Leaf responses to iron nutrition and low cadmium in peanut: anatomical properties in relation to gas exchange

Aims

This study evaluated how iron nutrition affect leaf anatomical and photosynthetic responses to low cadmium and its accumulation in peanut plants.

Methods

Seedlings were treated with Cd (0 and 0.2 μM CdCl₂) and Fe (0, 10, 25, 50 or 100 μM EDTA-Na₂Fe) in hydroponic culture.

Results

Cadmium accumulation is highest in Fe-deficient plants, and dramatically decreased with increasing Fe supply. The biomass, gas exchange, and reflectance indices were highest at 25 μM Fe²⁺ treatments, indicating the concentration is favorable for the growth of peanut plants. Both Fe deficiency and Cd exposure impair photosynthesis and reduce reflectance indices. However, they show different effects on leaf anatomical traits. Fe deficiency induces more and smaller stomata in the leaf surface, but does not affect the inner structure. Low Cd results in a thicker lamina with smaller stomata, thicker palisade and spongy tissues, and lower palisade to spongy thickness ratio. The stomatal length and length/width ratio in the upper epidermis, spongy tissue thickness, and palisade to spongy thickness ratio were closely correlated with net photosynthetic rate, stomatal conductance, and transpiration rate.

Conclusions

Cd accumulation rather than Fe deficiency alters leaf anatomy that may increase water use efficiency but inhibit photosynthesis.     

遮荫对金莲花光合特性和叶片解剖特征的影响

以金莲花为材料,系统研究了不同遮荫处理(全光照、40%、60%和80%遮荫)对金莲花叶片光合特性和叶解剖结构的影响,为金莲花引种驯化和抚育管理提供理论依据。结果表明:全光照下金莲花叶净光合速率有明显光合"午休"现象,遮荫处理下"午休"现象不明显。其中,40%遮荫处理显著促进了金莲花叶片净光合速率,80%遮荫处理则起抑制作用;40%和60%遮荫处理促进金莲花叶蒸腾速率和气孔导度的提高,降低了蒸汽压亏缺,有利于光合作用的进行;80%遮荫处理下胞间CO2浓度提高,气孔限制值和蒸汽压亏缺均显著高于其它处理,抑制了金莲花叶蒸腾作用和光合作用;遮荫处理降低了金莲花叶光补偿点和暗呼吸速率,40%和60%遮荫处理下表观量子效率和最大净光合速率提高,说明金莲花有一定耐荫性;金莲花叶光合特性与解剖结构关系密切。遮荫降低了金莲花叶气孔密度和气孔指数,但气孔纵轴长和横轴长显著高于对照。40%和60%遮荫处理下,单叶气孔数量没有明显变化;遮荫下金莲花叶片增厚。栅栏组织厚度和细胞层数减少,栅栏细胞宽度加大,且第二层栅栏细胞有向海绵组织过度的趋势。海绵组织厚度增加,栅栏组织/海绵组织比值减小。随遮荫度增高,栅栏组织和海绵组织细胞间隙增大。该结构有利于提高对光能的利用效率;遮荫处理促进维管束的发育,维管束数量增加,木质部、韧皮部分化明显,其中中脉导管直径和导管数目增加,有利于蒸腾作用和光合作用的进行。     

Photosynthetic and anatomic responses of peanut leaves to cadmium stress

Cadmium (Cd) treatments caused an inhibition in the net photosynthetic rate (P _N) of peanut (Arachis hypogaea) plants, due to the reduction of stomatal conductance (g _s) and photosynthetic pigment contents, as well as the alteration in leaf structure. The decrease of the transpiration rate and g _s might result from the Cd-induced xerophyte anatomic features of leaves (i.e. thick lamina, upper epidermis, palisade mesophyll, high palisade to spongy thickness ratio, as well as abundant and small stomata). The decline of P _N was independent of the impairment in photosystem 2.     

Variation in leaf traits at different altitudes reflects the adaptive strategy of plants to environmental changes

Leaf anatomical traits play key roles in plant functions and display evolutionary adaptive changes to suit the surrounding environment. To reveal the adaptive mode and mechanisms of plants in response to global warming, we analyzed leaf morphology and anatomical structures in three different species, Epilobium amurense Hausskn., Pedicularis densispica Franch., and Potentilla fulgens Wall. ex Hook., growing along an elevational gradient (3,000–4,600 m) in the Yulong Mountains. The results showed leaf length and width decreased, whereas leaf thickness increased with increasing altitude in all three species. Thickness of leaf upper epidermis, lower epidermis, palisade and spongy mesophyll, and main vein increased with rising altitude. Stomatal density in each species increased with rising elevation. These results illustrate that plants can adapt to the environmental changes that accompany high altitudes by decreasing leaf area and increasing leaf thickness, mesophyll tissue thickness, and stomatal density. Such morphological and anatomical plasticity would lead to lower transpiration rates, enhanced internal temperature and water status, and improved photosynthetic capability.     

山茶属五种植物叶片解剖特征及与光合生理相关性研究

利用扫描电子显微镜观察山茶属（ＣａｍｅｌｌｉａＬｉｎｎ．）五种植物叶片解剖特征,并与光合生理相关性进行分析。实验选择的五种植物均为Ｃ３植物,其中小黄花茶（ＣａｍｅｌｌｉａｌｕｔｅｏｆｌｏｒａＹ．Ｋ．Ｌｉ）和四球茶（Ｃ．ｔａｃｈａｎｇｅｎｓｉｓＦ．Ｃ．Ｚｈａｎｇ）为阴性植物．叶绿素含量较高;咸宁短柱茶（Ｃ．ｓａｌｕｅｎｅｎｓｉｓＳｔａｐｆ．ｅｘＢｅａｎ．）为偏阴性植物,叶绿素含量较低;美丽红山茶（Ｃ．ｄｅｌｉｃａｔａＹ．Ｋ．Ｌｉ）和狭叶瘤果茶（Ｃ．ｎｅｒｉｉｆｏｌｉａＣｈａｎｇ）则属于半阴性植物。它们的光合生理指标值差异取决于叶片解剖特征的差异;与光补偿点,光饱和点,净光合速率（Ｐｎ）,光呼吸速率（Ｐｒ）,暗呼吸速率（Ｄｒ）相关性最密切的解剖指标是栅栏组织厚度、栅栏组织中的细胞层数、栅栏细胞内的叶绿体数,栅栏组织厚与叶厚的比值、与栅栏细胞粗度的比值、与海绵组织厚度的比值．叶片上表皮厚、上表皮角质层厚及下表皮气孔密度亦与上述光合生理指标呈弱正相关,叶片大小、气孔器大小及气孔口长则与上述光合生理指标呈弱负相关;下表皮厚、海绵组织厚度及栅栏细胞内叶绿体大小与各项光合生理指标几乎无关。该五种植物的叶片解剖和光合生理特     

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。     

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.     

干旱胁迫下石灰花楸幼苗叶片的解剖结构和光合生理响应

采用盆栽控水设置4种水分梯度,研究1年生石灰花楸幼苗叶片解剖结构和光合生理指标对干旱胁迫的响应。结果显示:(1)干旱胁迫下,石灰花楸叶片逐渐变薄,轻度和中度胁迫下栅海比显著升高,而重度胁迫下显著降低。(2)干旱胁迫抑制了光合色素合成,降低了叶绿素和类胡萝卜素含量,同时使叶绿素a/b和叶绿素/类胡萝卜素值升高。(3)随干旱程度加剧,净光合速率、蒸腾速率和气孔导度日变化整体下降,胞间CO2浓度整体上升,光合限制以非气孔因素为主。研究表明,石灰花楸能根据水分亏缺程度调整叶片结构和光合生理特征以维持生存和生长,具有较强的耐旱性。     

Observations on the Three-dimensional Structure of the Leaf Cuticle in Certain Plants

The three-dimensional structure of the leaf cuticle has beenstudied microscopically on cuticular preparations from a numberof plants. The cuticle is one- to several-layered, the latterincluding, besides the epidermis, sub-epidermal layers suchas hypodermis, palisade, and spongy mesophyll.     

酸枣叶片结构可塑性对自然梯度干旱生境的适应特征

叶片是植物体暴露于环境中面积最大的器官,其最易感知环境变化而发生形态和结构上的改变。为探究植株叶片结构对不同生境的适应机理,研究以生长在烟台-石家庄-宁夏-新疆不同地域气候条件形成的自然梯度干旱环境中的酸枣为试验材料,应用植物显微技术研究酸枣叶片的结构的可塑性对不同自然梯度干旱环境的适应特征。结果表明:酸枣叶表皮着生有表皮毛,表皮细胞外覆有角质层与蜡质。叶肉为全栅型,栅栏组织发达,海绵组织退化,叶肉中有晶体及大量的分泌细胞。从烟台至新疆随生境梯度干旱加剧,酸枣叶片叶面积逐渐变小,叶片厚度依次增加,叶表皮角质层加厚,且上角质层厚度大于下角质层厚度;叶片上下表皮细胞长径及短径先增后降,栅栏组织总厚度和密度依次增大、层数减少,各层栅栏组织细胞的长径逐渐增加。叶脉薄壁细胞相对厚度逐渐减小,导管管径增大,晶体(草酸钙晶体)数增多。在梯度干旱环境中酸枣植株通过减小叶面积、提高栅栏组织密度、增加叶片及角质层厚度降低蒸腾作用,减少水分散失;通过增大导管管径提高水分利用率;通过增加晶体数量提高叶片机械性能,改变细胞的渗透势、提高吸水和保水能力。上述叶片结构的变化是酸枣植株长期对不同自然梯度干旱生境的适应特征。由此可知,叶片形态结构中叶面积、叶片厚度、角质层及叶肉组织(栅栏组织)随环境变化的可塑性较大。     

Photosynthesis and ultrastructure of photosynthetic apparatus in tomato leaves under elevated temperature

The microstructure of leaves and ultrastructure of chloroplasts were examined in tomato (Lycopersicon esculentum L.) plants treated with elevated temperature. Plants were exposed to 35°C for 30 d after florescence. The plants grown continuously under 25°C served as controls. Compared with the controls, the net photosynthetic rate (P _N) in stressed plants decreased significantly. Stomatal conductance, intercellular CO₂ concentrations, the rate of transpiration, and the limitation of stomatal conductance showed that the decrease in P _N was caused mainly by nonstomatal restrictions. Meanwhile, stomata density increased significantly in the stressed plants. The stomata status of opening and closing became disorganized with a prolonged 35°C exposure. The damage of chloroplast membrane occurred earlier and was more serious in the plants under elevated temperature. At the same time, the thylakoids were loosely distributed with lesser grana, but the number of lipid droplets increased in chloroplasts. The number of starch grains in chloroplasts increased first and then decreased. In addition, the length of the main nerve in leaves increased and the main vein showed distortion in the plants stressed by 35°C. An increase was observed in the number of cells on the abaxial side of the main vein and these cells were overly congregated. The thickness of a vertical section became thinner in the stressed leaves. The cells of the upper epidermis thinned, and the ratio of palisade tissue to spongy tissue decreased. Generally, the photosynthetic apparatus of tomato changed significantly and the changed chloroplast ultrastructure might be one of the important reasons that caused the decrease of P _N under 35°C.     

玉龙雪山不同海拔草血竭叶片形态与解剖结构的比较研究

采用常规石蜡制片技术和显微观察方法,对生长于滇西北玉龙雪山3 000~4 600m之间9个不同海拔高度的草血竭(Polygonum paleaceum)叶片表型及解剖结构进行比较研究,其目的是揭示该植物对不同海拔环境条件的响应特征,进而探讨植物对全球气候变化的生态适应性。结果显示:(1)草血竭叶片长度、宽度、长宽比均与海拔呈显著负相关关系,其降幅分别达84.64%、74.87%和37.50%。(2)草血竭叶片为异面叶,由表皮、叶肉、叶脉三部分组成;表皮细胞单层,角质层厚;叶肉的栅栏组织具2层或3层细胞;叶脉中的维管束呈1圈不连续的环状排列。(3)叶片厚度、栅栏组织厚度、海绵组织厚度、主脉厚度、以及主脉中维管束的长度和宽度均与海拔呈显著正相关关系,增幅分别达38.32%、47.30%、47.39%、156.46%、173.51%及337.42%;上表皮厚度、下表皮厚度、叶片紧密度、叶片疏松度、栅海比以及维管束的个数与海拔梯度之间的相关性则均不显著。研究表明,随着海拔升高,草血竭叶片形态结构的改变主要是有利于植物降低蒸腾,增强储水性和提高光合效率。     

Physiology of Eucalyptus plantlets grown photoautotrophically in a scaled-up vessel

Summary Nodal cuttings of Eucalyptus camaldulensis L. plantlets were cultured photoautotrophically (sugar-free nutrient medium and with enriched CO₂ and high photosynthetic photon flux) in a scaled-up vessel (volume 4.0 liters) under forced ventilation (SV-treatment). After 28 d of culture, physiological aspects of the plantlets were compared with plantlets grown photomixotrophically (20 g l⁻¹ sucrose in the medium) in a Magenta vessel (volume 0.4 liters) under natural ventilation (control). In the SV-treatment net photosynthetic rates were enhanced, normal stomatal closing and opening were observed, and the epicuticular leaf-wax content was significantly higher than the control. The anatomical study showed well-organized palisade and spongy mesophyll layers of SV leaves. The SV-treatment also allowed in vitro acclimatization, and after transplanting ex vitro, the transpiration rate and the percent water loss was lower than those of the control and thus the SV plantlets acclimatized easily ex vitro.     

青藏高原不同海拔3种菊科植物叶片结构变化及其生态适应性

采用常规石蜡制片技术和显微观察方法,对分布于青藏高原祁连山东部山地冷龙岭3600—4400 m不同海拔高度的蒲公英(Taraxacum mongolicum Hand-Mazz)、火绒草(Leontopodium leontopodioides(Willd.)Beauv)和美丽风毛菊(Saussurea superba Anthony)的叶片形态解剖结构进行了研究,并探讨了其对海拔高度的响应及生态适应性。结果表明:随海拔高度的升高,3种植物气孔器外拱盖内缘、角质层纹饰、气孔与表皮细胞的位置关系以及上、下表皮气孔器内缘呈现不同的变化趋势;火绒草上、下表皮气孔密度随海拔升高而增加,而蒲公英和美丽风毛菊的气孔密度则降低;3种植物上、下表皮气孔指数随海拔高度的增加均未出现规律性变化;3种植物叶片厚度、上下表皮厚度、上下角质层厚度、栅栏细胞系数均随海拔升高而增加;解剖学指标之间大多呈明显的协同进化;叶片结构,尤其是气孔密度对海拔高度变化表现出较大的可塑性。研究表明3种植物采取不同的响应机制来适应海拔高度的变化,植物对高原环境变化的适应具有多样性。     

Physiological and anatomical changes induced by drought in two olive cultivars (cv Zalmati and Chemlali)

Photosynthetic gas exchange, vegetative growth, water relations and fluorescence parameters as well as leaf anatomical characteristics were investigated on young plants of two Olea europaea L. cultivars (Chemlali and Zalmati), submitted to contrasting water availability regimes. Two-year-old olive trees, grown in pots in greenhouse, were not watered for 2 months. Relative growth rate (RGR), leaf water potential (Ψ_LW) and the leaf relative water content (LWC) of the two cultivars decreased with increasing water stress. Zalmati showed higher values of RGR and LWC and lower decreased values of Ψ_LW than Chemlali, in response to water deficit, particularly during severe drought stress. Water stress also caused a marked decline on photosynthetic capacity and chlorophyll fluorescence. The net photosynthetic rate, stomatal conductance, transpiration rate, the maximal photochemical efficiency of PSII (F _v/F _m) and the intrinsic efficiency of open PSII reaction centres (F′ _v/F′ _m) decreased as drought stress developed. In addition, drought conditions, reduced leaf chlorophyll and carotenoids contents especially at severe water stress. However, Zalmati plants were the less affected when compared with Chemlali. In both cultivars, stomatal control was the major factor affecting photosynthesis under moderate drought stress. At severe drought-stress levels, the non-stomatal component of photosynthesis is inhibited and inactivation of the photosystem II occurs. Leaf anatomical parameters show that drought stress resulted in an increase of the upper epidermis and palisade mesophyll thickness as well as an increase of the stomata and trichomes density. These changes were more characteristic in cv. ‘Zalmati’. Zalmati leaves also revealed lower specific leaf area and had higher density of foliar tissue. From the behaviour of Zalmati plants, with a smaller reduction in relative growth rate, net assimilation rate and chlorophyll fluorescence parameters, and with a thicker palisade parenchyma, and a higher stomatal and trichome density, we consider this cultivar more drought-tolerant than cv. Chemlali and therefore, very promising for cultivation in arid areas.     

赤皮青冈幼苗叶片解剖结构对干旱胁迫的响应

以浙江庆元、湖南洞口和湖南靖州3个种源的赤皮青冈1年生幼苗为材料,采用盆栽称量法控制土壤水分含量,设置土壤相对含水量分别为75%～80%(对照)、55%～60%(轻度干旱胁迫)、45%～50%(中度干旱胁迫)和30%～35%(重度干旱胁迫),研究不同程度干旱胁迫对赤皮青冈幼苗叶片解剖结构和蒸腾作用的影响.结果表明: 干旱胁迫下,3个种源赤皮青冈幼苗叶片的总厚度、上下表皮厚度和栅栏组织厚度均显著降低.栅栏组织厚度与海绵组织厚度比(栅海比)、气孔长度和宽度随干旱程度的增强而显著减小,而气孔密度随干旱胁迫程度的增强而显著增加.干旱处理对叶片主脉厚度无显著影响,木质部厚度的变化因种源而异.叶片结构的变化引起生理功能改变,随着胁迫程度的加强,赤皮青冈幼苗的蒸腾速率显著降低.对照和各干旱处理湖南洞口种源幼苗叶片的下表皮厚度、栅海比、气孔密度显著大于其他2个种源,蒸腾速率显著小于其他2个种源,表明湖南洞口赤皮青冈对干旱胁迫的适应性较强.     

Changes of Leaf Morphological, Anatomical Structure and Carbon Isotope Ratio with the Height of the Wangtian Tree （Parashorea chinensis） in Xishuangbanna, China

Leaf morphological and anatomical structure and carbon isotope ratio （δ^13C） change with increasing tree height. To determine how tree height affects leaf characteristics, we measured the leaf area, specific leaf mass （ratio of leaf mass to leaf area [LMA]）, thickness of the total leaf, cuticle, epidermis, palisade and sponge mesophyll, stomata traits and δ^13C at different heights of Parashorea chinensis with methods of light and scanning electron microscopy （SEM） and isotope-ratio mass spectrometry. The correlation and stepwise regression between tree height and leaf structure traits were carried out with SPSS software. The results showed that leaf structures and δ^13C differed significantly along the tree height gradient. The leaf area, thickness of sponge mesophyll and size of stomata decreased with increasing height, whereas the thickness of lamina, palisade mesophyll, epidermis, and cuticle, ratios of palisade to spongy thickness, density of stomata and vascular bundles, LMA and δ^13C increased with tree height. Tree height showed a significant relationship with all leaf indices and the most significant relationship was with epidermis thickness, leaf area, cuticle thickness, δ^13C. The δ^13C value showed a significantly positive relationship with LMA （R = 0.934）. Our results supported the hypothesis that the leaf structures exhibited more xeromorphic characteristics with the increasing gradient of tree height.     

Interactive effects of elevated CO2 and temperature on the anatomical characteristics of leaves in eleven species

The anatomical features of leaves in 11 species of plants grown in a temperature gradient and a temperature + CO₂ gradient were studied. The palisade parenchyma thickness, the spongy parenchyma thickness and the total leaf thickness were measured and analyzed to investigate the effects of elevated temperature and CO₂ on the anatomical characteristics of the leaves. Our results show that with the increase of temperature, the leaf thickness of C₄ species increased while the leaf thickness of C₃ species showed no constant changes. With increased CO₂, seven out of nine C₃ species exhibited increased total leaf thickness. In C₄ species, leaf thickness decreased. As for the trend on the multi-grades, the plants exhibited linear or non-linear changes. With the increase of temperature or both temperature and CO₂ for the 11 species investigated, leaf thickness varied greatly in different plants (species) and even in different branches on the same plant. These results demonstrated that the effect of increasing CO₂ and temperature on the anatomical features of the leaves were species-specific. Since plant structures are correlated with plant functions, the changes in leaf anatomical characteristics in elevated temperature and CO₂ may lead to functional differences.     

Anatomical responses of leaves of Black Locust (<Emphasis Type="Italic">Robinia pseudoacacia</Emphasis> L.) to urban pollutant gases and climatic factors

This study investigates responses in the leaf anatomy of Black Locust (Robinia pseudoacacia L.) to the atmospheric pollutants, SO₂, NO₂ and O₃ and climate in Tehran. The anatomical variables studied include thickness of the leaf lamina and of its main constituent tissues and the length and density of stomata. We present evidence that, in response to urban air pollution, the spongy mesophyll layer is thinner, the upper cuticle of the leaf thicker and stomatal density and the ratio of palisade parenchyma to spongy parenchyma are increased. Similar responses were also detected in relation to a climatic gradient. Stomatal density and thickness of the leaf lamina and of its mesophyll layer were all higher under warmer drier conditions. This overlap in anatomical response to two very different suites of environmental variables may reflect a functional overlap between mechanisms designed to restrict water loss in dry climates and those that minimize the uptake of toxic gases in polluted habitats.     

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。