Ethylene is required in tobacco to successfully compete with proximate neighbours

Plants sense neighbours even before these cause a decrease in photosynthetic light availability. Light reflected by proximate neighbours signals a plant to adjust growth and development, in order to avoid suppression by neighbour plants. These phenotypic changes are known as the shade‐avoidance syndrome and include enhanced shoot elongation and more upright‐positioned leaves. In the present study it was shown that these shade‐avoidance traits in tobacco (Nicotiana tabacum) are also induced by low concentrations of ethylene. Furthermore, it was shown that transgenic plants, insensitive to ethylene, have a delayed appearance of shade‐avoidance traits. The increase in both leaf angles and stem elongation in response to neighbours are delayed in ethylene‐insensitive plants. These data show that ethylene is an important component in the regulation of neighbour‐induced, shade‐avoidance responses. Consequently, ethylene‐insensitive plants lose competition with wild‐type neighbours, demonstrating that sensing of ethylene is required for a plant to successfully compete for light.     

The function, action and adaptive significance of phytochrome in light-grown plants

Abstract. It has previously been proposed that the fundamental function of phytochrome in the natural environment is the perception of the relative proportions of red and far-red light, i.e. the red: far-red ratio. This paper re-evaluates this hypothesis, for vegetative green plants, in the light of recent findings. Essentially, three issues are considered: (a) the modulation of the response to red: far-red by fluence rate: (b) the anticipation of competition for light by perception of changes in red: far-red that precede actual shading: and (c) characteristics of phytochrome that may be important in the mechanism of photoperception (i.e. the accumulation of photoconversion intermediates, and the stability of Pfr). We conclude: (a) the red: far-red ratio provides a reliable signal of plant density, even before shading by neighbours occurs: (b) plants are able to perceive and respond to these signals, and that possible ambiguities due to low red: far-red at low solar angles may be avoided by modulation of the perception process by fluence-rate dependent mechanisms; (c) although direct experimental evidence does not yet exist, circumstantial evidence suggests that the perception of red: far-red may confer positive adaptive advantage; and (d) plants of certain species perceive and respond to fluence rate changes, mediated perhaps by a blue-light absorbing photoreceptor or by phytochrome, but that these responses do not necessarily lead to shade avoidance reactions and their ecological relevance is not fully understood.     

Plant-plant signalling, the shade-avoidance response and competition

Plants generate and perceive informational signals. An interaction (between plants) is considered to be informational when it involves the exchange of an insignificant amount of matter or energy, in quantitative terms, but in spite of this has a profound effect on plants by modulating their developmental programme. This article discusses how plants 'use' light signals to detect neighbouring plants and outlines the implications of shade-avoidance responses for the development of size and fitness hierarchies in canopies. The role of shade avoidance as a stabilizing force in mono-specific canopies is noted.Keywords: Competition, informational signals, photomorphogenesis, phytochrome, shade avoidance      

Characterization of Shade Avoidance Responses in Lotus japonicus

Sessile plants must continuously adjust their growth and development to optimize photosynthetic activity under ever-fluctuating light conditions. Among such light responses in plants, one of the best-characterized events is the so-called shade avoidance, for which a low ratio of the red (R):far-red (FR) light intensities is the most prominent stimulus. Such shade avoidance responses enable plants to overtop their neighbors, thereby enhancing fitness and competitiveness in their natural habitat. Considerable progress has been achieved during the last decade in understanding the molecular mechanisms underlying the shade avoidance responses in the model rosette plant, Arabidopsis thaliana. We characterize here the fundamental aspects of the shade avoidance responses in the model legume, Lotus japonicus, based on the fact that its phyllotaxis (or morphological architecture) is quite different from that of A. thaliana. It was found that L. japonicus displays the characteristic shade avoidance syndrome (SAS) under defined laboratory conditions (a low R:FR ratio, low light intensity, and low blue light intensity) that mimic the natural canopy. In particular, the outgrowth of axillary buds (i.e., both aerial and cotyledonary shoot branching) was severely inhibited in L. japonicus grown in the shade. These results are discussed with special emphasis on the unique aspects of SAS observed with this legume.     

Phytochrome D acts in the shade-avoidance syndrome in Arabidopsis by controlling elongation growth and flowering time

Shade avoidance in higher plants is regulated by the action of multiple phytochrome (phy) species that detect changes in the red/far-red ratio (R/FR) of incident light and initiate a redirection of growth and an acceleration of flowering. The phyB mutant of Arabidopsis is constitutively elongated and early flowering and displays attenuated responses to both reduced R/FR and end-of-day far-red light, conditions that induce strong shade-avoidance reactions in wild-type plants. This indicates that phyB plays an important role in the control of shade avoidance. In Arabidopsis phyB and phyD are the products of a recently duplicated gene and share approximately 80% identity. We investigated the role played by phyD in shade avoidance by analyzing the responses of phyD-deficient mutants. Compared with the monogenic phyB mutant, the phyB-phyD double mutant flowers early and has a smaller leaf area, phenotypes that are characteristic of shade avoidance. Furthermore, compared with the monogenic phyB mutant, the phyB-phyD double mutant shows a more attenuated response to a reduced R/FR for these responses. Compared with the phyA-phyB double mutant, the phyA-phyB-phyD triple mutant has elongated petioles and displays an enhanced elongation of internodes in response to end-of-day far-red light. These characteristics indicate that phyD acts in the shade-avoidance syndrome by controlling flowering time and leaf area and that phyC and/or phyE also play a role.     

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.