leaves, and most of the root. The varied expression patterns may indicate that KNAT3 plays several different roles in plants, depending on when and where it is expressed. Previous work on KNAT3 (Serikawa et al. , 1996) indicated that expression of its RNA is regulated by light. Promoter-GUS seedlings were grown under different light conditions (continuous white, red and far-red light) to examine more closely the light regulation of the KNAT3 promoter. Continuous white light resulted in stronger overall GUS staining in the same patterns seen in seedlings grown under long-day conditions (cotyledons, upper hypocotyl and roots). Continuous red light resulted in reduced GUS expression in those same tissues. Continuous far-red light led to seedlings showing stronger staining in the hypocotyl and cotyledons than red light-grown plants but no staining in the roots. Thus, the KNAT3 promoter responds differently to red and far-red light.     

Cotton plant size and fiber developmental responses to FR/R ratio reflected from the soil surface

Cotton ( Gossypium hirsutum L.) plants were grown in irrigated field plots over red, green or white soil covers (mulches). The far-red (FR) to red (R) light ratios were higher in upwardly reflected light over the red and green surfaces than in incoming sunlight. Plants that grew over the white mulches received higher photosynhetic photon flux (PPF), but the reflected FR/R ratio did not differ significantly from that in incoming sunlight. At five weeks after emergence, seedling stern lengths were significantly longer over red and green than over white surfaces. At maturity, plants that had grown over the red and green surfaces had longer stems, larger shoots, more bolls (fruit), more seed cotton, and longer fibers than plants grown over the white surfaces even though those grown over the white surfaces had received more reflected photosynthetic light during growth and development.     

Oak Seedlings Grown in Different Light Qualities

Seedlings of oak (Quercus robur) were germinated in darkness for 3 weeks and then given continuous light or short pulses of light (5–8 min every day). The morphological development was followed during 25 days. In continuous white, blue, and red light the stem growth terminated after about 10 days by formation of a resting bud. At that time the seedlings were about 100 mm high. In con tinuous long wavelength farred light (wavelength longer than 700 nm) the stem growth including leaf formation was continuous without the formation of resting buds, and the stem length was about 270 mm after 25 days. The number of nodes developed became twice that of the seedlings grown in while light. The leaves became well developed in all light colours, but leaf areas were largest in plants cultivated in white light. Compared to dark grown seedlings the mean area per leaf was increased about five times in continuous long wavelength far red light. A supplement with short (5 min) pulses of red light each day increased the leaf area up to 20 times. The stem elongation showed a high energy reaction response, i.e. the stem length increased only in continuous long wavelength far-red light but was not influenced by short pulses of red light or far-red light. The leaf expansion, however, was increased by short pulses of red light with a partial reversion of the effect by a subsequent pulse of far-red light. The fraction of the plant covered with periderm was higher in plants given continuous light. In respect to periderm inhibition continuous long wavelength far red light was the most effective. The transfer of seedlings from darkness to continuous white light gave anthocyanin formation in the stem 10–20 mm below the apex. This formation took place in the cortex and was evident in plants grown in darkness or under short pulses of light. Plants grown in continuous red, blue or long wavelength Far red light showed only traces of anthocyanin.     

Biomass allocation, shade tolerance and seedling survival of the invasive species

Berberis darwinii (Berberidaceae) is a serious environmental weed in New Zealand, capable of invading a range of different light environments from grazed pasture to intact forest. According to optimal partitioning models, some plants optimise growth under different environmental conditions by shifting biomass allocation among tissue types (e.g. roots, shoots) to maximise the capture of limiting resources (e.g. water, light). We examined patterns of growth, biomass allocation, and seedling survival in Berberis darwinii to determine whether any of these factors might be contributing to invasion success. Growth and biomass allocation parameters were measured on seedlings grown for 7 months in five natural light environments in the field. Survival was high in the sunniest sites, and low in the shadiest sites. Seedlings grown in full sun were an order of magnitude taller and heavier, had five times as many leaves, and proportionally more biomass allocated to leaves than seedlings grown in other light environments. In the shade, leaves were bigger and thinner, and leaf area as a proportion of total plant biomass increased, but the proportion of above- to below-ground biomass was similar across all light and soil moisture environments. In summary, although leaf traits were plastic, patterns of biomass allocation did not vary according to optimal partitioning models, and were not correlated with patterns of seedling survival. Implications for the management of this invasive species are discussed.     

Morphology and Photosynthetic Efficiency of Tobacco Leaves That Received End-of-Day Red and Far Red Light during Development

Shaded leaves in plant canopies receive a higher proportion of far red relative to red light than is received by unshaded leaves. Brief end-of-day irradiations with red or far red light, acting through the phytochrome system, reversibly control morphological development of tobacco plants. Leaves that received far red light for 5 minutes at the end of each day during development were longer and narrower than those that received end-of-day red light. The far red treated leaves weighed less, had fewer stomata, and had less chlorophyll per unit area of leaf. Net CO₂ assimilation rates did not differ significantly between red- and far red-treated leaves on an area basis; however, the far red-treated leaves assimilated significantly more CO₂ on a leaf weight basis.     

Maize leaves turn away from neighbors

In commercial crops, maize (Zea mays) plants are typically grown at a larger distance between rows (70 cm) than within the same row (16-23 cm). This rectangular arrangement creates a heterogeneous environment in which the plants receive higher red light (R) to far-red light (FR) ratios from the interrow spaces. In field crops, the hybrid Dekalb 696 (DK696) showed an increased proportion of leaves toward interrow spaces, whereas the experimental hybrid 980 (Exp980) retained random leaf orientation. Mirrors reflecting FR were placed close to isolated plants to simulate the presence of neighbors in the field. In addition, localized FR was applied to target leaves in a growth chamber. During their expansion, the leaves of DK696 turned away from the low R to FR ratio signals, whereas Exp980 leaves remained unaffected. On the contrary, tillering was reduced and plant height was increased by low R to FR ratios in Exp980 but not in DK696. Isolated plants preconditioned with low R/FR-simulating neighbors in a North-South row showed reduced mutual shading among leaves when the plants were actually grouped in North-South rows. These observations contradict the current view that phytochrome-mediated responses to low R/FR are a relic from wild conditions, detrimental for crop yield.     

How portulaca seedlings avoid their neighbours

Summary Portulaca oleracea L. seedlings do not develop in the direction of neighbours, even when these neighbours are small and distant. Neighbouring plants could be simulated by small rectangles of a plastic that resembled leaves in its spectral characteristics. Unlike seedlings, mature plants did not respond to objects that do not influence photosynthetic light. When light of equal intensity was received from all directions, Portulaca seedlings avoided the direction with higher far-red light. Portulaca is thus able to use spectral composition and direction of light as clues for the probability of the direction of future shade.     

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.