Grassland canopy composition and spatial heterogeneity in the light quality

Both vertical and horizontal heterogeneity in red light levels and in the red/far red ratio were measured in a mountain grassland and correlated with the vegetation structure. Vertical change in the red/far red ratio was measured in three communities; canopy structure and density was investigated by the point-quadrat method and biomass harvesting. Quantities of both grasses and dicots were significantly correlated with R/FR ratio and red light intensity at ground level. In dicots, percentage cover determines their effect on light levels; in contrast, no single parameter suffices to capture the effect of grasses: grass biomass, number of hits (cover) and the mean height (distance from the soil surface) of grass hits are of importance. This indicates strikingly different geometry and optical properties of both plant groups.The fine-scale horizontal heterogeneity was assessed by means of a grid of 3.3×3.3 cm cells (plots of 8× cells) in the species-poor community. Both red light and R/FR ratio were determined at soil level in all the grid cells; fine-scale vegetation recording was done by counting all rooted stems (leaves for large rosette plants) in these cells. Spatial autocorrelation (Moran's I) of fine-scale heterogeneity in light levels, biomass per cell and individual species occurrences revealed many significant autocorrelations. Light levels (particularly red light) show autocorrelations at the distance of 1–2 cells (3.3–6.7 cm); biomass shows little autocorrelation. A multivariate analysis (redundancy analysis) showed that cellwise densities of two species had significant and systematic correlations with the light levels (both red light and R/FR ratio): a grass species with wide, mainly horizontal leaves, Anthoxanthum alpinum, was negatively correlated with light levels; Nardus stricta, with upright and narrow leaves and stems was positively correlated with light levels.     

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.     

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.     

Temperature‐dependent shade avoidance involves the receptor‐like kinase ERECTA

Plants detect the presence of neighbouring vegetation by monitoring changes in the ratio of red (R) to far‐red (FR) wavelengths (R:FR) in ambient light. Reductions in R:FR are perceived by the phytochrome family of plant photoreceptors and initiate a suite of developmental responses termed the shade avoidance syndrome. These include increased elongation growth of stems and petioles, enabling plants to overtop competing vegetation. The majority of shade avoidance experiments are performed at standard laboratory growing temperatures (>20°C). In these conditions, elongation responses to low R:FR are often accompanied by reductions in leaf development and accumulation of plant biomass. Here we investigated shade avoidance responses at a cooler temperature (16°C). In these conditions, Arabidopsis thaliana displays considerable low R:FR‐mediated increases in leaf area, with reduced low R:FR‐mediated petiole elongation and leaf hyponasty responses. In Landsberg erecta, these strikingly different shade avoidance phenotypes are accompanied by increased leaf thickness, increased biomass and an altered metabolite profile. At 16°C, low R:FR treatment results in the accumulation of soluble sugars and metabolites associated with cold acclimation. Analyses of natural genetic variation in shade avoidance responses at 16°C have revealed a regulatory role for the receptor‐like kinase ERECTA.     

Ecological significance of light quality in optimizing plant defence

Plants balance the allocation of resources between growth and defence to optimize fitness in a competitive environment. Perception of neighbour‐detection cues, such as a low ratio of red to far‐red (R:FR) radiation, activates a suite of shade‐avoidance responses that include stem elongation and upward leaf movement, whilst simultaneously downregulating defence. This downregulation is hypothesized to benefit the plant either by mediating the growth‐defence balance in favour of growth in high plant densities or, alternatively, by mediating defence of individual leaves such that those most photosynthetically productive are best protected. To test these hypotheses, we used a 3D functional–structural plant model of Brassica nigra that mechanistically simulates the interactions between plant architecture, herbivory, and the light environment. Our results show that plant‐level defence expression is a strong determinant of plant fitness and that leaf‐level defence mediation by R:FR can provide a fitness benefit in high densities. However, optimal plant‐level defence expression does not decrease monotonically with plant density, indicating that R:FR mediation of defence alone is not enough to optimize defence between densities. Therefore, assessing the ecological significance of R:FR‐mediated defence is paramount to better understand the evolution of this physiological linkage and its implications for crop breeding.     

Canopy studies on ethylene-insensitive tobacco identify ethylene as a novel element in blue light and plant-plant signalling

Plants growing at high densities express shade avoidance traits as a response to the presence of neighbours. Enhanced shoot elongation is one of the best researched shade avoidance components and increases light capture in dense stands. We show here that also leaf movements, leading to a more vertical leaf orientation (hyponasty), may be crucial in the early phase of competition. The initiation of shade avoidance responses is classically attributed to the action of phytochrome photoreceptors that sense red:far-red (R:FR) ratios in light reflected by neighbours, but also other signals may be involved. It was recently shown that ethylene-insensitive, transgenic (Tetr) tobacco plants, which are insensitive to the gaseous plant hormone ethylene, have reduced shade avoidance responses to neighbours. Here, we report that this is not related to a reduced response to low R:FR ratio, but that Tetr tobacco plants are unresponsive to a reduced photon fluence rate of blue light, which normally suppresses growth inhibition in wild-type (WT) plants. In addition to these light signals, ethylene levels in the canopy atmosphere increased to concentrations that could induce shade avoidance responses in WT plants. Together, these data show that neighbour detection signals other than the R:FR ratio are more important than previously anticipated and argue for a particularly important role for ethylene in determining plant responses to neighbours.     

Keeping up with the neighbours: phytochrome sensing and other signalling mechanisms

Plants ‘forage’ for light in plant canopies using a variety of photosensory systems. Far-red radiation (FR) reflected by neighbours is an early signal of competition that elicits anticipatory shade-avoidance responses. In Arabidopsis and cucumber, perception of reflected FR requires phytochrome B. Horizontal blue (B) light gradients also guide plant shoots to canopy gaps in patchy vegetation, and these B light signals are perceived by specific photoreceptors. When plants are shaded by neighbours they undergo extensive reprogramming of their morphological development. Although phytochromes and B light receptors are certainly involved in these responses to shading, other sensory systems probably play important roles in the field. Recent studies of plant–plant signalling are unveiling a paradigm of sensory diversity and sophistication, which has important implications for understanding the functioning of plant populations and communities.     

Canopy height and seed yield in winter linseed (Linum usitatissimum) can be influenced by proximity to pink canes

Changes to growth and development in a winter linseed crop were observed adjacent to pink canes used as markers in 1997. Notably, canopy height was increased in plants positioned up to 40 cm from the pink canes. In 1998 these effects were confirmed and further characterised in a replicated experiment: extension growth occurred approximately one week earlier and canopy height was increased by 14.5 cm in plants adjacent to pink canes. The number of seed capsules was 953 m^?2 in control plots and 2567 m^?2 in plots with pink canes. Seed yield, calculated from a 0.25 m² quadrat, was 12.4 g m² in control plots and 56.1 g m^?2 in plots with pink canes. The R:FR (red:far red) ratio of light reflected from canes was approximately 1.7 compared with approximate R:FR ratios of 1.4 in incident daylight and 0.2 in light reflected from a linseed canopy (assessed using a spectral analysis system). Possible mechanisms to explain the changes in crop performance are discussed.     

Dual effect of light on flowering and sprouting of rose shoots

Shade, caused by a dense leaf canopy in the light conditions of a normal greenhouse, reduced sprouting of the third axillary bud (from the top) on decapitated rose branches ( Rosa hybrida cv. Marimba) in comparison to less shaded buds on branches protruding above the canopy and sparsely spaced. Flowering of the third young shoot on shaded branches bearing 3 lateral shoots was totally inhibited. Mixed fluorescent and incandescent light in a growth chamber reduced sprouting of the third bud on decapitated rose branches in comparison to decapitated branches on rose plants held in fluorescent light of similar photon flux density. This was attributed to the higher R:FR ratio in fluorescent vs mixed light that reached the third bud, and in exposed vs shaded branches. Flowering of the third shoot was promoted by several factors: high photon flux density, 0.5 m M gibberellic acid (GA) or 0.2 m M benzyladenine (BA). BA was the most effective treatment. Treatments promoting flowering of the third shoot did not reduce growth or flowering of the upper shoots. However, spraying the uppermost shoot with BA suppressed the growth of the shoots below. It is concluded that light affects flowering in two ways. The effect on bud sprouting is related mainly to R:FR ratios, while the effect on flower development is related mainly to photon flux density. Cytokinins may substitute for the light effect on flower development.     

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.     

红光与远红光比值对温室切花菊形态指标、叶面积及干物质分配的影响

以切花菊品种‘神马’(Chrysanthemum morifolium Ramat ‘Jinba’)为试材,于2010-2011年设计不同红光(R: (660 ±10) nm)与远红光(FR: (730±10)nm)比值(R/FR分别为0.5、2.5、4.5、6.5)的LED灯照射处理,研究不同R/FR值对温室切花菊形态指标、叶面积形成及干物质分配的影响。结果显示R/FR=2.5处理的植株叶片数、株高、茎粗、花径、叶面积及总干重均为各个处理中最高,R/FR=0.5处理的节间最长。所有R/FR处理的单株地上干物质重量与光质处理天数呈指数-线性模型。随处理天数的增加不同R/FR值处理菊花植株地上部分及地下部分干物质分配指数差异均不显著,叶片和花的干物质分配指数随处理天数的增加分别呈降低和升高的趋势,茎干物质分配指数则呈现先升高后降低的趋势,R/FR=2.5处理下,菊花叶片干物质分配指数和花干物质分配指数最高,而茎干物质分配指数却为最低;R/FR=6.5处理茎干物质分配指数最高,叶片干物质分配指数最低;0.5处理花朵干物质分配指数最低,说明远红光比例增加能够促进干物质向茎中分配,R/FR=2.5处理利于干物质向花朵中分配。     

Tree seedling canopy responses to conflicting photosensory cues

Light with decreased red:far-red (R:FR) ratios may signal neighbor presence and trigger plant developmental responses. There is some evidence that plant canopies forage towards increased R:FR ratios, but it is unclear to what extent R:FR versus the total amount of photosynthetically active radiation (PAR) influences canopy foraging responses among forest trees. The objective of this study was to examine the relative importance of PAR and R:FR as photosensory cues leading to tree canopy foraging responses. Seedlings of Betula papyrifera Marshall (paper birch) were grown in an experimental garden. Each seedling was germinated and grown in its own shading structure and exposed to two spatially separated light environments, in a factorial design of PAR and R:FR. Plant canopy foraging was evaluated at the end of one growing season in terms of canopy displacement, canopy area, leaf number, direction of stem lean, petiole aspect, and lamina aspect with respect to experimental light treatments. Leaf number and canopy area were greater on the high PAR sides of plants, irrespective of the R:FR treatment. Seedling canopies were displaced towards the direction of high PAR, but this relationship was not significant across all treatments. Petiole aspect was random and showed no significant directedness towards any of the light treatments. Lamina aspect and the direction of stem lean were distributed towards the direction of high PAR, irrespective of the R:FR treatment. Overall, first-year B. papyrifera seedlings used PAR, rather than R:FR ratio, as a photosensory cue for canopy light foraging.     

THE GENETIC ARCHITECTURE OF PLASTICITY TO DENSITY IN IMPATIENS CAPENSIS

Plant responses to crowding may be mediated by resource availability and/or by a specific environmental cue, the ratio of red:far red wavelengths (R:FR) perceived by phytochrome. This study examined the contribution of phytochrome-mediated photomorphogenesis to genetic variation in plastic responses to density in the annual plant Impatiens capensis. Inbred lines derived from open and woodland populations were grown under low density high density, and high density with selective removal of FR wavelengths to block phytochrome-mediated perception of neighbor proximity. Genetic variation in plasticity to density and to the R:FR cue was detected for several traits Plants grown at high density displayed increased internode elongation; decreased branch, flower, and node production; increased menstem dormancy; and decreased leaf area and specific leaf weight compared to plants grown at low density. Stem elongation responses to density were suppressed when phytochrome perception was blocked at high density. For these phytochrome-mediated traits, a genotype's plasticity to density was strongly correlated with its response to R:FR. Phytochrome-mediated traits were tightly correlated with one another, regardless of the density environment. However, the responses to density of meristem allocation to branching and leaf traits were less strongly phytochrome-mediated. These traits differed in patterns of plasticity, and their genetic correlations often differed across environments. In particular, genetic trade-offs involving meristem allocation to branching were expressed only at low density. The observed density dependence of phenotypic and genetic correlations implies that indirect selection and the potential for correlated response to selection will depend upon the competitive environment. Thus the differential sensitivity of characters to the R:FR cue can influence the evolution of integrated plastic responses to density.     

The regulation of cell wall extensibility during shade avoidance: a study using two contrasting ecotypes of Stellaria longipes

Shade avoidance in plants involves rapid shoot elongation to grow toward the light. Cell wall-modifying mechanisms are vital regulatory points for control of these elongation responses. Two protein families involved in cell wall modification are expansins and xyloglucan endotransglucosylase/hydrolases. We used an alpine and a prairie ecotype of Stellaria longipes differing in their response to shade to study the regulation of cell wall extensibility in response to low red to far-red ratio (R/FR), an early neighbor detection signal, and dense canopy shade (green shade: low R/FR, blue, and total light intensity). Alpine plants were nonresponsive to low R/FR, while prairie plants elongated rapidly. These responses reflect adaptation to the dense vegetation of the prairie habitat, unlike the alpine plants, which almost never encounter shade. Under green shade, both ecotypes rapidly elongate, showing that alpine plants can react only to a deep shade treatment. Xyloglucan endotransglucosylase/hydrolase activity was strongly regulated by green shade and low blue light conditions but not by low R/FR. Expansin activity, expressed as acid-induced extension, correlated with growth responses to all light changes. Expansin genes cloned from the internodes of the two ecotypes showed differential regulation in response to the light manipulations. This regulation was ecotype and light signal specific and correlated with the growth responses. Our results imply that elongation responses to shade require the regulation of cell wall extensibility via the control of expansin gene expression. Ecotypic differences demonstrate how responses to environmental stimuli are differently regulated to survive a particular habitat.     

Simulating the effects of localized red:far-red ratio on tillering in spring wheat (Triticum aestivum) using a three-dimensional virtual plant model

The outgrowth of tiller buds in Poaceae is influenced by the ratio of red to far-red light irradiance (R:FR). At each point in the plant canopy, R:FR is affected by light scattered by surrounding plant tissues. This paper presents a three-dimensional virtual plant modelling approach to simulate local effects of R:FR on tillering in spring wheat (Triticum aestivum). R:FR dependence of bud outgrowth was implemented in a wheat model, using three hypothetical responses of bud extension to R:FR (unit step, curvilinear and linear response). Bud break occurred when a threshold bud length was reached. Simulations were performed for three plant population densities. In accordance with experimental observations, fewer tillers per plant were simulated for higher plant population densities. The linear and curvilinear responses caused a delay in the increase in tiller number compared with experimental data. The unit step response approached experimental results best. It is suggested that a model based on relatively simple relations can be used to simulate degree of tillering. This study has shown that the virtual plant approach is a promising tool with which to address crop morphological and ecological research questions where the determining factors act at the level of the individual plant organ.     

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     

Contrasting shade avoidance responses in two perennial grasses: a field investigation in simulated sparse and dense canopies

We designed an experiment with potted plants grown outdoors to investigate the expression of shade avoidance in simulated sparse and dense canopies by two perennial grasses known to express contrasting responses to low red:far-red ratios (R:FR). Plants were grown in canopy microenvironments designed to lower the R:FR by reflection of horizontally propagated FR from neighbors and by direct attenuation of R by filters located above plants. Two specific hypotheses were tested: (1) Paspalum dilatatum will express greater shade avoidance than Schizachyrium scoparium to low R:FR in both sparse and dense canopies, and (2) low R:FR will produce greater expressions of shade avoidance in sparse than in dense canopies in both species. P. dilatatum was more responsive to low R:FR than S. scoparium in both the sparse and dense canopies and lower ramet number plant^–1 was the only common shade avoidance response between species in sparse canopies. P. dilatatum also showed significant reductions in juvenile ramet initiation, juvenile ramet mass, total shoot mass, and shoot:root ratios in sparse canopies, but only juvenile ramet initiation was reduced in dense canopies. The suppression of juvenile ramet initiation in the dense canopy was at least partially modulated by the vertically propagated R:FR because a similar reduction in PFD and horizontally propagated R:FR showed 42% greater juvenile ramet initiation in the respective control. S. scoparium only showed a significant reduction in ramet number plant^–1 and a significant increase in blade length in sparse canopies, but no significant responses occurred in dense canopies. Consequently, neither hypothesis was rejected. Variable shade avoidance responses between species and canopy densities indicate that both interspecific variation and various proportions of vertically and horizontally propagated low R:FR can influence the expression of shade avoidance responses of perennial grasses in field settings.     

The reflectance properties of plant internodes modify elongation responses to lateral far-red radiation

The quality and quantity of light reflected from Nicotinia tabacum L. internodes was monitored as the angle of incidence was varied. Reflectance of incident light, which was either normal or longitudinal to the internode axis, was investigated. Increasing the angle of incidence caused a greater proportion of incident light to be reflected. Light striking N. tabacum internodes was always modified prior to reflection, and smaller incident angles produced greater modification of the reflected light quality. At larger angles, interactions with the internode were reduced. As a parallel investigation, the extension growth rate of light-grown Sinapis alba L. seedlings was monitored continuously using sensitive linear displacement transducers. When the angle of illumination by fibre-optic probes, presenting far-red light to the growing internode, was changed from 0° to 45°, an increase was observed in stem extension rate. There is a possibility that this rate increase was the result of a fall in the red:far-red ratio (R:FR) present inside the plant internode as the angle of incidence was altered. However, it is more likely that it was due to the larger surface area of stem being illuminated. The consequences of these observations are discussed in relation to the potential influence of such modified reflections on canopy light environments and resultant shade responses which may occur when light of known R:FR impinges on plant internodes at angles other than 0°. The possibility is discussed that plants may perceive the quality of reflected radiation from neighbouring plants to be substantially different dependent upon the angle at which it is reflected.     

The Shade Avoidance Syndrome in Arabidopsis: The Antagonistic Role of Phytochrome A and B Differentiates Vegetation Proximity and Canopy Shade

Light limitation caused by dense vegetation is one of the greatest threats to plant survival in natural environments. Plants detect such neighboring vegetation as a reduction in the red to far-red ratio (R:FR) of the incoming light. The low R:FR signal, perceived by phytochromes, initiates a set of responses collectively known as the shade avoidance syndrome, intended to reduce the degree of current or future shade from neighbors by overtopping such competitors or inducing flowering to ensure seed production. At the seedling stage these responses include increased hypocotyl elongation. We have systematically analyzed the Arabidopsis seedling response and the contribution of phyA and phyB to perception of decreased R:FR, at three different levels of photosynthetically active radiation. Our results show that the shade avoidance syndrome, induced by phyB deactivation, is gradually antagonized by phyA, operating through the so-called FR-High Irradiance Response, in response to high FR levels in a range that simulates plant canopy shade. The data indicate that the R:FR signal distinguishes between the presence of proximal, but non-shading, neighbors and direct foliar shade, via a intrafamily photosensory attenuation mechanism that acts to suppress excessive reversion toward skotomorphogenic development under prolonged direct vegetation shade.