Leaf movement in Calathea lutea (Marantaceae)

Summary Calathea lutea is a broad-leaved, secondary successional plant which shows complex leaf movements involving both elevation and folding of the leaf surface about the pulvinus. In the plants studied, mean leaf elevation increased from approximately 34 degrees in the early morning to 70 degrees at noon while the angle of leaf folding increased from 13 degrees to 50 degrees over the same time period. During the period from early morning to noon, these movements resulted in a significant decrease in the cosine of the angle of incidence, a measure of the direct solar radiation intercepted. The observed changes in elevational angle significantly reduce the cosine of angle of incidence while folding does not significantly reduce the fraction of direct solar radiation intercepted during the period of direct exposure of the leaf surface to the solar beam. Since elevational changes seem to account for the reduction in exposure to direct solar radiation, the role of folding remains unclear.     

Harnessing Genetic Variation in Leaf Angle to Increase Productivity of Sorghum bicolor

The efficiency with which a plant intercepts solar radiation is determined primarily by its architecture. Understanding the genetic regulation of plant architecture and how changes in architecture affect performance can be used to improve plant productivity. Leaf inclination angle, the angle at which a leaf emerges with respect to the stem, is a feature of plant architecture that influences how a plant canopy intercepts solar radiation. Here we identify extensive genetic variation for leaf inclination angle in the crop plant Sorghum bicolor, a C4 grass species used for the production of grain, forage, and bioenergy. Multiple genetic loci that regulate leaf inclination angle were identified in recombinant inbred line populations of grain and bioenergy sorghum. Alleles of sorghum dwarf-3, a gene encoding a P-glycoprotein involved in polar auxin transport, are shown to change leaf inclination angle by up to 34° (0.59 rad). The impact of heritable variation in leaf inclination angle on light interception in sorghum canopies was assessed using functional-structural plant models and field experiments. Smaller leaf inclination angles caused solar radiation to penetrate deeper into the canopy, and the resulting redistribution of light is predicted to increase the biomass yield potential of bioenergy sorghum by at least 3%. These results show that sorghum leaf angle is a heritable trait regulated by multiple loci and that genetic variation in leaf angle can be used to modify plant architecture to improve sorghum crop performance.     

Leaf orientation and light interception by juvenile Pseudopanax crassifolius(Cunn.) C. Koch in a partially shaded forest environment

Leaf orientations and light environments were recorded for 40 juvenile Pseudopanax crassifolius trees growing in New Zealand in a partially shaded, secondary forest environment. Efficiencies of interception of diffuse and direct light by the observed leaf arrangments were calculated relative to those of three hypothetical leaf arrangements. Canopy gaps above the study plants were unevenly distributed with respect to azimuth and elevation above the horizon. Our results indicate that photosynthetically active radiation (PAR) received from the sides is more important than that received from directly above. In 33 of the plants leaf orientation was found to be significantly clustered towards one azimuth. The mean azimuth and the mean angle of declination were different for each plant. Leaves were steeply declined, and oriented towards the largest canopy gap at each site. Steep leaf angles reduced interception of direct and diffuse PAR when compared to interception by plant with a hypothetical horizontal leaf arrangement. When compared to a hypothetical arrangement with steep leaf declination and a uniform azimuth distribution, the observed leaf arrangement increased the efficiency of interception of diffuse PAR, but had a variable effect on the interception of direct PAR. Results indicate that the developing leaves of juvenile P. crassifolius orient towards the strongest sources of diffuse light, regardless of their value as a source of direct light. By maximising diffuse light interception while reducing direct light interception, leaf orientation may be a partial determinant of the types of habitats exploited by this species. This study emphasises the importance of considering diffuse light interception for plants growing in partially shaded environments.     

Solar tracking response to drought in a desert annual

Summary The responses to drought of the solar tracking winter annualLupinus arizonicus (Wats.) were examined under field and laboratory growth regimes. Under drought conditions tracking movements were maintained until the plant reached the wilting point. The leaves and leaflets were observed to cup in response to decreases in the xylem water potential. This resulted in a negative, but linear relationship between the cosine of the angle of incidence of the direct solar beam upon the leaf and water potential. Leaf conductance was also significantly related to xylem water potential in natural populations. Leaf and leaflet cupping seem to be responding directly to changing plant water status, resulting in reduced solar radiation load to drought stressed plants. It is hypothesized that this property may have adaptive consequences in relation to the water relations, energy budget, and carbon balance of the plant.     

AXIAL ROTATION OF ERYTHRINA HERBACEA LEAFLETS

The leaflets of Erythrina herbacea, a leguminous plant native to subtropical North America, undergo daily movements. During the night, leaflets are oriented with their tips pointing downwards and show no rotation around the midrib of each leaflet. In the morning, leaflets rise, passing through a horizontal orientation and, somewhat before noon, reach a maximum average midrib elevation of approximately sixty degrees above the horizontal plane. Simultaneously, each leaflet rotates about the midrib by an average angle of forty to fifty degrees. These leaflet movements result in an initial increase in the relative leaflet surface area presented to the sun during the morning, a decrease in interception of direct solar radiation around noon, followed by a second maximum in light interception in early afternoon.     

Environmental control of the prostrate growth form in two high arctic grasses

In the High Arctic, leaf angles of graminoids are lower than would be expected to optimize absorption of incident solar radiation. Inflorescence, leaf, and culm angle of two prostrate grasses, Phippsia algida and Puccinellia vaginata , were measured in a variety of plant communities with differing microtopographic and microenvironmental conditions (surface relief, soil stability, soil moisture, and soil and air temperature). Growth form was found to be controlled primarily by environmental factors. Both species exhibited a plastic response to an amelioration of physical factors in situ. Leaf angles of both species were higher under conditions of increased soil moisture and temperature, and decreased vapor pressure deficit and windspeed. Adaptation to reduce the severity of the environment apparently holds a greater selective advantage than adaptation to maximize leaf orientation to a low sun angle.     

Modeling the light interception and carbon gain of individual fluttering aspen (<Emphasis Type="Italic">Populus tremuloides</Emphasis> Michx) leaves

Poplars (Populus spp.) have a particular petiole morphology that enhances leaf flutter even in light winds. Previous studies have shown that this trait enhances whole canopy carbon gain through changes in the temporal dynamics and spatial distribution of light in the lower canopy. However, less is known about the effects of flutter for leaves at the top of the canopy ("sun leaves"). A computer simulation model was developed that uses latitude, time of day, day of year, azimuth and a slope component, which was varied at a 3 Hz frequency over an arc of rotation to create the flutter motion, and generate data on light interception for both surfaces of a fixed or fluttering leaf. The light files generated (10 Hz) were input into a dynamic model of photosynthesis to estimate the carbon gain for both fluttering and fixed leaves. As compared to leaves fixed at various angles and azimuths, fluttering leaves had a more uniform light interception. Depending on their angle and azimuth, fixed leaves may not always be intercepting high photon flux density (PFD) even when exposed to full sun. Leaf flutter continuously randomizes leaf angles creating uniform PFD inputs for photosynthetic reactions regardless of variation in leaf orientation and solar position. These effects on light interception could have positive impacts on carbon gain for leaves at the top of the canopy.     

Solar radiation transmission in and around canopy gaps in an uneven-aged <Emphasis Type="Italic">Nothofagus betuloides</Emphasis> forest

The transmission of direct, diffuse and global solar radiation in and around canopy gaps occurring in an uneven-aged, evergreen Nothofagus betuloides forest during the growing season (October 2006–March 2007) was estimated by means of hemispherical photographs. The transmission of solar radiation into the forest was affected not only by a high level of horizontal and vertical heterogeneity of the forest canopy, but also by low angles of the sun’s path. The below-canopy direct solar radiation appeared to be variable in space and time. On average, the highest amount of transmitted direct solar radiation was estimated below the undisturbed canopy at the southeast of the gap centre. The transmitted diffuse and global solar radiation above the forest floor exhibited lower variability and, on average, both were higher at the centre of the canopy gaps. Canopy structure and stand parameters were also measured to explain the variation in the below-canopy solar radiation in the forest. The model that best fit the transmitted below-canopy direct solar radiation was a growth model, using plant area index with an ellipsoidal angle distribution as the independent variable (R ² = 0.263). Both diffuse and global solar radiation were very sensitive to canopy openness, and for both cases a quadratic model provided the best fit for these data (R ² = 0.963 and 0.833, respectively). As much as 75% and 73% of the variation in the diffuse and global solar radiation, respectively, were explained by a combination of stand parameters, namely basal area, crown projection, crown volume, stem volume, and average equivalent crown radius.     

A tropical epiphytic orchid uses a low‐light interception strategy in a spatially heterogeneous light environment

Light is considered a non‐limiting factor for vascular epiphytes. Nevertheless, an epiphyte's access to light may be limited by phorophyte shading and the spatio‐temporal environmental patchiness characteristic of epiphytic habitats. We assessed the extent to which potential light interception in Rodriguezia granadensis, an epiphytic orchid, is determined by individual factors (plant size traits and leaf traits), or environmental heterogeneity (light patchiness) within the crown of the phorophyte, or both. We studied 104 adult plants growing on Psidium guajava trees in two habitats with contrasting canopy cover: a dry tropical forest edge, and isolated trees in a pasture. We recorded the number of leaves and the leaf area, the leaf position angles, and the potential exposure of the leaf surface to direct irradiance (silhouette area of the leaf blade), and the potential irradiance incident on each plant. We found the epiphytes experience a highly heterogeneous light environment in the crowns of P. guajava. Nonetheless, R. granadensis plants displayed a common light interception strategy typical of low‐light environments, resembling terrestrial, forest understory plants. Potential exposure of the total leaf surface to direct irradiance correlated positively with plant size and within‐plant variation in leaf orientation. In many‐leaved individuals, within‐plant variation in leaf angles produced complementary leaf positions that enhanced potential light interception. This light interception strategy suggests that, in contrast to current wisdom, enhancing light capture is important for vascular epiphytes in canopies with high spatio‐temporal heterogeneity in light environments.     

THE ROLE OF PHYLLOTACTIC PATTERN AS A “DEVELOPMENTAL CONSTRAINT” ON THE INTERCEPTION OF LIGHT BY LEAF SURFACES

Computer simulations of model plants are used to assess the influence of leaf shape, size, and pattern of arrangement (= phyllotaxy) on the direct solar radiation intercepted by leaf surfaces. Changes in phyllotaxy significantly influence light interception (and, by inference, net assimilation rate) for rosette growth habits. However, changes in leaf shape and orientation and in stem length can compensate for the negative effects of leaf overlap produced by phyllotactic patterns. Phyllotaxy is viewed as a developmental limiting factor in photobiology that may necessitate compensatory changes in other morphological features not directly controlled by patterns of leaf initiation. This distinguishes it from functioning as a “developmental constraint” sensu stricto and may provide a paradigm for other features in plant evolution.     

The effects of nitrogen, light and water availability on tropic leaf movements in soybean (Glycine max)

Abstract. Rapid, tropic leaf movements and photo-synthetic responses of the heliotropic plant, soybean, Glycine max cv. Cumberland, grown under two different nitrogen, three different light and two different water treatments were examined. Measurements of leaf orientation during midday periods outdoors, and tropic reorientation of leaflets in response to vertical illumination indoors, revealed a positive, linear relationship between leaf water potential and the cosine of the angle of incidence between the leaf and the direct beam of the excitation light. This relationship was altered by nitrogen availability, such that a lower cosine of incidence (lower leaf irradiance) for a given leaf water potential was measured for plants grown under low nitrogen compared to those grown under high nitrogen. Additionally, plants grown under low nitrogen and low water availability showed more rapid rates of leaf movement compared to plants receiving high levels of these resources. Light regime during growth had no effect on the relationship between the cosine of incidence and leaf water potential. Reduced water and nitrogen availabilities during growth resulted in lower photosaturated rates of photosynthesis and stomatal conductance, as well as alterations in the relationship between these parameters. Thus, higher values for the ratio of intercellular CO₂/ambient CO₂ were measured for low-N grown plants (higher nitrogen use efficiencies) and lower values of this ratio for water stressed plants (higher water use efficiencies). The results show that environmental growth conditions other than water availability have the potential to modify leaf orientation responses to vectorial light in heliotropic legumes such as soybean. This has implications for the potential of heliotropic movements to minimize environmental stress-induced damage to the photosynthetic apparatus, and to modulate leaf-level resource use efficiencies.     

CHANGES IN LEAF CHARACTERISTICS OF SPECIES ALONG ELEVATIONAL GRADIENTS IN THE WASATCH FRONT,UTAH

Leaf absorptances and leaf angles were measured on 159 of the most common vascular plant species along a precipitation/elevation transect in the Wasatch Mountains. At the community level, leaf angles were steepest in the driest saltbush community sites, became progressively more horizontal in wetter communities, and reached a minimum in the midelevation mountain brush communities. At higher elevation plant communities, where growth is delayed until summer months because of heavy snowfall, leaf angles were again steeper. Leaf absorptances among all life forms were uniformly high along the entire transect until the driest saltbush communities where a number of species exhibited reduced leaf absorptances. The results are discussed within the context of mechanisms for reducing solar radiation absorption along an aridity transect.     

Exploring the spatial distribution of light interception and photosynthesis of canopies by means of a functional-structural plant model

Background and Aims

At present most process-based models and the majority of three-dimensional models include simplifications of plant architecture that can compromise the accuracy of light interception simulations and, accordingly, canopy photosynthesis. The aim of this paper is to analyse canopy heterogeneity of an explicitly described tomato canopy in relation to temporal dynamics of horizontal and vertical light distribution and photosynthesis under direct- and diffuse-light conditions.

Methods

Detailed measurements of canopy architecture, light interception and leaf photosynthesis were carried out on a tomato crop. These data were used for the development and calibration of a functional–structural tomato model. The model consisted of an architectural static virtual plant coupled with a nested radiosity model for light calculations and a leaf photosynthesis module. Different scenarios of horizontal and vertical distribution of light interception, incident light and photosynthesis were investigated under diffuse and direct light conditions.

Key Results

Simulated light interception showed a good correspondence to the measured values. Explicitly described leaf angles resulted in higher light interception in the middle of the plant canopy compared with fixed and ellipsoidal leaf-angle distribution models, although the total light interception remained the same. The fraction of light intercepted at a north–south orientation of rows differed from east–west orientation by 10 % on winter and 23 % on summer days. The horizontal distribution of photosynthesis differed significantly between the top, middle and lower canopy layer. Taking into account the vertical variation of leaf photosynthetic parameters in the canopy, led to approx. 8 % increase on simulated canopy photosynthesis.

Conclusions

Leaf angles of heterogeneous canopies should be explicitly described as they have a big impact both on light distribution and photosynthesis. Especially, the vertical variation of photosynthesis in canopy is such that the experimental approach of photosynthesis measurements for model parameterization should be revised.     

利用多光谱扫描仪测定空心莲子草冠层光谱的影响因素

【背景】太阳光的入射辐射是影响多光谱扫描仪测定植物冠层光谱反射率的主要因素。本研究旨在探讨不同测定条件对空心莲子草冠层光谱特征值的影响,确定应用多光谱扫描仪测定空心莲子草冠层光谱特征的适宜环境条件。【方法】采用多光谱扫描仪,在不同天气条件、仪器探头高度、太阳高度角、叶片湿度等外界因素条件下,测定空心莲子草冠层光谱反射率的变化。【结果】天气条件、仪器探头高度、太阳高度角、叶片湿度4个因素对空心莲子草冠层在绿光区560nm、黄光区660nm和近红外光区810nm3个特征波段处的光谱反射率均有显著影响。【结论与意义】利用多光谱扫描仪测定空心莲子草冠层光谱数据,应选择晴朗无云的天气,仪器探头距离冠层1.3m,适宜的数据采集时间为10：00～14：00,以保障适度的太阳高度角和干燥的叶面,从而保持稳定的光谱测量结果。本研究可为完善多光谱扫描评价技术提供理论依据。     

Growth of Lettuce, Onion, and Red Beet. 1. Growth Analysis, Light Interception, and Radiation Use Efficiency

A field experiment was carried out to analyse the growth oflettuce, onion and red beet in terms of: (a) canopy architecture,radiation interception and absorption; (b) efficiency of conversionof absorbed radiation into biomass; and (c) dry matter partitioning.Growth analysis, total solar radiation interception, PAR interceptionand absorption by the crop canopy, ground cover, maintenancerespiration of onion bulbs and red beet storage roots were measured.Models for different leaf angle distribution and ground coverwere used to simulate light transmission by the crop canopy. The three crops are shown to have contrasting growth patternsfrom both a morphological and a physiological point of view.Lettuce showed very high light interception and growth afterthe early growth stages but, throughout the growth cycle, thisleafy crop showed the lowest radiation use efficiency due tothe respirational cost of the high leaf area. Onion showed alower early relative growth rate than lettuce and red beet.This was due partly to the low light interception per unit leafarea in the later stages of growth and partly to the low initialradiation use efficiency compared with the other two crops.On the other hand, thanks to more uniform distribution of theradiation inside the canopy, to the earlier termination of leafdevelopment and to the very low level of bulb respiration, onionshowed high radiation use efficiency and was able to producea large amount of dry matter. Red beet leaf posture and canopystructure resulted in high light interception and absorption.Its radiation use efficiency was lower than that of onion, partlyperhaps because of the more adverse distribution of the interceptedradiation fluxes within the canopy and partly because of thehigh respiration cost of a continuous dry-matter allocationto the leaves. However, this crop can accumulate a very largeamount of dry matter as leaf blade development and storage rootgrowth can both continue almost indefinitely, providing continuouslyavailable sinks. Ground cover gave a good estimate of the PAR interception onlyat low values of light interception but, in general, it underestimatedPAR interception in all three crops. Ratios between attenuationcoefficients established by considering PAR or total solar radiationand LAI or ground cover were calculated. Lettuce,Lactuca sativa L. var.crispa ; onion,Allium cepa L.; red beet; Beta vulgaris L. var.conditiva ; growth analysis; light interception and absorption; canopy architecture; ground cover; radiation use efficiency; maintenance respiration rate; dry matter distribution     

小麦LAI-2000观测值对辐亮度变化的响应

利用LAI-2000植物冠层分析仪和ASD光谱仪,通过固定点拔节期冬小麦叶面积指数(LAI)观测实验和同步光谱辐亮度实验,测量了晴天条件下,固定点冬小麦从中午至傍晚24个不同时刻的LAI值及对应的辐亮度。继之分析了此段时间内测得的定点冬小麦LAI值分别与对应时刻可见光和近红外谱段的天空光辐亮度、总辐亮度和太阳直射辐亮度值之间的相关性。结果表明,无论波长小于490nm的谱段,还是波长大于490 nm的谱段,LAI与天空光辐亮度、总辐亮度和太阳直射辐亮度都呈负相关,相关系数(R2)高达0.8左右;尤其LAI与天空光辐亮度的负相关性最高,这种相关性随着波长的增大而减小。LAI与各谱段天空光辐亮度的相关性特征可为LAI-2000晴天观测的LAI值归一化修正处理提供一种新思路和技术途径,以消除太阳直射光的影响,从而解决LAI-2000只能在晴天观测的局限性,拓展LAI-2000在晴天观测条件下的适用性。     

紫耳箭竹克隆形态可塑性对典型冠层结构及光环境的响应

在重庆金佛山国家自然保护内,选择了3种典型群落类型(落叶阔叶林、常绿落叶阔叶混交林和常绿阔叶林),使用Hemiview数字植物冠层分析系统量化群落冠层结构和光环境特征,并对林下紫耳箭竹(Fargesia decurvata)的形态可塑性特征进行调查,分析冠层结构和光环境特征改变下紫耳箭竹形态可塑性的差异,并探讨它们之间的相互关系。结果表明:(1)随着落叶阔叶林"常绿落叶阔叶混交林"常绿阔叶林演替的进行,群落的冠层开度降低,叶面积指数增加,平均叶倾角变小,趋于水平化,冠层对光的截获能力提高,林下光照的强度降低(P0.05)。(2)随着光照强度的降低,紫耳箭竹分株矮小化,叶片变窄,生物量积累降低,但通过增大比茎长、叶面积率和比叶面积提高对光的利用效率,并增大分枝角度和比隔长有效适应弱光环境。(3)在光照条件差的常绿阔叶林下,紫耳箭竹降低对地下茎的投资,将较多的生物量用于秆的增高增长和叶片的生长;而在光照条件好的落叶阔叶林环境下,紫耳箭竹降低对枝、叶生物量的分配,则加大对地下茎的投资,可认为是克隆植物对水分资源所表现的一种觅食行为。研究表明,紫耳箭竹种群随着冠层结构的改变发生了明显的可塑性变化,这些可塑性变化是种群对冠层结构和光环境差异的适应性反应的结果,有利于增强种群对异质生境中光资源的获取和利用;群落内部可以通过调控冠层结构的改变协调和控制小径竹种群的发展。     

Leaf angle as a leaf and canopy trait: Rejuvenating its role in ecology with new technology

Life on Earth depends on the conversion of solar energy to chemical energy by plants through photosynthesis. A fundamental challenge in optimizing photosynthesis is to adjust leaf angles to efficiently use the intercepted sunlight under the constraints of heat stress, water loss and competition. Despite the importance of leaf angle, until recently, we have lacked data and frameworks to describe and predict leaf angle dynamics and their impacts on leaves to the globe. We review the role of leaf angle in studies of ecophysiology, ecosystem ecology and earth system science, and highlight the essential yet understudied role of leaf angle as an ecological strategy to regulate plant carbon–water–energy nexus and to bridge leaf, canopy and earth system processes. Using two models, we show that leaf angle variations have significant impacts on not only canopy-scale photosynthesis, energy balance and water use efficiency but also light competition within the forest canopy. New techniques to measure leaf angles are emerging, opening opportunities to understand the rarely-measured intraspecific, interspecific, seasonal and interannual variations of leaf angles and their implications to plant biology and earth system science. We conclude by proposing three directions for future research.     

Comparing the impact of cloudiness on carbon dioxide exchange in a grassland and a maize cropland in northwestern China

Light quantity and quality strongly influence plant growth. However, different ecosystems have different capabilities to assimilate solar radiation. In this study, the effects of cloudiness intensity on the net ecosystem exchange of carbon dioxide (NEE) were compared between an alpine grassland (with lower leaf area index) at A??Rou and an oasis maize cropland (with higher leaf area index) at Yingke, using flux data obtained during the middle of the growing season (July?CAugust) in 2008 and 2009. The results showed that the response of NEE to photosynthetically active radiation (PAR) was more negative (carbon uptake) under cloudy than under clear skies at both sites. The maximum NEE occurred when the clearness index (CI) ranged from 0.4 to 0.7 under cloudy skies. The maximum enhancements were 11.9% for solar elevation angles of 60?C65° in the grassland, and 34.9% for solar elevation angles of 60?C65° and 10.3% for angles of 35?C40° in the maize cropland before the irrigation period. The response of NEE to CI changed slightly with solar elevation angle in the grassland compared to the maize cropland. The results indicate that enhanced NEE under cloudy skies can be attributed to increasingly diffuse PAR and interactions with environmental factors (air temperature and vapor pressure deficit).     

Structural and functional variability within the canopyand its relevance for carbon gain and stress avoidance

The functional variability in leaf angle distribution within the canopy was analysed with respect to regulation of light interception and photoprotection. Leaf orientation strongly determined the maximum photochemical efficiency of PSII (F_v/F_m) during summer: horizontal leaves were highly photoinhibited whereas vertical leaf orientation protected the leaves from severe photoinhibition. The importance of leaf orientation within the canopy was analysed in two Mediterranean macchia species with distinct strategies for drought and photoinhibition avoidance during summer. The semi-deciduous species (Cistus monspeliensis) exhibited strong seasonal but minimal spatial variability in leaf orientation. Reversible structural regulation of light interception by vertical leaf orientation during summer protected the leaves from severe photoinhibition. The evergreen sclerophyll (Quercus coccifera) exhibited high spatial variability in leaf angle distribution throughout the year and was less susceptible to photoinhibition. The importance of both strategies for plant primary production was analysed with a three-dimensional canopy photoinhibition model (CANO-PI). Simulations indicated that high variability in leaf angle orientation in Q. coccifera resulted in whole-plant carbon gain during the summer, which was 94 % of the maximum rate achieved by theoretical homogeneous leaf orientations. The high spatial variability in leaf angle orientation may be an effective compromise between efficient light harvesting and avoidance of excessive radiation in evergreen plants and may optimize annual primary production. Whole plant photosynthesis was strongly reduced by water stress and photoinhibition in C. monspeliensis; however, the simulations indicated that growth-related structural regulation of light interception served as an important protection against photoinhibitory reduction in whole-plant carbon gain.