Structural Adaptation of the Leaf Mesophyll to Shading

Structural characteristics of the mesophyll were studied in five boreal grass species experiencing a wide range of light and water supply conditions. Quantitative indices of the palisade and spongy mesophyll tissues (cell and chloroplast sizes, the number of chloroplasts per cell, the total cell and chloroplast surface area per unit leaf surface area) were determined in leaves of each of the species. The cell surface area and the cell volume in spongy mesophyll were determined with a novel method based on stereological analysis of cell projections. An important role of spongy parenchyma in the photosynthetic apparatus was demonstrated. In leaves of the species studied, the spongy parenchyma constituted about 50% of the total volume and 40% of the total surface area of mesophyll cells. The proportion of the palisade to spongy mesophyll tissues varied with plant species and growth conditions. In a xerophyte Genista tinctoria, the total cell volume, cell abundance, and the total surface area of cells and chloroplasts were 30–40% larger in the palisade than in the spongy mesophyll. In contrast, in a shade-loving species Veronica chamaedris, the spongy mesophyll was 1.5–2 times more developed than the palisade mesophyll. In mesophyte species grown under high light conditions, the cell abundance and the total cell surface area were 10–20% greater in the palisade mesophyll than in the spongy parenchyma. In shaded habitats, these indices were similar in the palisade and spongy mesophyll or were 10–20% lower in the palisade mesophyll. In mesophytes, CO₂ conductance of the spongy mesophyll accounted for about 50% of the total mesophyll conductance, as calculated from the structural characteristics, with the mesophyll CO₂ conductance increasing with leaf shading.     

岷江上游干旱河谷海拔梯度上白刺花叶片生态解剖特征研究

对岷江上游干旱河谷海拔梯度上（1 650～1 950 m）白刺花(Sophora davidii)叶片进行生态解剖学研究.观测指标包括叶片形态特征（叶长宽比、叶面积、叶片厚度）、解剖结构（表皮厚度、栅栏组织厚度(P)、海绵组织厚度(S)、P/S比值、表皮角质膜厚度）及叶表皮特征（气孔器密度和面积、表皮细胞密度和面积、表皮毛密度和长度）.结果表明,白刺花叶片面积为0.144～0.208 cm²,叶总厚度为171.58～195.83 μm;叶肉组织分化明显,栅栏组织厚度与海绵组织厚度分别为69.83～82.42和62.00～ 80.67 μm,P/S的比值为1.14～1.01,上下表皮厚度分别为14.03～15.33和13.88～16.17 μm,上下角质膜厚度分别为2.66～4.56和2.76～2.02 μm;气孔密度为13.71～15.02个·mm^-2,其面积为249.86～280.43 μm²;表皮细胞密度为160.54～178.43个·mm^-2,其面积为557.43～626.85 μm²;表皮毛长度为186.51～260.99 μm,其密度为18.29～32.27个·mm^-2.随海拔升高叶面积、叶厚度、栅栏组织和海绵组织的厚度、气孔器面积、表皮细胞面积以及表皮毛密度呈增加趋势,而角质膜厚度、表皮细胞密度和表皮毛长度则呈减小趋势;叶长宽比、P/S的比值、表皮厚度与气孔器密度无明显差异.     

Changes in chloroplast morphology of different parenchyma cells in leaves of Haberlea rhodopensis Friv. during desiccation and following rehydration

The size, shape, and number of chloroplasts in the palisade and spongy parenchyma layers of Haberlea rhodopensis leaves changed significantly during desiccation and following rehydration. The chloroplasts became smaller and more rounded during desiccation, and aggregated in the middle of the cell. The size and number of chloroplasts in the palisade parenchyma cells were higher than in spongy parenchyma. The good correlation observed between the size or number of chloroplasts and the cross-sectional area of mesophyll cells, the cross-sectional width of the leaf and its water content suggested that the palisade cells were more responsive to water availability than the spongy cells. Changes in chloroplast number during desiccation and rehydration process are characteristic features for desiccation-tolerant plants (especially in homoiochlorophyllous strategy).     

The effect of light quality on leaf production and development of in vitro-cultured plants of Alternanthera brasiliana Kuntze

We investigated the influence of light quality on the leaf development of Alternanthera brasiliana Kuntze (Amaranthaceae) grown in vitro. Growth parameters including specific leaf mass, thickness, and leaf density were lowest in plants grown under red light. Blue light induced the largest number of leaves/plant, and the largest thickness and area of the leafblade. Green and red lights induced the smallest leaf areas. The thickness of the abaxial-face epidermis and spongy parenchyma of the plants was significantly reduced in plants grown under red light. The thickness of the palisade parenchyma and upper epidermis were significantly increased in plants grown under blue light, compared to the other fluorescent-light treatments. The specific spectral band also influenced the differentiation of mesophyll cells. In the dark and under red light, the mesophyll was homogenous; and in the dark and under green light, the leaves were more compact. Under blue light, the cells displayed the characteristic palisade morphology. The results showed that the increase of a specific parenchyma type was related to a specific spectral band. All spectral-quality treatments reduced the numbers of stomata and trichomes. The results for green light were in some respects similar to those for red light, and in other respects similar to those for blue light, probably because phytochromes and cryptochromes are green-light receptors. This study indicated that Alternanthera plants have strong morphological plasticity induced by light. The results suggest that high-quality Alternanthera can be achieved by culturing the plants in vitro under a combination of blue and red light.     

青藏高原草地植物叶解剖特征

运用常规石蜡制片技术对我国青藏高原66种草地植物优势种的叶解剖特征进行研究,并分析了叶解剖特征与海拔、生长季降水及生长季均温之间的关系.结果表明:青藏高原草地植物叶片具有很多适应高寒环境的结构特征,如表皮层厚且表皮细胞大小差异显著,表皮毛等表皮附属物发达,异细胞丰富,通气组织普遍发达等;叶片各组成部分厚度的变异程度不同,其中海绵组织厚度变异最大,其次为上角质层、下表皮层、下角质层、上表皮层、栅栏组织,叶片厚度的变异最小;青藏高原草地植物叶片各组成部分的厚度存在协同进化,上下角质层厚度呈强烈正相关,海绵组织厚度与叶片厚度相关性最强;青藏高原草地植物叶片各组成部分的厚度与海拔、生长季降水、生长季均温3个重要环境变量呈较弱的相关性,总体表现为随海拔升高叶片各组成部分的厚度减小,而随生长季降水和生长季均温的增加叶片厚度增加.     

The functional significance of palisade tissue: penetration of directional versus diffuse light

Light gradients were measured in leaves that had different types of anatomical development of the mesophyll but similar pigment content. Leaves of the legume, Thermopsis montana, had columnar palisade and spongy mesophyll whereas leaves of the monocot, Smilacina stellata, had spongy mesophyll only. Light gradients were measured at 550 nm in both types of leaves when they were irradiated with collimated or diffuse light. When irradiated with collimated light, light gradients were steeper in leaves with spongy mesophyll in comparison to those that had palisade tissue. On the other hand, light gradients were similar between both leaf types when they were irradiated with diffuse light. Thus, columnar palisade cells facilitated the penetration of collimated light over diffuse light. These results suggest that palisade tissue may help distribute light more uniformly to chloroplasts within the leaf. Moreover, the functional significance of palisade tissue may be related to the amount of collimated light within the natural environment.     

Altered Leaf Structure and Function in Triazine-Resistant Common Groundsel (Senecio vulgaris)

Anatomical and physiological characteristics of leaves of triazinesusceptible and -resistant biotypes of common groundsel (Senecio vulgaris L.) were studied in order to explain the differences in light-saturated photosynthetic rates previously reported. Leaves were of uniform leaf plastochron index from greenhouse-grown plants. Susceptible plants had greater leaf fresh and dry weights and leaf areas, while resistant plants had greater specific leaf mass (mg fresh weight/cm²). Susceptible plants had greater amounts of total chlorophyll per unit leaf weight and a higher chlorophyll a/b ratio. Soluble protein in leaves was higher in susceptible chloroplasts on a weight and area basis, but similar to resistant chloroplasts on a unit chlorophyll basis. Activity of ribulose 1,5-bisphosphate carboxylase was higher in resistant plants on a fresh weight, leaf area, and milligram chlorophyll basis. Stomatal frequency, length, and arrangement were similar between biotypes, as were transpiration and conductance. Resistant leaves had less air space (v/v), more cells in palisade and spongy mesophyll, and a greater volume of palisade tissue than spongy, when compared to susceptible leaves. Differences in leaf structure and function between biotypes are probably due to a complex of developmental adaptations which may be only indirectly related to modified photosystem II in resistant plants. These results indicate that the consistently lower rates of net photosynthesis and yield in resistant plants cannot be explained solely on the basis of these leaf characteristics. Several possible mechanisms to account for reduced productivity are suggested.     

风毛菊属3种植物叶的解剖结构比较

采用石蜡切片法对分布于祁连山海拔5 000 m左右流石滩上菊科风毛菊属水母雪兔子（Saussurea medusa Maxim）、鼠曲雪兔子（Saussurea gnaphalodes (Royle) Sch.）、红叶雪兔子（Saussurea paxiana Diels.）3种植物叶片的解剖结构进行了比较研究,结果表明：叶片表皮细胞均为单层,上下表皮都有气孔分布,气孔不下陷;角质层较厚,叶表面均被单列细胞的表皮毛。3种植物均为异面叶;叶肉栅栏组织较发达,通常由2～3层细胞组成,但栅栏组织细胞排列较疏松;海绵组织存在大量的细胞间隙;叶肉中通气组织发达,且均有不规则裂生性气腔。叶脉维管束中韧皮部都具有异细胞存在。这些共同特征是3种植物对高山地区缺氧、低温、强辐射等自然条件长期适应的结果。但是,3种植物在叶片的外部形态特征、叶肉栅栏组织细胞的特点、维管束发育程度、内分泌结构、不规则裂生气腔等方面又存在明显的不同,表现出3种植物对环境的适应也是存在差异的。     

Morphactin effects on soybean leaf anatomy and chlorophyll content

The effect of chlorflurenol (methyl 2-chloro-9-hydroxyfluorene-9-carboxylate) (CF) on chlorophyll (chl) content was studied in intact plants and floating leaf disks. For intact soybean (Glycine max (L.) Merrill) plants grown in the growth chamber, 2.5 μg/ml CF applied 10 to 20 d after planting retarded chl decline in senescing tissues such as cotyledons and unifoliate leaves and increased chl content in recently expanded tissues such as trifoliate leaves. CF did not retard chl decline in the dark unless regulator application was followed by a period of 24 h in the light prior to darkness. In floating leaf disk tests, CF retarded chl decline in dock (Rumex obtusifolius L.) and radish (Raphanus sativus L.) at concentrations of 10^?4 M, but was ineffective at lower concentrations. Chl decline was significantly hastened by CF in tobacco (Nicotiana tabacum L.) and soybean, but was unchanged in barley (Hordeum vulgare L.). CF treatment increased tissue weight (g fresh wt/cotyledon; g dry wt/ cm² for unifoliate and trifoliate leaves), decreased moisture content, and increased leaf thickness, palisade layer thickness, and palisade and spongy mesophyll cell counts. We conclude that plants treated with morphactins show greater green coloration predominantly because of growth effects, and only in small part because of prevention of chl decline in senescing tissues.     

新疆野苹果不同地理分布与树龄的叶片解剖结构特征

以新疆野苹果为材料,对不同居群及不同树龄叶片解剖结构特征进行了分析研究。结果表明：新疆野苹果叶片属于异面叶,由上表皮、栅栏组织、叶脉、海绵组织和下表皮组成;不同居群新疆野苹果叶片厚度及细胞排列方式有差异,巩留、额敏及托里的野苹果栅栏组织增厚,且栅栏组织/海绵组织比值增大;新源的野苹果叶片及栅栏组织较薄,栅栏组织/海绵组织比值偏低;不同树龄新疆野苹果叶片厚度随着树龄的增大而呈变薄的趋势,多年生野苹果及野苹果“树王”栅栏组织及海绵组织层数最少,厚度较薄。这些结果暗示新疆野苹果通过各部分结构的协调生长来保证光合作用效率,以适应不同的生长环境以及处于不同发育时期的生长状况。本研究为新疆野苹果原位保护、保育提供了实验依据。     

The paraveinal mesophyll of soybean leaves in relation to assimilate transfer and compartmentation

Nitrogen and carbohydrate assimilates were temporally and spatially compartmented among various cell types in soybean (Glycine max L., Merr.) leaves during seed filling. The paraveinal mesophyll (PVM), a unique cell layer found in soybean, was demonstrated to function in the synthesis, compartmentation and remobilization of nitrogen reserves prior to and during the seed-filling stages. At anthesis, the PVM vacuoles contain substantial protein which completely disappears by two weeks into the seed filling. Distinct changes in the PVM cytoplasm, tonoplast and organelles were correlated with the presence or absence of the vacuolar material. Microautoradiography following the accumulation of several radiolabeled sugars and amino acids demonstrated the glycoprotein nature of the vacuolar material. Incorporation of methionine, leucine, glucose, and glucosamine resulted in heavy labelling of the PVM vacuole, in contrast to galactose, proline, and mannose which resulted in a much reduced labelling pattern. In addition, starch is unequally compartmented and degraded among the various leaf cells during seed filling. At the end of the photoperiod at the flowering stage, the highest starch accumulation was in the second palisade layer followed by the spongy mesophyll and the first (uppermost) palisade layer. Starch in the first palisade layer was completely degraded during the dark whereas the starch in the second palisade and spongy mesophyll was not remobilized to any appreciable extent. By mid-podfilling (approximately five weeks postanthesis) starch was absent in the first palisade layer at the end of the photoperiod while the second palisade and spongy mesophyll layers contained substantial starch. Starch was remobilized from these latter cells during the remainder of seed filling when current photosynthetic production is low. Structural changes associated with cell senescence first appear in the upper palisade layer and then progress (excluding the PVM) to the second palisade and spongy mesophyll layer. The PVM and phloem appear to retain their structural integrity into the leaf yellowing stage. Reducing sink capacity by pod removal resulted in a continued accumulation of vacuolar protein, an increase in cytoplasmic volume, and fragmentation of the vacuole in the PVM. Pod removal also resulted in an increased amount of accumulated starch (which did not turn over) in all mesophyll layers, and an increase in cell size and cell-wall thickness.     

岷江上游干旱河谷矮探春叶片特征与环境因子的关系

对岷江上游干旱河谷矮探春(Jasminumhumile L.)的叶片形态解剖特征进行了显微观察,分析了海拔梯度上叶片形态与环境因子的关系。结果表明,矮探春叶片呈长椭圆形,叶肉组织分化明显;随着海拔升高,叶面积、厚度、干重、饱和含水量、海绵组织厚度,中脉厚度以及厚角组织厚度/中脉厚度之比(M/C)等呈增大趋势,而栅栏组织厚度/海绵组织厚度之比(P/S)则减小;叶片长/宽比、表皮厚度、栅栏组织厚度和比叶重在海拔梯度上无明显差异。叶面积、干重、饱和含水量、叶厚度和海绵组织厚度等参数两两之间呈显著正相关,而它们与P/S均呈显著负相关。叶面积、干重、海绵组织厚度和M/C等主要受土壤含水量的影响,并随着土壤含水量增加而增大;P/S随着土壤含水量和年降水量增加而减小;叶片厚度、饱和含水量和中脉厚度均随着温和度的降低而增大。岷江上游干旱河谷区土壤水份和生长季温度可能是影响矮探春叶片形态解剖特征的主要因子。     

Leaf anatomy during leaf development of photoautotrophically <Emphasis Type="Italic">in vitro</Emphasis>-grown tobacco plants as affected by growth irradiance

Tobacco (Nicotiana tabacum L.) plants were cultured in vitro photoautotrophically at three levels of irradiance (PAR 400–700 nm): low (LI, 60 μmol m⁻² s⁻¹), middle (MI, 180 μmol m⁻² s⁻¹) and high (HI, 270 μmol m⁻² s⁻¹). Anatomy of the fourth leaf from bottom was followed during leaf development. In HI and MI plants, leaf area expansion started earlier as compared to LI plants, and both HI and MI plants developed some adaptations of sun species: leaves were thicker with higher proportion of palisade parenchyma to spongy parenchyma tissue. Furthermore, in HI and MI plants palisade and spongy parenchyma cells were larger and relative abundance of chloroplasts in parenchyma cells measured as chloroplasts cross-sectional area in the cell was lower than in LI plants. During leaf growth, chloroplasts crosssectional area in both palisade and spongy parenchyma cells in all treatments considerably decreased and finally it occupied only about 5 to 8 % of the cell cross-sectional area. Thus, leaf anatomy of photoautotrophically in vitro cultured plants showed a similar response to growth irradiance as in vivo grown plants, however, the formation of chloroplasts and therefore of photosynthetic apparatus was strongly impaired.     

冠层位置对5种阔叶树叶片解剖结构与氮含量的影响

不同树冠位置的叶片为了达到功能的最大化,形成了不同的结构与功能的特征。然而,在相同的环境条件下,不同树种之间叶片对环境的反应是否存在一致规律,我们仍然缺乏了解。本研究采用石蜡切片和化学分析方法,对东北林区5个常见阔叶树种（蒙古栎、白桦、水曲柳、胡桃楸和黄波罗）不同树冠位置叶片的形态（叶厚度）、解剖（气孔密度、保卫细胞长度、栅栏和海绵组织厚度）和氮（N）含量特征进行了研究。结果表明：叶片的特征在种间和种内不同树冠位置均存在明显差异,并存在较强的规律性。在种内,5个树种的叶片和栅栏组织厚度均是上层外部最大,而保卫细胞和海绵组织的变化不明显,N含量的变异与树种有关。叶片所处的树冠高度和暴露程度对叶片的结构与N含量变异有重要影响。树种之间,蒙古栎的气孔密度最大,叶片厚度和海绵组织厚度最小,保卫细胞最短,黄波罗恰恰相反。结果表明,为了更好地执行整个树冠的功能,不同树种叶片均出现了与冠层位置有关的结构特征适应。     

Leaf plasticity in peanut (Arachis hypogaea L.) in response to heavy metal stress

Phenotypic plasticity in morphological, anatomical and physiological traits of peanut (Arachis hypogaea L.) leaves was tested at four different concentrations of Cd, Cu and Zn under greenhouse conditions. Among 18 characteristics tested, nine were found to be the most sensitive and demonstrate the greatest phenotypic plasticity. These were: the leaf area (LA), the leaf mass per area (LMA), chlorophyll a concentration (Chl a), chlorophyll b concentration (Chl b), total chlorophyll concentration (Chl t), the effective quantum yield of photosystem II (ΦPS II), stomatal density of upper epidermis (SDU), palisade thickness (PT), and palisade to spongy thickness ratio (P/S). The plasticity of chlorophyll concentration and fluorescence parameters may be maladaptive and reflects metal toxicity to leaves, whereas the anatomical plasticity is adaptive, indicative of a tradeoff between the physiological and anatomic plasticity. The anatomical plasticity resulted in a xerophyte feature of leaves (i.e. small leaflets, thick lamina, upper epidermis, palisade mesophyll, as well as abundant and small stomata), which enhanced the capacity to resist drought caused by heavy metals via a decrease in root growth.     

Leaf structure and specific leaf mass: the alpine desert plants of the Eastern Pamirs, Tadjikistan

This study examines interrelationships between eight leaf attributes (specific leaf mass, area, dry mass, lamina thickness, mesophyll cell number per cm², mesophyll cell volume, chloroplast volume, and number of chloroplasts per mesophyll cell) in field-grown plants of 94 species from the Eastern Pamir Mountains, at elevations between 3800 and 4750 m. Unlike most other mountain areas, the Eastern Pamirs, Karakorum system, Tadjikistan provide localities where low temperatures and radiation combine with moisture stress at high altitudes. For all the attributes measured, significant differences were found between plants with different mesophyll types. Leaves with dorsiventral palisade structure (dorsal palisade, ventral spongy mesophyll cells) had thicker leaves with larger but fewer mesophyll cells, containing more and larger chloroplasts. These differences in mesophyll type are reflected in differences in the total surface of mesophyll cells per unit leaf area ( A _mes/ A ) or volume ( A _mes/ V ). Plants with isopalisade leaf structure (palisade cells under both dorsal and ventral surfaces) are more commonly xerophytes and their increased values of A _mes/ A and A _mes/ V decrease CO₂ mesophyll resistance, which is an important adaptation to drought. Path analysis shows the critical importance of mesophyll cell volume in leading to the covariance between the different leaf attributes and hence to specific leaf mass (SLM), even though mesophyll cell volume is not itself strongly correlated with SLM. This is because mesophyll cell volume increases SLM through its effects on leaf thickness and chloroplast number per cell, but decreases SLM through its negative effect on mesophyll cell density.     

构树雌雄株叶片解剖结构特征的比较研究

为了研究雌雄异株植物构树（Broussonetia papyrifera（L.） Vent.）的性别差异,以其叶片为材料,采用石蜡切片法,对其主要解剖结构特征进行观察和比较。结果显示：（1）构树雌、雄株叶片解剖结构组成一致,均由表皮、叶肉组织和叶脉3部分组成。其上、下表皮均由一层细胞构成,具有较厚的角质层及丰富的表皮毛,叶肉中栅栏组织高度发达,此外,维管系统在叶中所占比例很大;（2）构树雌、雄株叶片细胞大小及厚度在各类型组织间存在一定差异,雄株叶片上表皮厚度、栅栏组织厚度、主脉维管束木质部厚度及维管束厚度均显著大于雌株叶片,且在栅海比、组织结构紧密度、组织结构疏松度和主脉维管束木质部面积占维管束的比例等方面也与雌株有极显著差异。研究结果表明构树叶片的解剖结构不仅具有旱生植物叶片的典型特征,而且还表现出明显的性别差异,这可能与构树雌、雄株的生殖分配有关。     

Leaf anatomy and chloroplast ultrastructure of Mn-deficient orange plants

Leaf samples of Mn-deficient and Mn-sufficient (control) ‘Navelate’ orange plants grown in a greenhouse were taken to investigate the effects of Mn deficiency in leaf structure and chloroplast ultrastructure. Total leaf chlorophyll concentration was significantly lower in Mn-deficient plants than in control ones. Entire lamina thickness was not altered due to Mn deficiency. However, Mn deficiency resulted in disorganization of mesophyll cells, mainly of palisade parenchyma cells. The number of mesophyll chloroplasts per cellular area and their length were both affected negatively. The membranous system of chloroplasts was also disorganized. The percentages of starch grains and plastoglobuli per chloroplast of Mn-deficient leaves were significantly greater than those of control leaves.     

遮荫对南美蟛蜞菊、蟛蜞菊及其自然杂交种叶片显微结构及叶绿体超微结构的影响

利用JSM-6360LV型扫描电镜和JEM-1010型透射电镜,观察了南美蟛蜞菊、蟛蜞菊及其自然杂交种新近成熟和老熟叶片的解剖结构及叶绿体超微结构。结果表明:遮荫后该杂交种与其亲本新近成熟叶片均表现为上下表皮气孔密度、叶片总厚度及上下表皮厚度、栅栏组织、海绵组织厚度减小,叶绿体肿胀变形,基粒片层垛叠程度增加,淀粉粒增多变大;遮荫后杂交种老熟叶片总厚度及上表皮、栅栏组织、海绵组织厚度增加,入侵种的下表皮厚度及本地种的上表皮厚度增加,叶绿体超微结构在遮荫后均出现严重损伤,基粒片层类囊体结构边缘溶解等。说明三种蟛蜞菊属物种及各物种不同叶龄叶片对弱光条件的响应存在差异;杂交种叶片显微及超微结构在不同光照下的变化介于亲本之间,对遮荫环境能较好适应。