The influence of axial rotation upon interception of solar radiation by plant leaves

Axial rotation of a plant leaf about the midrib is an important factor which permits effective maximization or minimization of the interception of direct solar radiation. Through use of matrix rotations of coordinate systems, the cosine of the angle of incidence, a measure of the fraction of solar radiation intercepted by the leaf, is described as a function of angles which specify the relative orientation of the leaf to the sun. All of these angles can be measured directly except for the axial rotation angle, which may be calculated from more easily measured quantities. Computer modeling and theoretical calculations demonstrate that relatively large amounts of average axial rotation permit a plant to reduce the cosine of angle of incidence of each flat leaf to zero, resulting in no interception of direct solar radiation, while smaller amounts of average axial rotation allow a plant to intercept as much as 80% of direct radiation, even if leaves are distributed uniformly over 360° of azimuth.     

与变化光环境关联的刺槐光合气体交换和复叶运动

在夏季的晴天阳生叶和阴生叶方位角和方向角的日变化均很小 ,但是叶片的中脉角存在显著的日变化差异 ,这种叶片垂直方向上的显著运动与躲避中午的强光胁迫有关。而在阴天 ,阳生叶和阴生叶方位角、方向角和中脉角的日变化均很小。这说明变化的光环境是引起刺槐复叶运动的主要原因 ,而节律性运动不是刺槐复叶运动的诱导因素。机械固定叶片的控制实验显示 :机械固定叶片的光合速率、气孔导度和蒸腾速率均显著低于对照叶片 ,而机械固定叶片温度明显高于对照叶片 ,这可能是由于机械固定叶片和对照叶片两者在光辐射接受量上的差异造成的。光诱导下的刺槐复叶运动是植株维持最佳生理状态的形态调节表现。     

光诱导下杂种杨无性系叶角和叶绿体的运动

自然光结合人工光源对几个杂种杨无性系叶片进行处理后,对其在两种光环境下３种叶角（方位角、方向角、悬挂角）和叶绿体的运动方式进行了研究。结果发现：各无性系的叶角运动在晴天比比阴天强烈,特别是三倍体无性系最为明显,晴天上午９：００－１１：００,三倍体无性系ＺＨ６和Ｂ３４６通过方位角运动来避免强光胁迫,各无性系主要通过方向角和悬挂角的变化来调节获得最佳太阳辐射,在中午受到强光胁迫时存在明显的“避光运动”。三倍体无性系和某些二倍体无性系在避光运动方式上是不同的,三倍体无性系ＺＨ６和Ｂ３４６采用叶片下垂形式,而二倍体无性系Ｂ１１则采用叶片竖立方式,无论是晴天还是阴天,植物的节律性运动可能参与了叶角运动。受光胁迫时,栅栏组织的叶绿通过不同的运动排列方式来实现对光辐射的最佳吸收。强光胁迫下叶绿体沿径向细胞壁排列,以尽量养活接受过量的太阳辐射,处于弱光条件时,叶绿体则充满足整个细胞,以扩大上太阳辐射的表面积,三倍体无性系对光诱导的敏感程度程度要高于二倍体无性系。     

基于Campbell椭球分布函数的大兴安岭地区主要树种叶倾角分布模拟

叶倾角分布(LAD)直接决定着植被冠层对辐射的截获量,同时对入射太阳辐射的大小与走向也起着决定性作用,是定量遥感中的关键参数.本研究基于Campbell椭球分布函数和迭代方法拟合大兴安岭林区主要树种的LAD,定量分析冠层叶片分层与不分层时模型的拟合情况及不同龄组对LAD的影响.结果表明： 大兴安岭地区6种主要树种的LAD均属于横椭球分布,针叶树的平均叶倾角小于阔叶树;无论对叶片分层处理与否,模型拟合叶倾角的结果与实测结果基本一致;白桦和落叶松的拟合结果与实测结果线性回归的相关系数分别是0.8268、0.8192,均方根误差分别是3.7%、4.3%,说明Campbell模型应用于森林冠层是可靠的;考虑龄组的影响时,虽然分层处理时叶倾角的分布趋势与龄组无关,但幼龄落叶松的平均叶倾角小于成熟落叶松,表明龄组对叶倾角分布取值有正向影响,而对消光系数取值有负向
影响.     

基于地基激光雷达点云的植被表型特征测量

植物表型是基因型与外界环境共同作用的结果。精确测量植物表型对于植物生理特征与功能性状研究具有重要意义。本研究以加拿大一枝黄花(Solidago canadensis)为对象,对20株植株进行3个月室内培养,各月利用地基激光雷达扫描(terrestrial Li DAR scanning,TLS)系统对实验植株进行多站扫描和点云融合,实现对植株生长过程的连续观测。对于扫描获取的离散点云,利用多端点三维坐标重构法获取植株高度,并基于叶片点云的Delaunay三角网重构叶片表面,获得植株的真实高度、叶面积、叶倾角和方位角等结构参量。对比手动测量结果,发现基于点云重构获得的植株高度与真实植株高度对比,二者间相似性的决定系数(R2)为0.991,叶面积、叶倾角、方位角相似性R2分别为0.989、0.949和0.871;基于TLS点云重构法实现了非破坏性的植物表型测量,能够获得高精度的植物表型特征;多时相扫描能精确监测植物生长过程的表型特征变化。     

Estimating leaf inclination and G-function from leveled digital camera photography in broadleaf canopies

The effectiveness of using leveled digital camera for measuring leaf inclination angles was investigated in this study as an inexpensive and convenient alternative to existing approaches. The new method is validated with manual leaf angle measurements for various broadleaf tree species common to hemi-boreal region of Estonia and the tropical forests of Hawai’i Islands. The acquired leaf angle distributions suggest that planophile case might be more appropriate than the commonly assumed spherical as the general approximation of leaf orientation while modeling the radiation transmission through the canopies of (hemi)-boreal broadleaf stands. However, direct leaf inclination measurements should be obtained whenever possible, as there will always exist a large variety of leaf orientation, both among different species and in the space–time domain within a single species. The camera method tested in this study provides a new robust and affordable tool to obtain this information.     

Petiole mechanics, leaf inclination, morphology, and investment in support in relation to light availability in the canopy of Liriodendron tulipifera

To determine the role of leaf mechanical properties in altering foliar inclination angles, and the nutrient and carbon costs of specific foliar angle variation patterns along the canopy, leaf structural and biomechanical characteristics, biomass partitioning into support, and foliar nitrogen and carbon concentrations were studied in the temperate deciduous species Liriodendron tulipifera L., which possesses large leaves on long petioles. We used beam theory to model leaf lamina as a uniform load, and estimated both the lamina and petiole flexural stiffness, which characterizes the resistance to bending of foliar elements at a common load and length. Petiole and lamina vertical inclination angles with respect to horizontal increased with increasing average daily integrated photon flux density (Q_int). Yet, the light effects on lamina inclination angle were primary determined by the petiole inclination angle. Although the petioles and laminas became longer, and the lamina loads increased with increasing Q_int, the flexural stiffness of both lamina and petiole increased to compensate for this, such that the lamina vertical displacement was only weakly related to Q_int. In addition, increases and decreases in the petiole inclination angle with respect to the horizontal effectively reduced the distance of lamina load from the axis of rotation, thereby reducing the bending moments and lamina inclination due to gravity. We demonstrate that large investments, up to 30% of total leaf biomass, in petiole and large veins are necessary to maintain the lamina at a specific position, but also that light has no direct effect on the fractional biomass investment in support. However, we provide evidence that apart from light availability, structural and chemical characteristics of the foliage may also be affected by water stress, magnitude of which scales positively with Q_int.     

Phytohormones signaling and crosstalk regulating leaf angle in rice

Leaf angle is an important agronomic trait in rice (Oryza sativa L.). It affects both the efficiency of sunlight capture and nitrogen reservoirs. The erect leaf phenotype is suited for high-density planting and thus increasing crop yields. Many genes regulate leaf angle by affecting leaf structure, such as the lamina joint, mechanical tissues, and the midrib. Signaling of brassinosteroids (BR), auxin (IAA), and gibberellins (GA) plays important roles in the regulation of lamina joint bending in rice. In addition, the biosynthesis and signaling of BR are known to have dominant effects on leaf angle development. In this review, we summarize the factors and genes associated with the development of leaf angle in rice, outline the regulatory mechanisms based on the signaling of BR, IAA, and GA, and discuss the contribution of crosstalk between BR and IAA or GA in the formation of leaf angle. Promising lines of research in the transgenic engineering of rice leaf angle to increase grain yield are proposed.     

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.     

卫矛属刺果卫矛组的数量分类学分析

通过对1500余份标本的测量和统计,结合文献和野外考察,选取了30个形态学性状,对卫矛属（Euonymus）刺果卫矛组（sect. Echinococcus）13个种进行了数量分类学分析。聚类分析结果表明,美国卫矛（E.americanus）和陈谋卫矛（E.chenmoui）的亲缘关系最近,不支持Blakelock将本组划分为两个系的观点。本组中系统位置有争议的物种：厚叶卫矛（E.hemsleyanus）与紫刺卫矛（E.angustatus）、棘刺卫矛（E.echinatus）和无柄卫矛（E.subsessilis）、刺猬卫矛（E.hystrix）和蒙自卫矛（E.mengtseanus）、刺果卫矛（E.acanthocarpus）和腾冲卫矛（E.tengyuehensis）,在聚类图中显示出较远的亲缘关系,通过方差分析对这些物种间有较明显差异的性状进行进一步分析。结果表明,这些物种在叶尖角度、叶长、侧脉与主脉的夹角、花序梗长等性状上均具有显著差异。分析结果支持将蒙自卫矛与腾冲卫矛排除出刺果卫矛组,支持厚叶卫矛和紫刺卫矛作为独立的种,由于在交叉分布地域没有足够的研究资料,棘刺卫矛和无柄卫矛的系统位置有待进一步确认。因子分析结果表明,本实验所用性状间的独立性较强,叶长、侧脉与主脉的夹角、花序分支数、果实带刺的长度等18个性状在本组中具有重要的分类学意义。     

Leaf Biomechanics and Biomass Investment in Support in Relation to Long-Term Irradiance in Fagus

Abstract: We investigated biomass investment in support and assimilative leaf biomass in Fagus orientalis Lipsky and F. sylvatica L., and foliar biomechanical characteristics in F. orientalis to gain mechanistic insight into the determinants of leaf inclination in Fagus along the canopy light gradient. Because the leaf laminas of Fagus are elliptical, with petioles comprising only ca. 8 % of total leaf length, a leaf was approximated as a continuous sine load. Lamina load increased with increasing seasonal integrated quantum flux density in the canopy (Q_int), but leaf length was independent of irradiance. Despite greater load, leaf deflection under leaf own weight was lower for leaves at higher Q_int, indicating that foliage flexural stiffness (EI), that is a variable characterizing the resistance of beam‐like structures to bending, scaled positively with irradiance. The components of EI ‐ the leaf apparent Young's modulus of elasticity (E), which is a measure of leaf material properties, and lamina second moment of area (I), which characterizes the distribution of mass around the axis of bending ‐ were also related to irradiance, with E decreasing, but I increasing with Q_int. The positive scaling of I with Q_int was associated with increases in leaf thickness and, in particular, with increases in the degree of leaf rolling, allowing the distribution of leaf mass further away from the neutral axis. Decreases in E were correlated with decreased leaf biomass investments in the midrib at higher irradiance. Both lamina and midrib nitrogen concentrations decreased with increasing Q_int, suggesting that foliage dry mass based physiological activity was lower at higher irradiance, possibly because of an interaction of Q_int with water stress in the canopy. Given that the veins also provide a pathway for water and nutrient transport to the leaf cells, as well as for carbon translocation from the leaf, lower leaf physiological activity in high light may provide an explanation for the lower biomass investment in major veins in high light. We conclude that foliage biomechanical characteristics and leaf inclination in the canopy are significantly affected by irradiance, and that the light effects may be modified by the reverse correlation between light and water availabilities in the canopy.     

树木叶片面积与叶柄长、节间长和叶倾角的关系初探

树木的叶片面积与叶柄长、节间长之间有着明显的内在联系．本文分析研究了这个联系,在适当的理想假设下通过数学建模得到了反映树木叶片面积与叶柄长、节间长和叶倾角之间的关系模型．接着,文章对模型的不合理之处进行了分析与改进,并得到叶片面积与节间长、心茎距之间的关系表达式．最后,本文利用分别采集到的13种互生和对生树木叶子上的相关数据对模型进行了检验．检验结果表明建立的模型是正确的,能够反映树木的叶片面积与叶柄长、节间长和叶倾角之间的相互数量关系．     

Spatial distribution characteristics of stomata at the areole level in Michelia cavaleriei var. platypetala (Magnoliaceae)

Background and AimsIn hierarchically reticulate venation patterns, smaller orders of veins form areoles in which stomata are located. This study aimed to quantify the spatial relationship among stomata at the areole level.MethodsFor each of 12 leaves of M. cavaleriei var. platypetala, we assumed that stomatal characteristics were symmetrical on either side of the midrib, and divided the leaf surface on one side of the midrib into six layers equidistantly spaced along the apical–basal axis. We then further divided each layer into three positions equidistantly spaced from midrib to leaf margin, resulting in a total of 18 sampling locations. In addition, for 60 leaves, we sampled three positions from midrib to margin within only the widest layer of the leaf. Stomatal density and mean nearest neighbour distance (MNND) were calculated for each section. A replicated spatial point pattern approach quantified stomatal spatial relationships at different distances (0–300 μm).Key ResultsA tendency towards regular arrangement (inhibition as opposed to attraction or clustering) was observed between stomatal centres at distances <100 μm. Leaf layer (leaf length dimension) had no significant effect on local stomatal density, MNND or the spatial distribution characteristics of stomatal centres. In addition, we did not find greater inhibition at the centre of areoles, and in positions farther from the midrib.ConclusionsSpatial inhibition might be caused by the one-cell-spacing rule, resulting in more regular arrangement of stomata, and it was found to exist at distances up to ~100 μm. This work implies that leaf hydraulic architecture, consisting of both vascular and mesophyll properties, is sufficient to prevent important spatial variability in water supply at the areole level.     

Simulation of Codman's paradox reveals a general law of motion

Codman's paradox refers to a specific pattern of motion at the shoulder joint. It asked how a mysterious axial rotation about the longitudinal axis of the arm occurred during two or three sequential arm rotations that did not involve rotation about the long-axis. The objective of this paper was to find how the mysterious axial rotation occurred in the Codman's paradox. First, Codman's paradox and Codman's rotation were defined in general situations that involved arm rotations about orthogonal axes starting from the neutral attitude as well as rotations about non-orthogonal axes and starting from non-neutral attitudes. Then a general law of motion was proposed to answer the question of the Codman's paradox, which is stated as when the long-axis of the arm performs a closed-loop motion by three sequential rotations defined as Codman's rotation, it produces an equivalent axial rotation angle about the long-axis. The equivalent axial rotation angle equals the angle of swing-the second rotation in the three sequential long-axis rotations. Validity of the proposed law of motion is demonstrated by computer simulation of various Codman's rotations. Clinical relevance of the proposed law of motion is also discussed in the paper.     

Can one angle be simply subtracted from another to determine range of motion in three-dimensional motion analysis?

To determine the range of motion of a joint between an initial orientation and a final orientation, it is convenient to subtract initial joint angles from final joint angles, a method referred to as the vectorial approach. However, for three-dimensional movements, the vectorial approach is not mathematically correct. To determine the joint range of motion, the rotation matrix between the two orientations should be calculated, and angles describing the range of motion should be extracted from this matrix, a method referred to as the matrical approach. As the matrical approach is less straightforward to implement, it is of interest to identify situations in which the vectorial approach leads to insubstantial errors. In this study, the vectorial approach was compared to the matrical approach, and theoretical justification was given for situations in which the vectorial approach can reasonably be used. The main findings are that the vectorial approach can be used if (1) the motion is planar (Woltring HJ. 1994. 3-D attitude representation of human joints: a standardization proposal. J Biomech 27(12): 1399–1414), (2) the angles between the final and the initial orientation are small (Woltring HJ. 1991. Representation and calculation of 3-D joint movement. Hum Mov Sci 10(5): 603–616), (3) the angles between the initial orientation of the distal segment and the proximal segment are small and finally (4) when only one large angle occurs between the initial orientation of the distal segment and the proximal segment and the angle sequence is chosen in such a way that this large angle occurs on the first axis of rotation. These findings provide specific criteria to consider when choosing the angle sequence to use for movement analysis.     

The phloem necrosis virus disease of tea in Ceylon; further characterization of necrosis in the leaf

Phloem necrosis due to the virus disease of that name in tea is fully described as it affects the leaf, on which diagnosis is chiefly based. Originating in the protophloem, it may extend inwards to the metaphloem and outwards to the pericycle, causing the breakdown and discoloration of the cells and their eventual death and obliteration, cell enlargement, and the production of new thin-walled cells by hyperplasia. The condition is termed 'true necrosis' to distinguish it from the non-pathogenic 'false necrosis', of unknown cause, which originates typically in the metaphloem but may have the same histological effects except for the absence of hyperplasia. In the petiole, the visual distinction between 'true' and 'false' necrosis on the basis of their position as seen in a transverse section of the bundle is relatively easy, and the continued use of this method of diagnosis is recommended. No such distinction can reliably be made in the midrib, where 'false' necrosis often occurs in the same position as 'true' necrosis, i.e. immediately within the pericycle. This is interpreted ontogenetically in terms of the smaller total width of the phloem in the midrib as compared with the petiole bundle; it effectively prevents the use of the midrib for diagnosis. The observations are discussed in terms of the inherent properties of the phloem, as affected by viruses and other agencies reported to have caused necrosis, among plants in general.     

墨兰试管苗植株的发育解剖学研究

用石蜡切片和扫描电镜对墨兰试管苗植株的生长发育进行了研究。发现幼叶中脉附近的叶肉细胞类似栅栏组织,随着叶片的不断成熟,叶片基部中脉附近的叶肉细胞逐渐变为近圆形或椭圆形,而叶尖部和叶中部中脉附近的叶肉细胞仍似栅栏组织。茎的发育经历了原球茎、根状茎和假鳞茎3个阶段。原球茎的大部分细胞都含有淀粉粒,根状茎的皮层细胞含淀粉粒,而假鳞茎几乎不含淀粉粒。幼根没有髓,皮层细胞含淀粉粒：成熟根具含淀粉粒的髓。出瓶苗上即带有4个芽,一般只有最外侧叶腋的1个花芽和最内侧叶腋的1个叶芽发育。     

Light regime in relation to plant population geometry

A beam-emission experiment using a Monte Carlo technqiue was attempted in a computer model for the random distribution (RD) of foliage. The RD foliage consisted of 500 leaves of the same size, their individual areas being just 1/100 of the unit land area; i.e., a leaf area index (LAI) of RD foliage was equal to 5. Satisfactory agreement of light transmission under a uniform overcast sky was obtained between the observed values in the RD foliage and theoretical anticipations for any leaf inclination angle and/or leaf area density. This result demonstrated the usefulness of the Monte Carlo technique adopted in this experiment. The Monte Carlo experiment was also applied to a patch-like population. For such a foliage, additional effect of lateral incidence were worth noting in contrast with the microclimates in the corresponding infinite foliage. For every leaf inclination angle the mean light flux density below the middle stratum within a patch-like population was not further attenuate with increasing LAI.