环境与植物中硒形态研究进展

本文综述了近年来在环境及植物中的硒形态研究方面的进展。在土壤和水中各种不同形态硒的分布情况与pH值及其它特性之间的关系;植物中各种有机硒化合物的含量,可能的结构和硫素之间的关系,植物对硒的吸收转化过程等等。     

植物硒及其含硒蛋白的研究

硒是植物的有益元素,植物对硒的吸收与外源硒的有效性、硒的形态、植物的种类等有关;硒在植物中主要以有机硒形态存在,HPLC-ICP-MS联用已成为植物体内硒形态鉴定的最常用手段;含硒蛋白是植物体内最主要的有机大分子硒,具有抗肿瘤、抗氧化等多种生物活性。在环境安全和人类健康等方面,富硒植物具有很好的应用价值,所以利用分子生物学手段分析富硒植物的富硒机制,可以为富硒基因的筛选和利用提供理论依据。     

植物体内硒和硫的相互作用

植物体内硒,硫和同化途径类似,硒和硫的相互作用表现在植物中硒和硫的吸收,运输和积累过程以及植物硒甲基化过程中,可影响植物生长发育及品质成分,这种相互作用随植物种类,硒和硫形态,浓度,植物的生育期,以及生长部位的不同而呈现出不同的特点。     

土壤中硒的形态转化及其对有效性的影响研究进展

在对一些典型地区环境硒调研的基础上,系统研究了硒在土壤中形态和价态转化,探讨了硒在土壤固液相中的分配及其与土壤性质的关系;明确了土壤中硒的转化对其生物有效性的影响;系统归纳了硒的形态、土壤性质、植物的品种和种类等影响硒有效性的因素;建立了将形态和价态相结合的土壤硒形态测定方法,并对测定硒有效性的化学浸提法和梯度扩散薄膜技术(DGT)进行了比较。最后,提出了相关研究未来的发展方向。     

植物对土壤硒的吸收转化研究进展

硒是人类、动物和某些微生物必需的微量元素。植物因其具有吸收和转化土壤硒的能力而成为人类饮食补硒的主要来源。提高植物的富硒水平对于人类健康具有重要意义。为更好地了解影响植物富硒效果的因素,综述了土壤中硒的赋存形态,重点阐述了植物对土壤不同形态硒的吸收、迁移和代谢过程,并由此对硒生物强化未来的研究方向进行了展望。     

植物硒生理及与重金属交互的研究进展

硒是一种重要的微量元素,在低浓度时对生物有益,但高浓度时呈现与重金属类似的毒性。植物作为人体硒摄入的主要来源,其硒代谢对于植物硒积累乃至人体硒营养水平十分重要。研究植物硒吸收、代谢和积累机理能指导富硒粮食的生产,是解决人体硒摄入不足/超量问题的有效途径。本文在阐述土壤硒含量、形态、生物有效性及分布的基础上,综述了植物对硒的吸收、代谢机理的研究进展,并讨论了农业生物强化以及遗传育种生物强化等两种硒生物强化的实践方法,以及利用硒生物强化缓解重金属毒性减少积累;最后,提出了植物硒代谢及硒生物强化研究的前沿问题,以期为改善人体硒营养水平提高人体健康状况提供理论和实践依据。     

国产海桑属（Sonneratia Linn．f．）植物的形态解剖

本文描述我国海南岛产的卵叶海桑、海南海桑、杯萼海桑、拟海桑、海桑等５种海桑属植物的叶、花、果实、种子、木材等的形态结构;分析它们的主要特征在各种植物之间的异同．观察结果表明,海南海桑和拟海桑两新种的形态结构特征与其它各种植物有明显的不同,并讨论它们之间的分类关系。     

植物硒吸收转化机制及生理作用研究进展

硒是大多微生物、动物及人类的必要微量元素,但其在植物生长发育中的生理作用至今存在争议.较低浓度硒具有促进植物生长、提高植物耐受能力的功能,而大部分植物在高浓度下表现出中毒现象.随着人类对摄入硒及环境硒污染问题的认识加深,作物硒生物强化与硒污染植物修复问题引起重视,推动了对硒在植物中的吸收积累及代谢调控的研究.近年来对植物硒吸收及转化的研究表明,不同硒水平下植物对硒吸收积累及生理响应存在差异,土壤环境因素对植物硒吸收及转化具有重要影响,对高聚硒植物硒代谢研究逐渐揭示出硒在植物体内的转化过程和调控机理等.本文总结了目前硒生物强化与植物修复方面的研究进展,对环境中硒分布特点、植物硒吸收及其影响因素、植物体内硒转化及其过程调控关键酶,以及硒在植物中的生理作用等进行了综述,并对植物硒生理及分子机制未来研究方向进行展望.     

硒的生物地球化学研究团队介绍

<正>本课题组近10年来致力于土壤-植物体系中硒迁移转化及其生物有效性的研究。对陕西紫阳硒中毒区、青海平安富硒地区和泾惠灌区缺硒地区环境中硒及有效性进行了调研,系统研究了外源硒在土壤中环境过程及其影响硒有效性的因素;建立了形态和价态相结合的土壤硒测定方法,针对性地提出了评价硒有效性的指标,这些研究为硒有效性评价及缺硒地区硒的     

李花粉教授团队介绍

正本团队主要研究硒在土壤-植物系统中的迁移转化以及硒对重金属的解毒作用,建立了硒形态测定与分析技术。近年来在"New Phytologist","Plant and Soil","Journal of Agricultural and Food Chemistry"等刊物上发表多篇论文。相关科研项目有国家自然科学基金"水稻根际硒形态转化及吸收转移机理研究"和"硒缓解水稻累积镉的根际调控机制研究"以及公益性行业(农业)科研专项"优质高效富硒农产品关键技术研究与示范"等。     

Absorption,Distribution and Transformation of Selenium in the Tomato plants

Low concentration of selenium enhanced, but high concentration inhibited the growth of tomato seedlings, the higher the concentration of selenium, the more the inhibition of the growth was. Selenium was absorbed quickly by roots, and conveyed to the other organs. With prolonged time, the degree of increase in the distribution of total selenium content in the plants was in the orders of roots>leaves>stems. From lower to upper the distribution of selenium in the leaves increased progressively. At the flowering and bearing stage, the orders of selenium distribution were roots>fruits>stems>leaves. Only a part of selenium in the plants existed at the state of inorganic selenium, and most changed into organic selenium, and the majority of it was in the form of selenoproteins. 90.89% of its selenoproteins was found in the fruits which was more than that of the other organs of plants. Sulfur starvation enhenced the absorption and transportation of selenium from roots,accelerated the distribution of selenium in the leaves ,and increased the organic selenium content in the roots ,stems and fruits.     

Selenium speciation profiles in selenite-enriched soybean (Glycine Max) by HPLC-ICPMS and ESI-ITMS

Soybean (Glycine Max) plants were grown in soil supplemented with sodium selenite. A comprehensive selenium profile, including total selenium concentration, distribution of high molecular weight selenium and characterization of low molecular weight selenium compounds, is reported for each plant compartment: bean, pod, leaf and root of the Se-enriched soybean plants. Two chromatographic techniques, coupled with inductively coupled plasma mass spectrometry (ICPMS) for specific selenium detection, were employed in this work to analyze extract solutions from the plant compartments. Size-exclusion chromatography revealed that the bean compartment, well-known for its strong ability to make proteins, produced high amounts (82% of total Se) of high molecular weight selenospecies, which may offer additional nutritional value and suggest high potential for studying proteins containing selenium in plants. The pod, leaf and root compartments primarily accumulate low molecular weight selenium species. For each compartment, low molecular weight selenium species (lower than 5 kDa) were characterized by ion-pairing reversed phase HPLC-ICPMS and confirmed by electrospray ionization ion trap mass spectrometry (ESI-ITMS). Selenomethionine and selenocystine are the predominant low molecular weight selenium compounds found in the bean, while inorganic selenium was the major species detected in other plant compartments.     

Influence of selenium on uptake and toxicity of copper and cadmium in pea (Pisum sativum) and wheat (Triticum aestivum)

The detoxifying effect of selenium on animals toxicated with heavy metals is well known. In this study we examine if there is a similar effect in plants. Wheat ( Triticum aestivum L. cv. Sunny) and pea ( Pisum sativum L. cv. Fenomen) were grown for 21 days on a nutrient solution based on the nutrient proportions in healthy plants. Nutrients along with cadmium, copper, selenite, selenate or selenite and selenate in combinations with copper or cadmium were supplied in small amounts with a daily incremental increase of 0.12 (wheat) and 0.20 (pea). The metal and selenium uptake and distribution in the plants as well as the effects on growth were investigated.
The results show that selenium does not reduce the toxicity of heavy metals to plants. Instead, selenium enhances metal uptake and toxicity, especially in peas grown in the presence of metal and selenate. Selenite increased cadmium concentrations of pea roots up to 300% and selenate that of wheat shoots up to 50%.     

植物硒素营养及其机理研究进展

土壤生境中Se的丰缺对植物的Se素营养有重要的影响。本文阐述了植物对Se的吸收,积累和运移过程,概述了Se对植物的毒性,对植物产量和品质影响及其机理研究的进展,在此基础上,提出以研究Se对土壤酶活性影响来解释Se对植物的必要性的新途径。     

Selenium accumulation protects plants from herbivory by Orthoptera via toxicity and deterrence

To investigate whether selenium (Se) accumulation in plants provides a chemical defense against generalist insect herbivores, the feeding preference and performance of a mix of orthopteran species were investigated. The selenium hyperaccumulator Stanleya pinnata and accumulator Brassica juncea were used in herbivory studies in the laboratory, and S. pinnata was also used in a manipulative field experiment. In laboratory studies, both crickets and grasshoppers avoided plants pretreated with selenate, while those given no choice died after eating leaves with elevated Se (447 +/- 68 and 230 +/- 68 microg Se g(-1) DW, respectively). B. juncea has previously been shown to accumulate selenate, while S. pinnata hyperaccumulates methyl-selenocysteine. Thus, these findings demonstrate that both inorganic and organic forms of selenium protect plants from herbivory. Grasshoppers fed S. pinnata contained methylselenocysteine in their midgut and absorbed this form into surrounding tissues. In a manipulative field experiment, methylselenocysteine protected S. pinnata from invertebrate herbivory and increased its long-term survival rate over an entire growth season. * In native habitats of selenium hyperaccumulators, orthopterans represent a major group of insect herbivores. Protection offered by organic selenium accumulation against these herbivores may have promoted the evolution of selenium hyperaccumulation in plants.     

Uptake,distribution, and turnover rates of selenium in barley

The present communication elucidates initially the topographic distribution of selenium in barley grains. Then by the fluorimetric method the uptake of selenium (selenite) in 8–16 d old germinating barley was estimated. Finally by means of⁷⁵Se the anabolic and catabolic rates (turnover) of⁷⁵Se (selenite) was compared. The distribution of selenium in barley was evaluated after microdissection of barley grains. In dried grains the highest concentration was found in husk and pericarp with about 0.6 ppm Se. Then followed Scutellum with 0.4 and 0.3 ppm in embryon. The aleurone layer, embryonic leaves, and initial root did only have 0.2 ppm Se. In order to know more about the uptake and distribution of selenium in 8-d-old barley, the plants were cultivated for a further 8 d in the culture medium with variation in selenite concentration. In roots and leaves, the uptake did not arrive at saturation during the period studied since the dose-response curve increased up to 0.34 mM selenite in the medium, whereas the selenium levels were about 200 ppm in roots and 30 ppm in leaves. However, the uptake was linear, with concentration during 8 d of cultivation up to 0.84 μM selenite for grain and stem. At higher concentrations the dose-response curve diminished its slope. At 0.34 mM selenite the concentration in grain increased to 6.87 ppm and in the stem to 8.13 ppm. The uptake, distribution, and catabolic rate of selenium components in germinating barley were further evaluated by exposing the plants to 0.0492 μCi⁷⁵Se (12.6 μM selenite) for up to 4 d. Then the plants were moved to a selenium deficient medium for further 4 d. Then finally the medium was supplemented with high doses of cold selenite (0.126 mM selenite) for further 4 d. The first third period made it possible to estimate the rate of uptake. It was highest in roots (313 fmol/h/mg dw), i.e., about 10 times those of grains, stems, and leaves. The intermediate period where the barley was transferred to a selenium deficient medium made it possible to estimate the kinetics and eventual sparing mechanisms. The selenium losses were highest for leaves (39%), then followed by roots and stems (22 and 25%, respectively). The losses were lowest in grain with 9% Se losses. The losses were three times more pronounced during the first day than in the following 3 d. These data may argue that the selenium is distributed into different pools and that sparing mechanisms may be in function. The last period, i.e., the chase experiment, revealed the rate of elimination of selenium under conditions with surplus selenium. The catabolic rate was about 10 times faster in roots (169 fmol/h/mg dw) than in grains and about 8 times faster than in leaves.     

Expression of a mouse selenocysteine lyase in Brassica juncea chloroplasts affects selenium tolerance and accumulation

Selenium is an essential nutrient for many organisms, as part of certain selenoproteins. However, selenium is toxic at high levels, which is thought to be due to non-specific replacement of cysteine by selenocysteine leading to disruption of protein function. In an attempt to prevent non-specific incorporation of selenocysteine into proteins and to possibly enhance plant selenium tolerance and accumulation, a mouse selenocysteine lyase was expressed in Brassica juncea (Indian mustard) chloroplasts, the site of selenocysteine synthesis. This selenocysteine lyase specifically breaks down selenocysteine into elemental selenium and alanine. The transgenic cpSL plants showed normal growth under standard conditions. Selenocysteine lyase activity in the cpSL transgenics was up to 6-fold higher than in wild-type plants. The cpSL transgenics contained up to 40% less selenium in protein compared to wild-type plants, indicating that Se flow in the plant was successfully redirected. Surprisingly, the selenium tolerance of the transgenic cpSL plants was reduced, perhaps due to interference of produced elemental selenium with chloroplastic sulphur metabolism. Shoot selenium levels were enhanced up to 50% in the cpSL transgenics, but only during the seedling stage.     

A tale of two toxicities: malformed selenoproteins and oxidative stress both contribute to selenium stress in plants

Background

Despite selenium''s toxicity in plants at higher levels, crops supply most of the essential dietary selenium in humans. In plants, inorganic selenium can be assimilated into selenocysteine, which can replace cysteine in proteins. Selenium toxicity in plants has been attributed to the formation of non-specific selenoproteins. However, this paradigm can be challenged now that there is increasingly abundant evidence suggesting that selenium-induced oxidative stress also contributes to toxicity in plants.

Scope

This Botanical Briefing summarizes the evidence indicating that selenium toxicity in plants is attributable to both the accumulation of non-specific selenoproteins and selenium-induced oxidative stress. Evidence is also presented to substantiate the claim that inadvertent selenocysteine replacement probably impairs or misfolds proteins, which supports the malformed selenoprotein hypothesis. The possible physiological ramifications of selenoproteins and selenium-induced oxidative stress are discussed.

Conclusions

Malformed selenoproteins and oxidative stress are two distinct types of stress that drive selenium toxicity in plants and could impact cellular processes in plants that have yet to be thoroughly explored. Although challenging, deciphering whether the extent of selenium toxicity in plants is imparted by selenoproteins or oxidative stress could be helpful in the development of crops with fortified levels of selenium.     

Speciation, Distribution, and Bioavailability of Soil Selenium in the Tibetan Plateau Kashin–Beck Disease Area—A Case Study in Songpan County, Sichuan Province, China

To clarify the relationship between the soil selenium distribution and its bioavailability with the distribution of Kashin–Beck disease (KBD) endemic areas on the eastern edge of the Tibetan Plateau, samples of natural soil (0–20 cm), cultivated topsoil, and main crops of the region (highland barley) were collected at different altitudes according to topographical and geomorphological features in both KBD and non-KBD areas of Songpan County. These samples were used for determination and analysis of total selenium content in soil and highland barley and available selenium that can be absorbed and utilized by plants. The results showed that the average total selenium content of natural and cultivated topsoil in KBD areas was lower than that in non-KBD areas (natural soil, P?=?0.061; cultivated soil, P?=?0.002), which is in agreement with the geographical distributions of selenium in other KBD-affected areas. However, the total soil selenium content exhibits certain micro-spatial distribution features, namely, the total selenium content in some endemic areas was significantly higher than that of non-KBD areas. This result was contrary to the general distribution that total selenium content in a KBD-affected area is lower than that in a non-KBD area. We further studied the extraction rate and content of soil selenium in six different fractions. The results indicated that the content and extraction rate of available selenium in KBD-affected areas were significantly lower than those in non-KBD areas. There is a distinct positive correlation between plant-available selenium and highland barley selenium (r?=?0.875, P?=?0.001) and a distinct negative correlation with altitude (r?=??0.801, P?=?0.010). Therefore, in KBD endemic areas, the selenium content in crops decreases as the available selenium content in soil decreases and is closely related to the geographical environment features (such as altitude and precipitation). These results suggest that the soil available selenium and ecological features are important factors that restrict the dietary selenium flux for residents in KBD endemic areas of the Tibetan Plateau, providing a theoretical and experimental basis for implementing agricultural measures to regulate the ecological cycle of the selenium flux in the KBD endemic area.