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.     

光强对烟草幼苗形态和生理指标的影响

通过白纱布遮荫模拟不同光生境条件(透光率分别为100%、68.2%、35.4%和16.7%),研究了光强因子对烟草幼苗形态和生理指标的影响.结果表明:随相对光强的减弱,幼苗高度增加,茎粗、干鲜比、叶片厚度和单位叶面积质量均呈降低趋势,幼苗干物质积累减少,但其对叶数的影响不大.弱光条件下,叶片自由水、叶绿素、总氮和蛋白质含量增加,束缚水含量降低,叶绿素a/b值减小,转化酶活性降低;烟草幼苗根系相对不发达,根冠比和根生物量减小,根系活力降低.表明弱光条件不利于培育烟草壮苗,生产中应尽可能改善苗床的光照条件.     

不同光照质量和刈割强度对小糠草无性繁殖特性的影响

本实验于1996年在美国德克萨斯理工大学草原,野生动物和渔类系的温室和实验室中进行,实验材料均为盆栽糠草,实验设计为在自然光照下（R／FR＝1．1）,以不同留茬高度（10,15,20cm和不刈割）作为不同刈割强度,用不同红光／远红光比例（R／FR＝1．3和0．7）处理不同刈割强度的小糠草,观测不同处理对小糠草无性繁殖特性的影响,结果表明,小糠草净增蘖数随刈割强度增加而减少,除不刈割处理外,光照质量变化（R／FR的比例）对分蘖性能影响不大,所有刈割强度下都不存在分蘖的补偿性生长效应,但幼叶伸长速度随刈割强度的增大而加快,可以认为小糠草应适度利用,强度利用或不利均不利于该草生长发育。     

Light from below matters: Quantifying the consequences of responses to far‐red light reflected upwards for plant performance in heterogeneous canopies

In vegetation stands, plants receive red to far‐red ratio (R:FR) signals of varying strength from all directions. However, plant responses to variations in R:FR reflected from below have been largely ignored despite their potential consequences for plant performance. Using a heterogeneous rose canopy, which consists of bent shoots down in the canopy and vertically growing upright shoots, we quantified upward far‐red reflection by bent shoots and its consequences for upright shoot architecture. With a three‐dimensional plant model, we assessed consequences of responses to R:FR from below for plant photosynthesis. Bent shoots reflected substantially more far‐red than red light, causing reduced R:FR in light reflected upwards. Leaf inclination angles increased in upright shoots which received low R:FR reflected from below. The increased leaf angle led to an increase in simulated plant photosynthesis only when this low R:FR was reflected off their own bent shoots and not when it reflected off neighbour bent shoots. We conclude that plant response to R:FR from below is an under‐explored phenomenon which may have contrasting consequences for plant performance depending on the type of vegetation or crop system. The responses are beneficial for performance only when R:FR is reflected by lower foliage of the same plants.     

Regulation by light of chlorophyll synthesis in the cotyledons of Scots pine (Pinus sylvestris) seedlings

Seedlings of gymnosperms, unlike angiosperms, synthesize chlorophyll(ide) (Chl) in darkness (D). In Scots pine cotyledons ( Pinus sylvestris L.) Chl accumulation ceases in D at a low level but Chl accumulation is strongly increased by light, red light (R) being more effective than blue light (B), whereas in Pinus maritima Chi synthesis is almost light-independent. In Scots pine the capacity to form Chl can be increased by R pulses, fully reversible by far-red light, demonstrating the involvement of phytochrome. However, when B- or R–grown seedlings were transferred to D, Chl accumulation stopped immediately irrespective of the level of P_fr (far-red light absorbing form of phytochrome), indicating that the conversion of protochlorophyllide (PChl) is light-dependent. Dose response curves in R and B and simultaneous irradiation with R and B show that R and B are perceived by separate photoreceptors. The immunodetected NADPH-dependent protochlorophyllide oxidoreductase (POR, EC 1.6.99.1), assumed to regulate light-dependent Chl synthesis in angiosperms, is not correlated with the capacity of gymnosperm Chi accumulation in darkness. While two FOR bands could be separated in extracts from dark grown material (38 and 36 kDa) of Pinus sylvestris and P. maritima , only the 38 kDa band disappeared consistently in the light. However. the significance of the more light resistant 36 kDa band for chlorophyll synthesis remains unclear as well.     

水分和光照对厚皮甜瓜苗期植株生理生态特性的影响

以厚皮甜瓜‘西博洛托’为试材,研究了温室内育苗基质含水量和光照强度对厚皮甜瓜苗期植株生长发育和生理生态特性的影响.结果表明:不同基质含水量和光照强度对厚皮甜瓜苗期植株生长发育影响显著,随基质含水量的降低,苗期植株叶绿素a、叶绿素b和总叶绿素含量升高,叶片相对含水量和比叶面积下降,基质含水量为60%～80%时,幼苗净光合速率最高;随着光照强度的减弱,幼苗的叶绿素a、叶绿素b、总叶绿素含量以及叶片相对含水量、比叶面积和胞间CO2浓度升高,叶绿素a/b和净光合速率下降.在不同处理中,以80%基质含水量 100%温室光照处理的幼苗生长最健壮,壮苗指数、净光合速率等指标最高.     

中国木本植物幼苗生长对光照强度的响应

光照是影响植物幼苗生长的重要的环境因子,定量化研究光照对木本植物不同生活型幼苗生长的影响具有重要意义。系统收集了有关光照对我国木本植物幼苗生长影响的国内外文献,采用Meta分析的方法对幼苗的生长效应进行评估。研究结果表明:(1)与全光照下幼苗生长相比,由于遮阴处理对幼苗基径增长量的抑制作用强于对株高增长量的作用,幼苗出现株高基面积比增高的形态特征变化;(2)与灌木相比,乔木树种幼苗对不同光照强度的响应更加敏感;(3)在小于20%和20%—40%全光照条件下,落叶阔叶和常绿阔叶树种幼苗基径均受到显著抑制,而常绿针叶树种的基径并没有受到遮阴的明显抑制;(4)落叶阔叶和常绿阔叶树种的株高基面积比均显著高于全光照,而常绿针叶树种的株高基面积比与全光照相比差异较小。我们的这一研究将有助于更好的理解木本植物幼苗生长对光照的响应机理。     

Differential genetic variation in adaptive strategies to a common environmental signal in Arabidopsis accessions: phytochrome-mediated shade avoidance

Shade avoidance is a syndrome of plastic responses to light signals encountered in crowded plant communities and is a crucial component of competitive strategy in higher plants. The responses are mediated via signal perception by specific members of the phytochrome family of photoreceptors, which detect the relative proportions of red (R) and far‐red (FR) radiation within dense communities. We analysed two aspects of shade avoidance, the acceleration of flowering and the enhancement of elongation growth, displayed by more than 100 accessions of Arabidopsis thaliana (Heyn.) in response to FR‐proximity signals. Both traits showed wide variation between accessions, which was unrelated to the latitude of the location of original collection. Flowering acceleration is a major feature of shade avoidance in rosette plants such as Arabidopsis, and most accessions showed dramatic responses, but several were identified as being recalcitrant to the proximity signal. These accessions are likely to be informative in the analysis of quantitative variation in shade avoidance. Hypocotyl elongation, treated here as an indicator of elongation growth responses, also varied widely amongst accessions. The variations in flowering acceleration and elongation were not correlated, indicating that microevolution in the downstream pathways from signal perception has occurred separately.     

Root:shoot ratio in developing seedlings: How seedlings change their allocation in response to seed mass and ambient nutrient supply

Root:shoot (R:S) biomass partitioning is one of the keys to the plants' ability to compensate for limiting resources in the environment and thus to survive and succeed in competition. In adult plants, it can vary in response to many factors, such as nutrient availability in the soil or reserves in the roots from the previous season. The question remains whether, at the interspecific level, reserves in seeds can affect seedlings' R:S ratio in a similar way. Proper allocation to resource‐acquiring organs is enormously important for seedlings and is likely to determine their survival and further success. Therefore, we investigated the effect of seed mass on seedling R:S biomass partitioning and its interaction with nutrient supply in the substrate. We measured seedling biomass partitioning under two different nutrient treatments after 2, 4, 6, and 12 weeks for seventeen species differing in seed mass and covering. We used phylogenetically informed analysis to determine the independent influence of seed mass on seedling biomass partitioning. We found consistently lower R:S ratios in seedlings with higher seed mass. Expectedly, R:S was also lower with higher substrate nutrient supply, but substrate nutrient supply had a bigger effect on R:S ratio for species with higher seed mass. These findings point to the importance of seed reserves for the usage of soil resources. Generally, R:S ratio decreased over time and, similarly to the effect of substrate nutrients, R:S ratio decreased faster for large‐seeded species. We show that the seed mass determines the allocation patterns into new resource‐acquiring organs during seedling development. Large‐seeded species are more flexible in soil nutrient use. It is likely that faster development of shoots provides large‐seeded species with the key advantage in asymmetric above‐ground competition, and that this could constitute one of the selective factors for optimum seed mass.     

Growth and ethylene evolution by shade and sun ecotypes of Stellaria longipes in response to varied light quality and irradiance

Plants growing in the shade receive both low light irradiance and light enriched in far red (FR) (i.e., light with a low red (R) to FR ratio). In an attempt to uncouple the R/FR ratio effects from light irradiance effects, we utilized Stellaria longipes because this species has two distinct natural population ecotypes, alpine (dwarf) and prairie (tall). The alpine population occupies the open, sun habitat. By contrast, the prairie population grows in the shade of other plants. Both 'sun' and 'shade' ecotypes responded with increased stem elongation responses under low irradiance, relative to growth under 'normal' irradiance, and this increased growth was proportionally similar. However, only the shade ecotype had increased shoot elongation in response to a low R/FR ratio. By contrast, the sun ecotype showed increased stem elongation in response to increasing R/FR ratio. Varying the R/FR ratios had no significant effect on ethylene evolution in either sun or shade ecotype. Under low irradiance, only the sun ecotype showed a significantly changed (decreased) ethylene evolution. We conclude that R/FR ratio and irradiance both regulate growth, and that irradiance can also influence ethylene evolution of the sun ecotype. By contrast, R/FR ratio and irradiance, while having profound influences on growth of the shade ecotype, do not appear to regulate these growth changes via effects on ethylene production.     

伞形科两种植物幼苗生长对光照强度的可塑性响应

将明党参（Changium smyrnioides Wolff）和峨参（Anthriscus sylvestris Hoffm．）幼苗置于按全光照百分率为100％（S100）、65％（S65）和25％（S25）的人工控制光环境下处理。结果表明,光照强度显著影响明党参和峨参幼苗的生长：明党参在S6,生长最好,株高、冠幅、叶长、叶宽、地上地下生物量等均达到最大值,且S65的明党参生物量是S100的2倍、S25的3倍左右;而峨参地上部生长随着光照强度的减弱而增强,S25生长最好,但地下部生长则在S65生长最好,表现出与明党参同样的响应规律。明党参在不同光照强度下生长速率差异显著,S65显著高于其余两个处理;峨参虽然也在S65生长速率最大,但在不同光照强度间无差异。明党参生长速率明显小于峨参,总生物量约为峨参的1／3—1／8,相对于峨参而言明党参是一个在幼苗阶段生长缓慢的物种。     

不同比例红蓝光对番茄幼苗生长和光合作用的影响

为了探索番茄幼苗生长发育对红蓝组合光的响应机制,本试验采用发光二极管(LED)精量调制光源,以番茄品种‘SV0313TG’为试材,设红光(R)、蓝光(B)和红蓝组合光(9R1B、6R1B、3R1B、1R1B、1R3B)7个处理,以白光为对照,研究不同比例红蓝光质对番茄幼苗生长、光合色素含量、光合特性、叶绿素荧光参数及根系活力的影响.结果表明:不同比例红蓝光质处理对番茄幼苗生长的影响具有明显差异.红光显著促进幼苗株高增加,比叶面积增大,胞间CO₂浓度提高,但PSⅡ最大光化学效率(F_v/F_m)、PSⅡ实际光化学效率(Ф_PSⅡ)降低,根系生长受阻,根系活力下降,壮苗指数降低;蓝光下幼苗生长受到明显抑制,叶绿素含量降低,但叶绿素a/b 值升高;红蓝组合光有利于番茄幼苗的生长发育,3R1B处理下植株干物质量、叶绿素含量和光合性能均显著提高,幼苗生长健壮,壮苗指数最大.综上,红蓝组合光能够增加番茄幼苗叶片光合色素含量,提高光合效率,促进植株生长,尤以3R1B处理最佳.     

Cessation of tillering in spring wheat in relation to light interception and red : far-red ratio

BACKGROUND AND AIMS: The production of axillary shoots (tillering) in spring wheat (Triticum aestivum) depends on intraspecific competition. The mechanisms that underlie this competition are complex, but light within the wheat canopy plays a key role. The main objectives of this paper are to analyse the effects of plant population density and shade on tillering dynamics of spring wheat, to assess the canopy conditions quantitatively at the time of tillering cessation, and to analyse the relationship between the tiller bud and the leaf on the same phytomer. METHODS: Spring wheat plants were grown at three plant population densities and under two light regimes (25 % and 100 % light). Tiller appearance, fraction of the light intercepted, and red : far-red ratio at soil level were recorded. On six sampling dates the growth status of axillary buds was analysed. KEY RESULTS: Tillering ceased earlier at high population densities and ceased earlier in the shade than in full sunlight. At cessation of tillering, both the fraction of light intercepted and the red : far-red ratio at soil level were similar in all treatments. Leaves on the same phytomer of buds that grew out showed more leaf mass per unit area than those on the same phytomer of buds that remained dormant. CONCLUSIONS: Tillering ceases at specific light conditions within the wheat canopy, independent of population density, and to a lesser extent independent of light intensity. It is suggested that cessation of tillering is induced when the fraction of PAR intercepted by the canopy exceeds a specific threshold (0.40-0.45) and red : far-red ratio drops below 0.35-0.40.     

The ratio of leaf to total photosynthetic area influences shade survival and plastic response to light of green-stemmed leguminous shrub seedlings

Different plant species and organs within a plant differ in their plastic response to light. These responses influence their performance and survival in relation to the light environment, which may range from full sunlight to deep shade. Plasticity, especially with regard to physiological features, is linked to a greater capacity to exploit high light and is usually low in shade-tolerant species. Among photosynthetic organs, green stems, which represent a large fraction of the total photosynthetic area of certain species, are hypothesized to be less capable of adjustment to light than leaves, because of biomechanical and hydraulic constraints. The response to light by leaves and stems of six species of leguminous, green-stemmed shrubs from dry and high-light environments was studied by growing seedlings in three light environments: deep shade, moderate shade and sun (3, 30 and 100 % of full sunlight, respectively). Survival in deep shade ranged from 2 % in Retama sphaerocarpa to 74 % in Ulex europaeus. Survival was maximal at moderate shade in all species, ranging from 80 to 98 %. The six species differed significantly in their ratio of leaf to total photosynthetic area, which influenced their light response. Survival in deep shade increased significantly with increasing ratio of leaf to total photosynthetic area, and decreased with increasing plasticity in net photosynthesis and dark respiration. Responses to light differed between stems and leaves within each species. Mean phenotypic plasticity for the variables leaf or stem specific mass, chlorophyll content, chlorophyll a/b ratio, and carotenoid to chlorophyll ratio of leaves, was inversely related to that of stems. Although mean plasticity of stems increased with the ratio of leaf to total photosynthetic area, the mean plasticity of leaves decreased. Shrubs with green stems and a low ratio of leaf to total photosynthetic area are expected to be restricted to well-lit habitats, at least during the seedling stage, owing to their inefficient light capture and the low plasticity of their stems.     

Mutants of phospholipase A (pPLA‐I) have a red light and auxin phenotype

pPLA‐I is the evolutionarily oldest patatin‐related phospholipase A (pPLA) in plants, which have previously been implicated to function in auxin and defence signalling. Molecular and physiological analysis of two allelic null mutants for pPLA‐I [ppla‐I‐1 in Wassilewskija (Ws) and ppla‐I‐3 in Columbia (Col) ] revealed pPLA‐I functions in auxin and light signalling. The enzyme is localized in the cytosol and to membranes. After auxin application expression of early auxin‐induced genes is significantly slower compared with wild type and both alleles show a slower gravitropic response of hypocotyls, indicating compromised auxin signalling. Additionally, phytochrome‐modulated responses like abrogation of gravitropism, enhancement of phototropism and growth in far red‐enriched light are decreased in both alleles. While early flowering, root coils and delayed phototropism are only observed in the Ws mutant devoid of phyD, the light‐related phenotypes observed in both alleles point to an involvement of pPLA‐I in phytochrome signalling.