Structure of the Photosynthetic Apparatus in Leaves of Freshwater Hydrophytes: 2. Quantitative Characterization of Leaf Mesophyll and the Functional Activity of Leaves with Different Degrees of Submersion

The structure of leaf photosynthetic elements was investigated on 42 boreal plant species characterized by different degrees of submergence (helophytes, neustophytes, and hydatophytes). Six main types of mesophyll structures were identified. Quantitative characteristics for the mesostructure of the photosynthetic apparatus in these groups were determined, such as the size and abundance of cells and chloroplasts in the mesophyll and epidermis, the number of plastids per cell in each tissue, the total surface area of the mesophyll cells, epidermal cells, and chloroplasts per unit leaf area. Analysis showed that quantitative characteristics of the photosynthetic apparatus in hydrophytes are determined by two factors: (a) the degree of leaf submergence and (b) the type of mesophyll structure. With an increasing degree of immersion in water, the mesophyll types change in a sequence isopalisade dorsoventral homogeneous. The leaves become thinner, their weight per unit area diminishes, cells and chloroplasts become less numerous (on a per unit leaf area basis), but their dimensions become larger. Adaptation to aquatic medium is also manifested in the increasing contribution of the epidermal tissue to the overall photosynthesis: in submerged leaves, the epidermis accounts for more than 50% of the photosynthetic activity. The occurrence of six structural types of leaves contrasting in their characteristics was confirmed by discriminatory analysis according to the qualitative parameters of mesophyll.     

Comparative Characterization of the Pigment Complex in Emergent, Floating, and Submerged Leaves of Hydrophytes

The content of chlorophylls (Chls) and carotenoids was studied in the leaves of 42 species of boreal aquatic plants with different degree of submergence (emergent, floating, and submerged) and isopalisade, dorsoventral, and homogenous types of mesophyll structure. Hydrophytes were shown to have a low Chl content (1–2 mg/g fr wt) and low Chls/carotenoids ratio (2.3–3.5) as compared to terrestrial plants. The pigment content per dry wt unit and unit leaf area was dependent on the type of mesophyll structure. It was a consequence of the changes in the parameters of leaf mesophyll structure characterizing the density of photosynthetic elements. In a sequence emergent floating submerged forms, the content of Chls and carotenoids decreased, and the photosynthetic capacity decreased due to a reduction in the chloroplast number per unit leaf area. Adaptation of submerged leaves to low illumination and slow CO₂ diffusion changed the functional properties of chloroplasts. An increase in the pigment content in the chloroplasts of submerged leaves (7 × 10^–9 mg Chl, 2 × 10^–9 mg carotenoids) as compared to emergent and floating leaves was accompanied by a decline in the photosynthetic capacity per Chl comprising 1.6 mg CO₂/(mg Chl h) versus 3.9 and 3.8 mg CO₂/(mg Chl h) in emergent and floating leaves, respectively.     

Basic Types of Structural Changes in the Leaf Mesophyll during Adaptation of Eastern Pamir Plants to Mountain Conditions

Quantitative characteristics of mesophyll structure were compared in leaves of eleven alpine plant species grown under natural conditions in the Eastern Pamirs at various altitudes, from 3800 to 4750 m. Basic types of changes in mesophyll structure, associated with plant adaptation to mountain conditions, were characterized. These changes manifested themselves in different numbers of cell layers and cell sizes in the palisade tissue and, as a consequence, in changed leaf thickness and cell number per unit of leaf area. Three plant groups were identified by the changes in the leaf structural characteristics depending on the type of mesophyll structure, ecological group of plant species, and altitude of plant habitat. The first group comprised alpine xerophytes with an isopalisade structure, in which the volume of palisade cells decreased and their number per unit of leaf area increased with the altitude of plant habitat. The number of mesophyll layers and leaf thickness decreased or did not change with altitude. The second group comprised subalpine plant species with a dorsoventral structure of mesophyll; these species occur below the line of continuous night frost. In these plant species, the number of mesophyll layers, leaf thickness, and cell number per unit of leaf area increased with altitude. The third group comprised mesophyte plants with a dorsoventral and homogenous mesophyll structure, which are encountered in a wide range of habitats, including the nival belt (from 4700 to 5000 m). In this group, cell volume increased and cell number per unit of leaf area decreased with altitude. We present a general scheme of leaf structural changes, which explains the changes in the quantitative characteristics of mesophyll as a function of altitude and highland environmental conditions.     

A Method for Calculating the Volume and Surface Area in Rice Mesophyll Cells

A method was developed for determining the surface area and volume of rice mesophyll cells of elaborate configuration. The method was employed to calculate these indices in several types of rice mesophyll cells found in seventy samples (53 species) of diverse origin coming from Japan, China, Korea, India, Nepal, Australia, France, Italy, Uzbekistan, Afghanistan, and Krasnodar and Primorskii regions. The cultivars of diverse geographic origin varied in cell shape and size due to the number, size, and arrangement of chloroplasts. When the volumes and surface areas of leaf mesophyll cells were compared using the method reported herein and a simple empirical model of the cell as a single ellipsoid, the two methods produced relatively similar data for cell volume; however, the surface area calculated by the former method was about two times larger than in the latter case. The method described in this paper allows for accurate calculations of the volume and surface area of rice mesophyll cells when data are available on the cell shape and linear dimensions and the number of chloroplasts per cell.     

Dry Matter and Leaf Structure in Young Wheat Plants as Affected by Cadmium, Lead, and Nickel

The effects of 1 mM cadmium, lead and nickel on dry mass, Cd, Pb, and Ni contents, and changes in leaf structure in young wheat plants were studied. In leaves, Cd content was highest, followed by Pb and Ni, in roots Cd content was also highest, but followed by Ni and Pb. Roots accumulated considerably larger amounts of the three heavy metals than leaves. Largest reductions of leaf and root mass were obtained with Cd. Pb and Ni effects were almost equal. Ni excess had a strong negative effect on mesophyll thickness, while Cd mostly reduced the number and size of vascular bundles and vessel diameter. High Pb reduced the diameter of vessels causing their different deformations. This revised version was published online in July 2006 with corrections to the Cover Date.     

Structural and Functional Changes in the Leaves of Plants from Steppe Communities as Affected by Aridization of the Eurasian Climate

Morphological and physiological characteristics of leaves from plant species collected in steppe communities in the various climatic zones in Eurasia were compared. The changes in leaf structure correlated with the major climatic factors. The mean thickness of leaves increased with increasing mean temperature of July and decreasing mean precipitation, which corresponded to aridity increase. The increased leaf thickness correlated with an increase in the specific leaf weight. The content of chlorophylls (a + b) in leaves greatly varied with plant habitats, whereas the chlorophyll a/b ratio remained unchanged. The chlorophyll content in leaf tissues had a general tendency to decrease with increasing leaf thickness. The leaf chlorophyll content positively correlated (R ² = 0.77) with the proportion of chlorenchyma in leaf tissues. It is concluded that steppe plants adapt to climate aridization at the structural level by increasing the proportion of protective heterotrophic components of the leaf without changing the functional activity of photosynthetic tissues.     

不同倍性大青杨的光合特性及叶片解剖结构比较

通过对二年生二倍体、三倍体和四倍体大青杨叶片结构和光合特性的研究,探讨不同倍性大青杨生长性状差异的原因。结果表明：不同倍性大青杨的净光合速率（Pn）、光饱和点（LSP）、光补偿点（LCP）等光合指标差异显著。三倍体的最大净光合速率（Pmax）高出二倍体21%。叶片厚度、表皮细胞厚度、栅栏组织厚度、栅栏组织与海绵组织厚度的比值,都是三倍体和四倍体大青杨比二倍体高,且均呈现增加的趋势。其中,四倍体大青杨表皮细胞最厚,上下表皮分别高出二倍体64%和17%。三倍体大青杨的栅栏组织厚度及栅栏组织与海绵组织厚度的比值最大,比值高出二倍体50%。通过叶片光合特性和解剖结构比较证明,三倍体和四倍体大青杨对光的适应性和光合作用能力强于二倍体。     

植物叶片解剖结构的量化及其在C4植物高粱中的应用

测量切片内所有细胞或细胞器的总周长,并乘以切片厚度和校正因子是一种量化植物叶片解剖结构的常用方法。前人在使用该方法时因忽视了细胞大小、切片厚度等影响估算精确度的因素,从而导致计数样本偏少并产生较大误差。通过分析该方法误差产生的原因及其与精确度的关系,明确了该方法在实际应用中不同精确度所要求对应的切片数量。在此基础上,将该方法用于C4植物高粱(Sorghum bicolor)叶片结构的量化,提出叶肉细胞与维管束鞘细胞接触面积(CABM)等参数,发展并完善了植物叶片解剖结构的量化方法。     

Shoot Growth and Changes in the Mesophyll Cell Structure in the Biennial Needles of Abies sibirica Ledeb.

Structural aspects of source–sink relations in current-year and second-year shoots of Siberian fir were investigated. The period of linear growth of current-year shoots was accompanied by a decrease in the size of starch granules in chloroplasts and the number of mitochondria, as well as by the disappearance of lipid inclusions in the mesophyll cells of second-year needles. Growth processes did not affect seasonal changes in the structure of chloroplast grana in these cells. After the termination of shoot growth, large polysaccharide globules appeared in the cytoplasm of mesophyll cells in second-year needles. It is possible that the formation of these inclusions prevents the inhibition of the photosynthesis by starch, which was accumulated in chloroplasts.     

田间大豆叶片成长过程中的光合特性及光破坏防御机制

田间大豆叶片在成长进程中光饱和光合速率持续提高,但气孔导度的增加明显滞后.尽管叶片在成长初期就具有较高的最大光化学效率,但是仍略低于发育成熟的叶片.随着叶片的成长,光下叶片光系统Ⅱ实际效率增加;非光化学猝灭下降.幼叶叶黄素总量与叶绿素之比较高,随着叶面积的增加该比值下降,在光下,幼叶的脱环氧化程度较高.因此认为大豆叶片成长初期就能够有效地进行光化学调节;在叶片生长过程中依赖叶黄素循环的热耗散机制迅速建立起来有效抵御强光的破坏.     

沙棘雌雄株叶片解剖结构比较研究

为了研究沙棘雌、雄株叶片的第二性征,本文采用石蜡切片法观察了沙棘雌、雄株叶片结构的差异。结果表明：（1）沙棘雌、雄株叶片均由表皮、叶肉和叶脉3部分组成,表皮均由1层细胞构成,表皮毛发达,上表皮有拟泡状细胞;叶肉栅栏组织与海绵组织分化明显。（2）雌株上表皮具更多的拟泡状细胞,其主脉韧皮部薄壁细胞及其下方的一些薄壁细胞含较多的后含物,下表皮的表皮毛更浓密;而雄株的叶片厚度、叶片上表皮厚度、栅栏组织厚度、栅栏组织厚度/海绵组织厚度均显著大于雌株,且其主脉维管束更发达。结果表明,沙棘雌雄株叶片解剖结构存在明显差异,这些差异是第二性征的表现,也是沙棘长期进化中形成的稳健的适应策略,可能有利于该物种的繁衍。     

沙生植物沙鞭叶表皮微形态特征及其生态适应性研究

利用光镜对沙生植物沙鞭50个代表种群的叶表皮微形态特征进行了观测、统计和分析,总结了代表种群叶表皮结构的异同,系统探讨了叶表皮微形态特征的分类学及生态适应性意义。结果表明,沙鞭叶表皮由长细胞、短细胞、气孔器细胞和刺毛组成,无微毛、大毛和乳突,表现为典型的狐茅型;脉间长细胞形态和壁波曲程度、短细胞形态和分布式样、气孔器体积大小与分布及副卫细胞形状、以及刺毛形态等在种群间具有明显差异,可将其鉴分为两大类。同时,沙鞭种群叶表皮气孔密度随环境干旱程度的增加而增加,第一类气孔密度大、第二类气孔密度小,这种结构有利于减少其体内水分的散失,进而更好地适应干旱环境,是对荒漠环境的一种微观形态解剖特征性适应。     

Optimization of the Leaf Mesophyll Structure in Alloploid and Diploid Wheat Species

Comparative analysis of the indices of plant growth and mesostructure of the photosynthetic apparatus was carried out with ten wheat (Triticum L.) species of various origins. Wheat alloploid forms (tetra- and hexaploids with the chromosome numbers of 28 and 42) exceeded the diploid forms (the chromosome number of 14) 2.3–2.4-fold by their absolute growth rate (AGR). As a result, the alloploid species developed a larger assimilation area; this change involved the internal reorganization of leaf phototrophic tissues and an increase in the cumulative internal assimilation area. In addition, the alloploid species manifested a higher correlation between the surface areas of cell and chloroplast membranes caused by a decrease in the cell number per the unit leaf area, a relative increase of the number of composite multifaveolate cells, a considerable expansion (in volume and surface area) of mesophyll cells, and an increase in chloroplast size and numbers. The decreased ratio between the characteristics of the cell membrane and chloroplast envelope presumes that CO₂ diffusion via cell and chloroplast membranes in the leaves was better balanced in the alloploid wheat species than in the diploid forms. All wheat species did not notably differ in their plastid–cytoplasm ratio (cell volume corresponding to one chloroplast and cell surface area per one chloroplast) and the ratio of surface area of cells to cell volume. The discriminant analysis revealed the indices of leaf growth and mesophyll structure instrumental in distinguishing between the diploid and alloploid species: leaf area, AGR, and cell size and number. The change in the latter indices optimized the structure of leaf phototrophic tissues in tetraploid and hexaploid species; as a result, the internal assimilation area was expanded and, consequently, leaf CO₂ conductance was increased.     

不同地区青檀叶片的解剖及表皮特征的比较研究

采用石蜡切片技术和叶表皮离析法,对我国8个省共15个居群的青檀(Pteroceltis tatarinowii Maxim.)叶片特征进行了比较和分析.研究发现:各居群的栅栏组织细胞层数多数为2层,显示出一定的稳定性,但不同居群的叶片横切面解剖特征存在显著性差异.叶片上表皮细胞有多边形和不规则形两种类型,而下表皮细胞均为不规则形.上表皮细胞垂周壁为平直形、圆形、浅波形或深波形,差异较明显,下表皮细胞垂周壁则仅有深波形和浅波形两种.叶表皮毛的分布位置在各居群间不同,而表皮毛的类型在居群间稳定,且在多数居群中表皮毛与周围细胞形成花环结构.各居群的气孔器类型均为不规则型且均仅在下表皮随机分布,但气孔器的密度、大小等特征在不同居群间存在显著性差异.生境地的气候因子对青檀叶的特征产生一定的影响.结果表明:青檀叶的解剖特征在不同居群间差异性较大,青檀叶表皮的气孔密度等特征在居群间有一定差异,而气孔器类型和表皮细胞形状等特征较稳定,有一定的分类学价值.     

A Remarkable New Leaf Type With Unusual Photosynthetic Tissue in a Central Asiatic Genus of Chenopodiaceae

Abstract: From the hygrohalophyte Borszczowia aralocaspica Bunge (Chenopodiaceae), a new leaf type with 1-layered chlorenchyma is described as "borszczovoid" and compared with other leaf types in subfamily Salsoloideae. The chlorenchyma is suspected to represent a unique C₄ type. Evidence is cited from anatomical studies and documented by micrographs and Carbon isotope determinations (ä¹³C values). The 1-layered photosynthetic tissue combines all essential anatomical characters of a 2-layered chlorenchyma of regular C₄ plants and is in intimate contact with concentrically arranged peripheral bundles. The ä¹³C values are − 13.03 ‰ from young stems and − 13.78 ‰ from leaves. The results are discussed in the anatomical, physiological and taxonomic framework. In addition, from distantly-related Suaeda species of section Conosperma the conospermoid leaf type is re-described. It is characterized by typical palisade and Kranz layers and differs from the C₄ suaedoid type by an external water-storaging hypodermis and an arrangement of Kranz cells reminiscent of the atriplicoid type from subfamily Chenopodioideae. From eight other species of Chenopodiaceae ä¹³C values are given for the first time.     

Photosynthetic and anatomical responses of Eucalyptus grandis leaves to potassium and sodium supply in a field experiment

Although vast areas in tropical regions have weathered soils with low potassium (K) levels, little is known about the effects of K supply on the photosynthetic physiology of trees. This study assessed the effects of K and sodium (Na) supply on the diffusional and biochemical limitations to photosynthesis in Eucalyptus grandis leaves. A field experiment comparing treatments receiving K (+K) or Na (+Na) with a control treatment (C) was set up in a K‐deficient soil. The net CO₂ assimilation rates were twice as high in +K and 1.6 times higher in +Na than in the C as a result of lower stomatal and mesophyll resistance to CO₂ diffusion and higher photosynthetic capacity. The starch content was higher and soluble sugar was lower in +K than in C and +Na, suggesting that K starvation disturbed carbon storage and transport. The specific leaf area, leaf thickness, parenchyma thickness, stomatal size and intercellular air spaces increased in +K and +Na compared to C. Nitrogen and chlorophyll concentrations were also higher in +K and +Na than in C. These results suggest a strong relationship between the K and Na supply to E. grandis trees and the functional and structural limitations to CO₂ assimilation rates.     

B9和PP333对人参生长和叶片光合特性的影响

人参叶片展叶前喷施3 000 mg*L-1的B9或400 mg*L-1的PP333后,人参植株的叶片面积、叶柄长、茎长和花梗长降低,茎直径增加,叶片叶绿素含量、叶绿素a/b比值、净光合速率、气孔导度和蒸腾速率显著提高,PP333的效果更为明显.     

Structural and Functional Rearrangements of Mesophyll as a Probable Basis for the Cytokinin-Dependent Assimilate Translocation in Detached Leaves

The effects of benzyladenine (BA) on the mesophyll functioning, such as osmotic potential (), the effect of the inhibitors of ⁺-ATPase on the influx of ¹⁴C-sucrose, the direction of carbon metabolism, and the rate of dark respiration, were followed in the detached leaves of pumpkin (Cucurbita pepo L.) and broad beans (Vicia faba L.). BA elevated and established a gradient of (_p) between the treated and untreated leaf regions. The inhibitors of H⁺-ATPase did not affect the BA-induced influx of ¹⁴C-sucrose. The changes were accompanied with the elevated synthesis of starch and other polymeric compounds and the diminished synthesis of the substances of relatively low molecular weight. The stimulation of dark respiration was short and inconsiderable. The author concludes that the BA-induced transport was a passive process related to a increase. Leaf expansion accompanied by the synthesis of high-molecular-weight substances essential for cell growth and by starch synthesis apparently increased the sink capacity of the BA-treated detached leaves. The diminished efflux from the leaf blade was probably related to a lowered level of the transportable carbon compounds restricting their entry into the phloem. The influx induction could result from the activation of growth and metabolic processes, the decline in the number of organic molecules per cell volume unit, and the development of _p between the source and sink leaf regions.     

Photosynthetic Characteristics of the Palisade Mesophyll and Spongy Mesophyll in the CAM/C4 Intermediate Plant,Peperomia camptotricha*

Several photosynthetic parameters were examined in the different tissue layers of leaves from the recently characterized crassulacean acid metabolism/C₄ intermediate plant, Peperomia camptotricha (Nishio and Ting, 1987). Light appears to control the development of certain photosynthetic characteristics within the tissue layers, while factors other than light seem to dictate others. Analysis of the chlorophyll content (including P700) and chlorophyll-proteins indicated that more light harvesting chlorophylls were associated with reaction centers in the tissues that were shaded by overlying tissue. Electron transport activity and chlorophyll-protein analysis indicated that the ratio of photosystem I to photosystem II in the spongy mesophyll (the abaxial tissue type and hence most shaded) was elevated relative to the overlying chlorophyll-rich median palisade mesophyll. The elevated photosystem I relative to photosystem II in the spongy mesophyll of Peperomia camptotricha may be related to C₄ metabolism and an increased requirement for ATP.     

林下与全光下地枫皮叶片形态和光合特性的比较

测量了林下与全光下地枫皮的叶片形态和光合-光响应曲线,探讨光强对地枫皮的形态和生理特性的影响。结果表明：林下与全光下地枫皮叶片净光合速率（Pn）、气孔导度（Gs）、蒸腾速率（Tr）和水分利用效率（WUE）对光强的响应趋势均基本一致,但全光下的Pn、Gs和Tr值较高,林下WUE值较高。全光下地枫皮的最大净光合速率、光饱和点和光补偿点均极显著高于林下,但弱光下的量子效率无显著差异;林下地枫皮的叶长、叶宽、干物质重、叶面积和比叶面积等叶片形态参数均极显著大于全光。推断地枫皮为耐阴性较弱的阳生植物,其光合能力和光饱和点较低,是对干旱环境的适应性反应;全光下地枫皮叶片狭小降低了吸光面积,有利于避免过高光强对叶光合器官的损伤。