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.     

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)     

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.     

Photomorphogenesis in Chenopodium album. Effects of supplementary far-red light on the kinetics of stem extension

The effects of far-red light given against a background of white light on the stem-extension kinetics of three-week-old, light-grown Chenopodium album seedlings were investigated. Under white light alone, the stems (cotyledon-to-apex) extended almost exactly logarithmically with time. Under these conditions the increase in log [stem length in mm] per hour was approx. 3.7·10^-3, equivalent to about 1% per h during both skoto-and photoperiods. Supplementary far-red given throughout each photoperiod massively stimulated extension. The calculated logarithmic extension rate, however, slowly returned to that of the controls, following an initial large increase. This is predicted by a model in which far-red light linearly increases the extension rate of individual internodes which arise at an exponentially increasing rate. The behaviour of the model is also consistent with critical experiments in which far-red was given as a pre-treatment or transiently, as well as with other published data. Far-red stimulation of logarithmic extension rate in successive photoperiods was closely and linearly correlated with calculated phytochrome photoequilibrium. Daily short periods of supplementary far-red were especially potent in accelerating extension; the plants seemed least responsive at the end of the photoperiod.Abbreviations FR supplementary far-red light - I stem length (mm) - LSER logarithmic stem extension rate - Pfr far-red absorbing form of phytochrome - R:FR red:far-red fluence rate ratio - WL white light - _c calculated phytochrome photoequilibrium     

Photocontrol of stem elongation in light-grown plants of Fuchsia hybrida

Stems of the caulescent long-day plant, Fuchsia hybrida cv Lord Byron, showed 2 types of response to light. In one, internode length was increased by far-red irradiation given at the end of an 8 h photoperiod: the response was no greater with prolonged exposure and was less when the start of far-red was delayed. The effect of far-red was reversible by a subsequent exposure to red light. Internode length was inversely proportional to the P_fr/P ratio established before entry to darkness and there was no evidence for loss of P_fr during a 16 h dark period. The inhibitory effect of P_fr acted at a relatively late stage of internode growth. With the development of successive internodes a second response appeared in which stems lengthened following prolonged daily exposures to red or far-red light, or mixtures of the two, or to brief breaks with red or white light. In these later internodes, a short exposure to far-red near the middle of the night was not reversible by red because red alone promoted elongation at this time. Internode length increased with increase in the daily duration of light and, when light was given throughout an otherwise dark period of 16 h, with increase in illuminance to a saturation value of 200 lx from tungsten lamps. Elongation increased as a linear function of decrease in photostationary state of phytochrome down to P_fr/P0.3; however, internodes were shorter in far-red light than in 25% red/red+far-red. It was concluded that stem length is a net response to two modes of phytochrome action. An inductive effect of P_fr inhibits a late stage in internode expansion, and a phytochrome reaction which operates only in light (and may involve pigment cycling) promotes an early stage of internode development. Stem elongation is thus a function both of the daily duration of light and its red/red+far-red content. The outgrowth of axillary buds was controlled by the first type of phytochrome action only.Abbreviations and symbols FR far red light - R red light - P phytochrome - P_fr phytochrome in the far-red light absorbing form - SD 8 h short days - LDP long-day plant - SDP short-day plant     

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.     

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.     

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 control of short-day-induced bud set in black cottonwood

In trees and other woody perennial plants, short days (SDs) typically induce growth cessation, the initiation of cold acclimation, the formation of a terminal bud and bud dormancy. Phytochrome control of SD-induced bud set was investigated in two northern clones of black cottonwood (Populus trichocarpa Torr. & Gray) by using night breaks with red light (R) and far-red light (FR). For both clones (BC-1 and BC-2), SD-induced bud set was prevented when R night breaks as short as 2 min were given in the middle of the night. When night breaks with 2 min of R were immediately followed by 2 min of FR, substantial reversibility of bud set was observed for BC-1 but not for BC-2. By comparing the effects of the R night breaks on bud set and the length of specific internodes, we determined that the R night breaks influenced internode elongation in two opposing ways. First, the addition of a R night break to the SD treatment prevented the cessation of internode elongation that is associated with bud set. Those internodes that would not have elongated under SDs (and would have been found within the terminal bud) elongated in the R treatment. Second, the R night breaks decreased internode length relative to the long-day (LD) control. In contrast to the clonal differences in reversibility that we observed for bud set, the decrease in internode length (i.e. the second effect of R) was R/FR reversible in both clones. Based on these results, we conclude that internode elongation is influenced by two distinct types of phytochrome-mediated response. The first response is a typical response to photpperiod, whereas the second response is a typical “end-of-day” response to light quality. Our results demonstrate that SD-induced bud set in black cottonwood is controlled by phytochrome but that clonal differences have an important influence on the R/FR reversibility of this response. The availability of an experimental system in which SD-induced bud set is R/FR reversible will be valuable for studying the physiological genetics of photoperiodism in trees.     

Phytochrome effects on stem carbon gain in light-grown mustard seedlings are not simply the result of stem extension-growth responses

Carbon allocation and partitioning were investigated in the first internode of light-grown Sinapís alba L. seedlings exposed to white light (WL) with or without supplementary far-red light (FR). In the internode, supplementary FR increased the rates of extension-growth and the accumulation of radiolabeled carbon (fed through the leaves), reducing sugars (even per unit volume), starch, hemicellulose and cellulose, but had no effect on the levels of sucrose and ammonium oxalate-solubilised cell wall carbohydrates, on invertase activity or on the use of additional sucrose fed through the leaves. In source leaves, supplementary FR had no effect on photosynthesis rates and reduced the accumulation of radiolabeled carbon. Mechanical reduction of stem extension-growth responses to supplementary FR did not affect internode carbohydrate or carbon accumulation responses. Supplementary FR provided only to one leaf had no effect on internode extension growth but increased carbon accumulation in the internode. provided that supplementary FR and radiolabeled carbon were both given to the same leaf. Phytochrome-mediated effects on carbon partitioning are not the mere consequence of internode extension-growth responses. Some additional control point(s) (e.g. leaf-source strength) must be under the direct influence of phytochrome.     

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.     

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.     

Effects of irradiance and spectral quality on seedling development of two Southeast Asian Hopea species

 Seedling developmental responses to understory shade combine the effects of reductions in irradiance and changes in spectral quality. We studied the seedling development of two Southeast Asian dipterocarp trees in response to differences in irradiance (photosynthetic photon flux density, PPFD) and spectral quality (red to far-red ratio, R:FR). The two species, Hopea helferei and H. odorata, are taxonomically closely related but differ in their ecological requirements; H. helferei is more drought-tolerant and typically grows in more open habitats. Seedlings were grown in six different replicated shadehouse treatments varying in percentage of solar PPFD and R:FR. The two species differed in the influence of light variables on most seedling characters, particularly for final height, internode distance, branch/trunk internodes, stem length/mass, leaf area/stem length, petiole length, and growth/mol of photons received. Most of the characters in both taxa were primarily influenced by PPFD, but spectral quality also influenced some characters – more so for H. odorata. The latter species grew more rapidly, particularly in the low PPFD treatments, and its leaves were capable of higher photosynthesis rates. However, growth in H. helferei was not reduced in direct sunlight. The growth of this taxon may be constrained by adaptations, particularly in leaves, for drought tolerance. Received: 14 April 1996 / Accepted: 8 October 1996     

Modulation of leaf elongation, tiller appearance and tiller senescence in spring barley by far-red light

Supplemental far-red (FR) illumination of light-grown grass seedlings inhibits tiller production while enhancing leaf elongation. Although much is known about FR enhancement of internode elongation in dicots, relatively little research has been conducted to determine the effects of FR on monocot development. In growth chamber experiments, fibre optics were used to direct supplemental FR to elongating leaf blades, main stem bases and mature leaf blades of light-grown barley (Hordeum vulgare L.) seedlings. Our objective was to identify specific sites of perception for FR enhancement of leaf elongation and inhibition of tiller production, and to assess potential FR effects on tiller senescence. Far-red illumination of elongating leaves or of the main stem base reduced the total number of tillers per plant, primarily by reducing secondary and tertiary tiller production, and enhanced leaf elongation. However, leaf elongation was less sensitive to stem base treatments than to illumination of the elongating blade. Increased leaf length resulted from increased leaf elongation rate, while the duration of leaf elongation was unaffected. Exposure of mature leaf blades to FR had no effect on tillering or leaf elongation. None of the FR treatments led to tiller senescence. Localization of FR perception in vertically oriented tissues such as elongating blades and stem bases permits early detection of reflected light from neighbouring plants, allowing rapid response to impending competition.     

Rapid photomodulation of stem extension in light-grownSinapis alba L.

Treatment of the whole of aSinapis alba plant with supplementary far-red light (FR), in back-ground white light (WL), induces a rapid increase in stem extension rate. This rapid increase is regulated by the light environment of the stem itself. Supplementary FR to the stem increases extension rate after a lag period of 10–15 min. A lag period of 3–4 h follows FR irradiation of the leaf, before an increase in extension rate is detectable. When the stem is given supplementary FR, the change in extension rate which is induced increases with increasing FR fluence rate, and with decreasing phytochrome photoequilibrium. There is no difference between the effects of supplementary FR _max 719 nm and supplementary FR _max 739 nm for these relationships. The increase in extension rate induced by supplementary FR is reversed by an increase in the fluence rate of red light (R). These data indicate that the response is controlled by phytochrome photoequilibrium.Abbreviations B blue light - FR far-red light - R red light - WL white light - Pfr far-red absorbing form of phytochrome - Pr red absorbing form of phytochrome - P_tot total phytochrome level (=Pr+Pfr); -Pfr/Ptot, measured - ER difference in stem extension rate, before and after treatment     

Branching responses of a plagiotropic clonal herb to localised incidence of light simulating that reflected from vegetation

Plants sense the presence of neighbouring vegetation through phytochrome photoreceptors perceiving a lowered red to far-red ratio (R:FR) of light reflected from such vegetation. We hypothesised that it would be advantageous for the grassland clonal herb, Trifolium repens, to have an inhibitory branching response to perception by leaves of light reflected from neighbouring vegetation (i.e. light with lowered R:FR ratio) but have no response to interception of such light by the plagiotropic stem. We tested whether photoreception of reflected light by plagiotropic stems resulted in a different branching response to photoreception by leaves and whether leaf ontogeny influenced the response. To simulate light reflected from vegetation, FR light-emitting-diodes were used to supplement controlled environment room light so that the R:FR ratio, but not the photosynthetic photon fluence rate, of light incident at the stem or leaf of a phytomer of T. repens was lowered from 1.20 to 0.25. The plagiotropic stems were unresponsive to light simulating that reflected from vegetation. This response differs from that of stems of orthotropic species, indicating that plagiotropic stems have evolved an organ-specific photobiology. Treatment of the mature leaf with light of lowered FR ratio reduced phytomer production only of the branch in the axil of the treated leaf. Similar treatment of the immature leaf retarded, in addition, branching at basal phytomers on the same side of the primary stem axis. Thus the response to light simulating that reflected from neighbouring vegetation depended upon whether the light was incident at the stem or the leaf and on the stage of leaf development. We argue that such responses improve the performance and fitness of T. repens within grassland habitats by allowing axillary buds on plagiotropic stems to branch freely when stems are in receipt of light reflected from vegetation while leaves are in full light.     

Implications of opposite phyllotaxis for light interception efficiency of Mediterranean woody plants

Opposite leaves lead to a greater leaf overlapping than leaves spirally arranged along a shoot, decreasing light interception efficiency (Ea, fraction of the light reaching the plant actually intercepted by the leaves) of the crown. However, Ea results from a whole suite of morphological traits. The interplay between phyllotaxis, crown architecture, leaf morphology and Ea was explored in 12 woody species from Mediterranean-type ecosystems, where the abundance of woody species with opposite phyllotaxis is unusually high. The three-dimensional model Y-plant was used to estimate Ea in unbranched, vertical shoots of each species encompassing the natural morphological variation found from moderate shade to open light environments. Ea exhibited significant interspecific differences, ranging from 0.25 in Daphne gnidium to 0.75 in Cistus ladanifer, Olea europaea and Salvia officinalis, decreasing with leaf inclination angle and leaf area ratio (LAR), and increasing with internode-to-leaf-length ratio and supporting biomass. Species with spiral vs. opposite phyllotaxis did not differ in their mean Ea. However, the former had higher Ea than the latter at short internode lengths. The natural range of variation in internode length had a larger effect on Ea than the natural range of leaf elevation angle. Principal component analysis segregated species with opposite phyllotaxis from those with spiral leaves because of their greater self-shading for high sun elevation angles (>45°); they were in turn distributed in two groups, one with high Ea, large investment in supporting biomass and long internodes, and another with low Ea and large LAR. Species with spiral phyllotaxis all had intermediate or low Ea and steep leaf elevation angles. Species with opposite phyllotaxis can compensate their less efficient leaf arrangement by decreasing leaf elevation angle and increasing internode length, but they may experience a real phylogenetic constraint for light interception when biomass allocation to supporting tissues (internodes and petioles) becomes very costly. This constraint could be involved in the shade intolerance of woody Mediterranean species exhibiting opposite phyllotaxis.     

Does far-red light affect growth and mycorrhizas of Scots pine seedlings grown in forest soil?

We studied the response of Scots pine (Pinus sylvestris L.) to supplementary far-red sidelight in seedlings grown in a forest soil substrate without additional nutrient supply. Our aim was to determine possible changes in the accumulation and allocation of dry weight and mineral nutrients and the presence of mycorrhizas. Half of the seedlings were grown in light conditions simulating reflected far-red light (FR) from neighbouring plants and the other half were controls not receiving additional FR. PAR irradiance was kept constant in both treatments. At the first harvest (41 d of treatment), FR+ had no effect on stem height, biomass accumulation or allocation. However, at the end of the experiment (93 d of treatment), an increase in stem extension rate and stem dry weight was observed in FR+ seedlings when compared to controls. Both control and treated plants had several morphological types of ectomycorrhizas, but no effect of FR+ on the frequency of these morphotypes was observed. Nor was the concentration of ergosterol or estimated mycorrhizal fungal biomass affected. Nutrients were more responsive to the light quality treatment: P concentration in roots and N and P contents in stems and roots were higher in FR+ plants than in control seedlings. These results are in contrast to those of a previous study in which the root system of pine seedlings, which were fertilised and had less developed mycorrhizas, was reduced by FR+. This revised version was published online in June 2006 with corrections to the Cover Date.     

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.     

Phytochrome action in light-grown mustard: kinetics,fluence-rate compensation and ecological significance

Internode extension in young, light-grown mustard plants was measured continuously to a high degree of resolution using linear voltage displacement transducers. Plants were grown in background white light (WL) and the first internode was irradiated with supplementary far-red (FR) from fibre-optic light guides, depressing the Pfr/P (ratio of FR-absorbing form of phytochrome to total spectrophotometrically assayable phytochrome) within the internode and causing an acceleration of extension rate. The internode was sensitive to periods of FR as brief as 1 min, with a sharp increase in extension rate occurring after the return to background WL only. The mean latent period of the response to FR was approx. 10 min. Periods of FR longer than approx. 35 min caused an apparently biphasic growth response, with an initial sharp acceleration in extension rate (Phase 1) being followed by a brief deceleration and a further acceleration to a more-or-less steady elevated rate, somewhat less than the first peak (Phase 2). With such longer-term FR, extension rate decelerated upon FR switch-off after a mean lag of approx. 6 min, achieving the prestimulation extension rate within 16 min. The magnitude of the FR-induced increase in extension rate, expressed as a percentage of the rate in WL alone, was an inverse, linear function of the phytochrome photoequilibrium (i.e. Pfr/P, measured in etiolated test material irradiated under the same geometry) over the range 0.17 to 0.63. This relationship was not significantly affected by variations in backround WL fluence rate over the range 50–150 mol·m^-2·s^-1 and was held both for Phase 1 and Phase 2 of the response. The data provide evidence for rapid coupling/uncoupling between phytochrome and its transduction chain in the light-grown plant and for fluence-rate compensation of the regulation of extension rate. The extensive linearity of the relationship between phytochrome photoequilibrium and proportional extension rate increment allows for fine tuning in shade avoidance. The results are discussed with respect to recent evidence on the nature of phytochrome in light-grown plants and in relation to the function of phytochrome in plants growing in the natural environment.Abbreviations FR far-red light - LVDT linear voltage displacement transducer - P total spectrophotometrically assayable phytochrome - PAR photosynthetically active radiation (400–700 nm) - Pfr FR-absorbing form of phytochrome - Pr R-absorbing form of phytochrome - R red light - WL white light