三叶鬼针草（Bidens pilosa L.）对重金属Cd、Pb胁迫的响应与修复潜能研究
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摘要
随着人类社会工业化和城镇化进程的广泛而深刻推进,物质文明达到空前繁荣,实现了当代人前所未有的便利和富足；同时也给生态环境造成了沉重负担,如重金属、永久性有机物污染和石漠化、荒漠化等。因为具有隐蔽性强、半衰期长、毒性大、难清除和易生物放大进入食物链危害人类健康等特点,重金属污染成为全球生态环境最大的威胁之一。重金属污染环境的修复方法很多,主要有物理修复、化学修复和生物修复,其中生物修复中的植物修复因具有成本低、环境友好、修复过程持久,尤其是能与当前重金属污染面J广一的特点相适合而被认为是最有发展前途的方法。植物修复的核心是超富集植物。本文以三叶鬼针草(B. pilosa L.)为研究对象,综合考察其在Cd、Pb胁迫下的光合生理、质膜氧化、有机酸及重金属形态、渗透调节物质、抗氧化酶、根际有机酸和营养素等响应,评估其对Cd、Pb污染土壤的修复潜能.最后以三叶鬼针草叶片诱导出毛状根,初步探索了其对Cd、Pb污染水体的修复潜能。通过实验本文主要得出如下结论：
1.光合生理对Cd、Pb (?)协迫的响应。单一胁迫时,叶绿素含量、净光合率、气孔导度和蒸腾速率随单一Cd、Pb胁迫浓度增加先升后降,胞间CO2浓度微弱增加。Cd-Pb复合胁迫,表现出浓度加和效应,低浓度复合胁迫促进光合作用,高浓度复合胁迫抑制光合作用。Cd、Pb胁迫通过对光抑制(Fv/Fm)、非辐射能量、光合电子传递效率和PS Ⅱ开放等的抑制或促进,使光合机构完整性与运作状况受损害的同时也激活了光合机构的自我修复。Cd、Pb胁迫促进或抑制光合作用的主要原因是促进或抑制叶绿素合成。
2.质膜氧化对Cd、Pb胁迫的响应。Cd、Pb胁迫导致三叶鬼针草植株内超氧阴离子的积累。随单—Cd、Pb胁迫浓度增加,体内超氧阴离子先增后减。低、中浓度Pb促进Cd-Pb复合胁迫植株超氧阴离子的积累,高浓度Pb抑制积累。Cd对Cd-Pb复合胁迫植株超氧阴离子积累无影响。植株体内丙二醛含量变化与超氧阴离子变化基本一致；脂氧合酶活性随重金属胁迫增加而增加,有组织器官差异。随Cd、Pb胁迫浓度增加,三叶鬼针草不饱和脂肪酸和长链饱和脂肪酸含量增加,并在高胁迫浓度维持高含量水平。同时构成膜脂肪酸不饱和系数(DBI)和不饱和脂肪酸含量(UFA%)在高胁迫浓度下均维持高水平。上述结果表明,在Cd、Pb胁迫下三叶鬼针草可以通过提高膜脂肪酸不饱和程度提高质膜的流动性,通过增加长链饱和脂肪酸来维持细胞膜结构的稳定性,从而提高对重金属胁迫导致氧伤害的忍耐能力。
3.渗透调节物质与抗氧化酶对Cd、Pb胁迫的响应。Cd、Pb(?)办迫均能导致三叶鬼针草叶片中可溶性糖和脯氨酸积累。积累程度与金属种类和胁迫浓度有关。随Cd胁迫浓度增加,叶片中可溶糖先增后减；随Pb肋迫浓度增加,则叶片中可溶糖持续增加。可溶性蛋白含量随Cd、Pb单一胁迫浓度增加而先增后降；相对于单一Cd胁迫,Cd-Pb复合胁迫能使可溶性蛋白和可溶性糖含量降低积累。随Cd、Pb及Cd-Pb胁迫浓度增加,叶片内SOD、CAT和POD变化没有一致性。随Cd、Pb胁迫浓度增加,SOD活性逐渐增大；Cd-Pb复合胁迫协同促进SOD活性提高。CAT活性随单一Cd胁迫浓度增加先升后降；随Pb胁迫浓度增加,CAT活性持续增加;高浓度Cd与Pb复合胁迫促进CAT活性提高。POD活性随单一Cd胁迫浓度增加先升后降,随单一Pb胁迫浓度增加持续缓慢升高。Cd-Pb复合胁迫在一定程度上提高了POD活性且与Pb浓度有关。Cd、Pb胁迫下,三叶鬼针草主要通过调节植物叶片脯氨酸、可溶性糖和可溶性蛋白含量维持Cd、Pb胁迫下的体内渗透平衡,调节抗氧化酶SOD、CAT和POD活性,提高对重金属诱导的氧伤害忍耐能力,从而提高对Cd、Pb胁迫的忍耐能力。
4.体内有机酸、重金属形态及分布对Cd、Pb胁迫的响应。Cd、Pb胁迫促进草酸在体内累积。同时还能调节植株根系和叶片中苹果酸、柠檬酸、醋酸和琥珀酸合成使进入植株体内重金属被螯合而毒性降低。三叶鬼针草体内Cd主要以与蛋白质和果胶结合形态(Fr3)存在,低肋迫浓度有利于Cd从根系向上转运,高浓度抑制Cd向叶片转运；Pb主要以草酸盐形态(Fr5)存在,高肋迫浓度下有利于Pb在叶片的积累。复合胁迫下Pb促进Cd的Fr3形态向Fr4和Fr5转变,把活性较高的形态向相对活性较低形态转变,表现的效应是缓解Cd毒害的效应：而Cd促进活性较低的Fr5的Pb向相对活性较高Fr3、Fr4、Fr2和Frl等形态转变,表现的效应是协同加和毒性效应。Cd、Pb在亚细胞分布以细胞壁(F1)与可溶部分(F4)为主,叶绿体或质体(F2)和其他细胞器(F3)含量很低。Cd-Pb复合肋迫,Pb能促进低浓度Cd肋迫植株的细胞壁和可溶部分含量与百分比的增加：Cd能促进叶绿体或质体内Pb的百分比和含量增加而细胞壁和可溶部分的Pb减小。表明,三叶鬼针草通过增加体内草酸、苹果酸和柠檬酸的含量提高对重金属的螯合能力、以活性较低的果胶和蛋白质结合态与草酸盐形态存贮和亚细胞分布等策略降低进入植物体内Cd、Pb毒性而提高植物对重金属忍耐能力。
5.根际土壤对Cd、Pb肋迫的响应。Cd、Pb胁迫下,根际均能积累大量草酸。随Cd胁迫浓度增加,草酸的含量先增后降,随Pb胁迫浓度增加持续升高。在低浓度Cd与Pb复合胁迫时,根际草酸含量随Pb浓度增加而上升；在高浓度Cd与Pb复合胁迫时,随Pb浓度增加先降后升。根际Cd以残渣态为主,并随Cd胁迫浓度增加而增加；Pb主要是以碳酸盐结合态,其次是Fe-Mn氧化结合态存在。Cd-Pb复合胁迫时Pb促进可交换态和碳酸盐结合态Cd的比例增加,且浓度为1000mg/kg时,促进效应最明显；Cd促进残渣态Pb增加而使碳酸盐结合态Pb减少。Pb是促进Cd从活性较低形态(残渣态)向活性较高形态(可交换态和碳酸盐结合态)转换；Cd则是促进Pb从活性较高形态(碳酸盐结合态)向较低形态(残渣态)转换。Cd、Pb肋迫对三叶鬼针草根际营养元素影响不明显,仅对K影响和Cd-Pb复合肋迫N、P、K的含量降低。三叶鬼针草可以通过分泌提高根际草酸含量与重金属结合形成不利于迁徙的草酸盐形态影响重金属根际生物活性,从而调节重金属活性系数与移动系数,控制重金属进入植物体的量与时间,提高植物对重金属的忍耐能力。
6.植株对Cd、Pb污染土壤的修复潜能。在一个生命周期内,植物地上部和根系重金属含量随生命进程的发展逐渐增加,这与生物量积累一致。在胁迫90d,植物处于旺盛生长时期,植物体内重金属含量和生物量达到最大,随着生命周期从营养生长向生殖生长发展,植物体内重金属含量维持较高水平并略下降。在单一胁迫时,植株体内重金属含量随浓度增加而增加。Cd-Pb复合胁迫,一般低、中浓度Pb有促进植物对低Cd吸收与转运,而高浓度Pb则严重抑制植株对Cd的吸收；高浓度Pb与高浓度Cd复合胁迫,Pb是抑制Cd向地上部转运。Cd对于植物吸收和转运Pb的效应影响不显著,当植株体内Cd含量高的实验组Pb含量相对比较低；Cd含量较低的组,Pb含量相对较高,基本维持重金属总量保持不变。在整个生命周期内,三叶鬼针草对Cd、Pb的富集能力与转运能力基本是随生命进程的发展呈现先升后降趋势。三叶鬼针草生命周期一般为180d,利用三叶鬼针草修复重金属污染土壤,最佳采收时期应在植物从营养生长向生殖生长过渡时期,即90-120d内。
7.三叶鬼针草毛状根对Cd、Pb污染水体的修复。低浓度Cd、Pb促进毛状根生物量积累,高浓度则抑制积累。Cd-Pb复合胁迫,低、中浓度Pb促进三叶鬼针草毛状根生物量积累；高浓度Pb与低浓度Cd、低中高浓度Pb与高浓度Cd复合胁迫均抑制生物量积累。胁迫浓度与胁迫时间协同作用促进毛状根对Cd、Pb的生物富集。毛状根在不同的胁迫浓度下均有不同的最大吸收范围。短期胁迫,高胁迫浓度有利于毛状根对重金属的富集,当达到最大吸收值后,随胁迫时间延长,富集能力逐渐减弱；低、中胁迫浓度下,随胁迫时间延长,逐渐达到吸收最大范围并维持到胁迫期末。Cd-Pb复合胁迫,低、中Pb促进毛状根对Cd富集,高浓度Pb则抑制Cd的富集;低浓度Cd仅对低浓度Pb有微弱促进积累效应,其余均抑制Pb富集。单一Cd、Pb胁迫,三叶鬼针草毛状根体内最大Cd含量2532mg/kg,最大Pb含量2011mg/kg; Cd-Pb复合胁迫,Cd的最大含量为2223mg/kg, Pb最大含量为1960mg/kg。Cd-Pb复合,Pb对Cd有促进效应,Cd对Pb为抑制效应。细胞壁固定作用和液泡区室化是三叶鬼针草毛状根忍耐重金属Cd、Pb胁迫的机制之一。
With the expansion and further enhancement of industrialization and urbanlization of human society, material civilization has reached its highest peak, thus it can provide human beings with more convenience and affluence that has never seen before. Meanwhile, more and more emergent situations were produced in the ecological environment, such as heavy metal pollution, persistent organic pollutants and desertification. With its ecological toxicology characteristic of strong concealment, long half-life, immense toxicity, it's hard to be eliminated and easy to be biological signifying and inyade the food chain, heavy metal pollution has become one of the biggest threat to ecological environment in the world. There are many ways to removal heavy metals from the contaminated environment, which mainly based on three kinds of remediation technologies including physical, chemical and bioremediation. Currently, phytoremediation is regarded as the most promising method because of its low cost, environmental friendly, enduring restoration process and its suitability for the large scale of heavy metal pollution in current situation. The key of phytoremediation is hyperaccumulator plants. Taking B. pilosa L. as research object, we studied the responding mechanisms of photosynthetic physiology, plasma membrane peroxidation, organic acid and chemical forms, osmotic regulation substances and antioxidant enzymes, organic acids and nutrient elements in the rhizosphere soil to B. pilosa under the stresses caused by cadmium(Cd)and lead(Pb)in order to estimate the phytoremediation's potential abilities to restore Cd, Pb pollution. We still explore its restoration to Cd and Pb in polluted water by using the induced hairy roots from leaves of B. pilosa. The followings are the main conclusions of this paper.
1. The response of photosynthesis by Cd and Pb. Content of chlorophyll, net photosynthesis, stomatal conductance and water vapor rate of B. pilosa has the trend of increase at the beginning and then decreas with the increase of the density of the stress of Cd or Pb, and the intercellular CO2concentration keeps weak increase. The compound stress of Cd-Pb shows addictive effect, in other words, low compound density of stress of Cd and Pb promotes photosynthesis, and high compound density restrains photosynthesis. Cd, Pb and Cd-Pb stresses destroy the compintegrality of photosynthetic apparatus and activate the self-restoration of photosynthetic apparatus by restraining the light and restraining or promoting the non-radicative energy, the transimiting rate of photosynthetic electricity and the opening of PS Ⅱ. Cd and Pb stress can promote or restrain photosynthesis mainly because of their promoting or restraining the composition of chlorophyll, thus result in the increase or decline of the content of chlorophyll.
2. Cd, Pb and Cd-Pb stresses all can lead to the accumulation of superoxide radicals in B. pilosa. By increase the density of single Cd or Pb stress, the content of superoxide radicals increases at the beginning and then decreases; low and middle concentration of Pb stress promotes the accumulation of superoxide radicals in Cd-Pb stressed plants and high concentration of Pb restrains the acuumulation. Cd stress has no influence on the accumulation of superoxide radicals in Cd-Pb stressed plants. The variation of the amounts of malondialdehyde in plants primarily is the same as that of superoxide radicals. The activity of lipoxygenase increases with the increase of heavy metal stress but different among organs of the plants. With the increase of Cd or Pb, unsaturated fatty acids and long chain saturated fatty acids increase and keep in high level with high metal stress. Both Double band index(DBI)and Unsaturated fatty acids to percentage(UFA%)keep in high level with high density stress too. That means in the station of Cd or Pb stress, B. pilosa can enhance the fluidity of membrane by increase the unsaturated degree of fatty acids and maintain the stableness of the membrane structure by increase the content of the long chain saturated fatty acids. So as to improve their ability of endure the oxygen destruction caused by heavy metal stress.
3. Cd, Pb and Cd-Pb stresses can lead to the accumulation of soluble sugar and proline in the leaves of the B. pilosa. The degree of the accumulation differs from one to another due to different kinds of heavy metal stress and the plant's senstineness to the concentration. With the increase of Cd concentration stress, soluble sugar in leave of B. pilosa first increases and then decreases. The amount increase continuingly with the increase of Pb. Soluble proteins in B. pilosa increase at the beginning and then decrease with the increase of Cd or Pb stress. The Cd-Pb compound stress can promote the contents decreased of soluble protein and soluble protein in B. pilosa. With the increase of Cd. Pb or Cd-Pb stress, the activity of main antioxidant enzyme SOD. CAT and POD do not show any uniformity. With the increase of Cd. Pb or Cd-Pb compound stress, the activity of SOD in leaves of B. pilosa gradually increases. With the increase of Cd stress, the activity of CAT in B. pilosa increases first and then decreases. With the increase of Pb stress, the activity of CAT in B. pilosa continuingly increases. Under the condition of Cd-Pb compound stress, high concentration of stress promotes the increase of the activity of CAT in B. pilosa. With the increase of Cd stress, the activity of POD in B. pilosa increases at first and then decreases. And the activity of POD increases gradually in leaves of the paint with increase of Pb stress. Cd-Pb compound stress can slightly promote the activity of POD in leaves of plant and it relies on the concentration of Pb. Under the stress of Cd, Pb or Cd-Pb, B. pilosa can maintain the osmosis balance mainly by adjusting the content of proline, soluble sugar and soluble protein in the leaves and adjust the activity of SOD. CAT and POD to improve their ability in order to endure the antioxidant injury by heavy metal. As a result, they can enhance their ability to endure Cd and Pb stress.
4. Cd, Pb stresses promote the accumulation of oxalate in B. pilosa. They also chelate the absorbed heavy metal by adjusting the composition of malic acid, citric acid, acetic acid and amber acid in the roots and leaves of plants to reduce the toxicity of the heavy metal. In plants' bodies. Cd mainly exists in B. pilosa with the form of Fr3, which combined with protein and pectin. In this situation, low stress is beneficial to the transmission of Cd from roots to other positions and high stress restian the transport of Cd to leaves; Pb mainly exists in form of Fr5, a kind of oxalate, and high stress promotes the accumulation of Pb in the leaves. In compound stress conditions. Pb promotes Fr3turning to Fr4or Fr5to endure the contamination. Compared with Fr3, Fr4or Fr5is inactive. But Cd promotes Fr5to turn to active forms of Fr3, Fr4, Fr2and Fr1resulting to more toxicity effects. Cd and Pb mainly distribute in the subcellulars in form of cytoderm(F1)and dissoluvable parts(F4), but the content of chloroplast or plastid(F2)and other organelles is sparse. Pb can promote the increase of the content and proportion of low density Cd stressed plants' cytoderm and dissoluvable parts in case of Cd-Pb compound stress Cd can promote the increase of the proportion and content of Pb in chloroplasts or plastids, but reduce the content and proportion of cytoderms and dissoluvable parts. The plants enhance their ability of chelating heavy metals and reduce the toxicity of absorbed Cd and Pb by strategies of increase oxalate, malic acid and citric acid. They also increase the inert chemical forms of heavy metal in the subcellular distribution in order to enhance their ability to endure Cd and Pb stress.
5. In the case of Cd, Pb and Cd-Pb stresses, B. pilosa can accumulate large amount of axalate in rhizosphere. With the increase of Cd stress, the content of oxalate increases first and then decreases, while its content continuingly increases with increase of the density of Pb stress. Under the condition of Cd-Pb compound stress. The content of oxalate raises with the increase of Pb in low density Cd-Pb compound stress but increases first and then decreases in high density Cd-Pb compound stress. Cd exists mainly in the form of residual and its content increases with the increase of Cd stress. Pb mainly exists in the form of carbonate combination and Fe-Mn oxygenized combination take second place. Under the condition of Cd-Pb compound stress, increase of Pb can promote the Cd increase in changeable and carbonate combination form. The promotion is most obvious when adding Pb is1000mg/kg. Cd can promote residual Pb increase and M3of Pb decrease. Pb promote Cd transforming from less active form(M5)to active form(M3)but Cd promote Pb transforming from active form(M2)to less active form(M5). Cd or Pb has no obvious influence on the rhizosphere nutrition elements but K in the plant. But Cd-Pb compound stress decrease the accumulation of N, P and K. B. pilosa can enhance secreting oxalic acid to combinate heavy metal in order to affect the bioactivity of heavy metal in roots. As a result, they adjust the activity ratio and mobility ratio of the heavy metal and control the amount and time of the absorbed heavy metal, so that plants can enhance their ability to endure heavy metal.
6. Within a life period of B. pilosa, content of heavy metal in upper ground parts and roots gradually increases with the growing of plants, which consistent with biomass. After90days' stressing, when growth of the plants reaches their climax, the content of heavy metal and biomass in plants achieve the largest scale. The amount of heavy metal maintains in their maximum or slightly declines with life period changes from nutritive growth to reproductive growth. Under the condition of single heavy metal stress, the content of heavy metal in plants increases with the increase of the stress. Generally, in Cd-Pb compound situation, low and moderate density of Pb can promote the absorbing and transporting of low density Cd, but high density Pb seriously restrains the absorbing of Cd. In high density Pb-Cd compound stress, Pb restrains the plants to transmit Cd to upper ground parts but Cd shows no obvious influence on the absorbing and transporting of Pb. Generally, in a test group, when the content of Cd is large, the content of Pb is relatively small and if the content of Cd is small, the content of Pb is large, but the total amount of heavy metal remains invariablenes. Within life period of the plants, they show ability to accumulate and transport heavy metal, which increase at the beginning and then decreasing with the growth of the plants. In generally, the life span of B. pilosa is180days. So the best time to collect plants is90-120days for B. pilosa to phytoextract the heavy metal in polluted soil in excellent performance.
7. Low density of Cd or Pb promotes the accumulation of biomass in hairy roots of B. pilosa but high density restrains. In case of Cd-Pb compound stress, low and moderate density of Pb stresses promote accumulation of biomass; while other density Pb compound to high Cd stress restrain the accumulation of biomass. Under the condition of Cd or Pb stress, the hairy roots' accumulation of Cd and Pb is the additive result of the density and time of stressing. Hairy roots have different maximum absorbing amount in different density of stresses. In short period stresses, high density stress avails the hairy roots to accumulate heavy metals. Under the condition of low and high density stress, the absorbing of heavy metal gradually approaches peak and then maintains to the end of the stress. In case of Cd-Pb compound stress, low and moderate density of Pb promotes the hairy roots to accumulate Cd, while high density of Pb restrains the accumulation; low density of Pb stress has weak promotion merely to the accumulation of low density of Pb but it restrains the accumulation of Pb in the case of other varieties of density. Under the condition of Cd or Pb stress, the maximum contents of Cd or Pb in the hairy roots are2532mg/kg and2011mg/kg respectively. Under the condition of Cd-Pb compound stress, the maximum contents of Cd and Pb are2223mg/kg and1960mg/kg respectively. Pb stress has the effect of promoting the absorbing of Cd while Cd has the effect of restraining the absorbing of Pb. Compartmentalization and fixed holding of detoxification mechanisms of heavy metal by cell wall vacuole are the strategies of the hairy roots' endurance to Cd and Pb stress.
1.光合生理对Cd、Pb (?)协迫的响应。单一胁迫时,叶绿素含量、净光合率、气孔导度和蒸腾速率随单一Cd、Pb胁迫浓度增加先升后降,胞间CO2浓度微弱增加。Cd-Pb复合胁迫,表现出浓度加和效应,低浓度复合胁迫促进光合作用,高浓度复合胁迫抑制光合作用。Cd、Pb胁迫通过对光抑制(Fv/Fm)、非辐射能量、光合电子传递效率和PS Ⅱ开放等的抑制或促进,使光合机构完整性与运作状况受损害的同时也激活了光合机构的自我修复。Cd、Pb胁迫促进或抑制光合作用的主要原因是促进或抑制叶绿素合成。
2.质膜氧化对Cd、Pb胁迫的响应。Cd、Pb胁迫导致三叶鬼针草植株内超氧阴离子的积累。随单—Cd、Pb胁迫浓度增加,体内超氧阴离子先增后减。低、中浓度Pb促进Cd-Pb复合胁迫植株超氧阴离子的积累,高浓度Pb抑制积累。Cd对Cd-Pb复合胁迫植株超氧阴离子积累无影响。植株体内丙二醛含量变化与超氧阴离子变化基本一致；脂氧合酶活性随重金属胁迫增加而增加,有组织器官差异。随Cd、Pb胁迫浓度增加,三叶鬼针草不饱和脂肪酸和长链饱和脂肪酸含量增加,并在高胁迫浓度维持高含量水平。同时构成膜脂肪酸不饱和系数(DBI)和不饱和脂肪酸含量(UFA%)在高胁迫浓度下均维持高水平。上述结果表明,在Cd、Pb胁迫下三叶鬼针草可以通过提高膜脂肪酸不饱和程度提高质膜的流动性,通过增加长链饱和脂肪酸来维持细胞膜结构的稳定性,从而提高对重金属胁迫导致氧伤害的忍耐能力。
3.渗透调节物质与抗氧化酶对Cd、Pb胁迫的响应。Cd、Pb(?)办迫均能导致三叶鬼针草叶片中可溶性糖和脯氨酸积累。积累程度与金属种类和胁迫浓度有关。随Cd胁迫浓度增加,叶片中可溶糖先增后减；随Pb肋迫浓度增加,则叶片中可溶糖持续增加。可溶性蛋白含量随Cd、Pb单一胁迫浓度增加而先增后降；相对于单一Cd胁迫,Cd-Pb复合胁迫能使可溶性蛋白和可溶性糖含量降低积累。随Cd、Pb及Cd-Pb胁迫浓度增加,叶片内SOD、CAT和POD变化没有一致性。随Cd、Pb胁迫浓度增加,SOD活性逐渐增大；Cd-Pb复合胁迫协同促进SOD活性提高。CAT活性随单一Cd胁迫浓度增加先升后降；随Pb胁迫浓度增加,CAT活性持续增加;高浓度Cd与Pb复合胁迫促进CAT活性提高。POD活性随单一Cd胁迫浓度增加先升后降,随单一Pb胁迫浓度增加持续缓慢升高。Cd-Pb复合胁迫在一定程度上提高了POD活性且与Pb浓度有关。Cd、Pb胁迫下,三叶鬼针草主要通过调节植物叶片脯氨酸、可溶性糖和可溶性蛋白含量维持Cd、Pb胁迫下的体内渗透平衡,调节抗氧化酶SOD、CAT和POD活性,提高对重金属诱导的氧伤害忍耐能力,从而提高对Cd、Pb胁迫的忍耐能力。
4.体内有机酸、重金属形态及分布对Cd、Pb胁迫的响应。Cd、Pb胁迫促进草酸在体内累积。同时还能调节植株根系和叶片中苹果酸、柠檬酸、醋酸和琥珀酸合成使进入植株体内重金属被螯合而毒性降低。三叶鬼针草体内Cd主要以与蛋白质和果胶结合形态(Fr3)存在,低肋迫浓度有利于Cd从根系向上转运,高浓度抑制Cd向叶片转运；Pb主要以草酸盐形态(Fr5)存在,高肋迫浓度下有利于Pb在叶片的积累。复合胁迫下Pb促进Cd的Fr3形态向Fr4和Fr5转变,把活性较高的形态向相对活性较低形态转变,表现的效应是缓解Cd毒害的效应：而Cd促进活性较低的Fr5的Pb向相对活性较高Fr3、Fr4、Fr2和Frl等形态转变,表现的效应是协同加和毒性效应。Cd、Pb在亚细胞分布以细胞壁(F1)与可溶部分(F4)为主,叶绿体或质体(F2)和其他细胞器(F3)含量很低。Cd-Pb复合肋迫,Pb能促进低浓度Cd肋迫植株的细胞壁和可溶部分含量与百分比的增加：Cd能促进叶绿体或质体内Pb的百分比和含量增加而细胞壁和可溶部分的Pb减小。表明,三叶鬼针草通过增加体内草酸、苹果酸和柠檬酸的含量提高对重金属的螯合能力、以活性较低的果胶和蛋白质结合态与草酸盐形态存贮和亚细胞分布等策略降低进入植物体内Cd、Pb毒性而提高植物对重金属忍耐能力。
5.根际土壤对Cd、Pb肋迫的响应。Cd、Pb胁迫下,根际均能积累大量草酸。随Cd胁迫浓度增加,草酸的含量先增后降,随Pb胁迫浓度增加持续升高。在低浓度Cd与Pb复合胁迫时,根际草酸含量随Pb浓度增加而上升；在高浓度Cd与Pb复合胁迫时,随Pb浓度增加先降后升。根际Cd以残渣态为主,并随Cd胁迫浓度增加而增加；Pb主要是以碳酸盐结合态,其次是Fe-Mn氧化结合态存在。Cd-Pb复合胁迫时Pb促进可交换态和碳酸盐结合态Cd的比例增加,且浓度为1000mg/kg时,促进效应最明显；Cd促进残渣态Pb增加而使碳酸盐结合态Pb减少。Pb是促进Cd从活性较低形态(残渣态)向活性较高形态(可交换态和碳酸盐结合态)转换；Cd则是促进Pb从活性较高形态(碳酸盐结合态)向较低形态(残渣态)转换。Cd、Pb肋迫对三叶鬼针草根际营养元素影响不明显,仅对K影响和Cd-Pb复合肋迫N、P、K的含量降低。三叶鬼针草可以通过分泌提高根际草酸含量与重金属结合形成不利于迁徙的草酸盐形态影响重金属根际生物活性,从而调节重金属活性系数与移动系数,控制重金属进入植物体的量与时间,提高植物对重金属的忍耐能力。
6.植株对Cd、Pb污染土壤的修复潜能。在一个生命周期内,植物地上部和根系重金属含量随生命进程的发展逐渐增加,这与生物量积累一致。在胁迫90d,植物处于旺盛生长时期,植物体内重金属含量和生物量达到最大,随着生命周期从营养生长向生殖生长发展,植物体内重金属含量维持较高水平并略下降。在单一胁迫时,植株体内重金属含量随浓度增加而增加。Cd-Pb复合胁迫,一般低、中浓度Pb有促进植物对低Cd吸收与转运,而高浓度Pb则严重抑制植株对Cd的吸收；高浓度Pb与高浓度Cd复合胁迫,Pb是抑制Cd向地上部转运。Cd对于植物吸收和转运Pb的效应影响不显著,当植株体内Cd含量高的实验组Pb含量相对比较低；Cd含量较低的组,Pb含量相对较高,基本维持重金属总量保持不变。在整个生命周期内,三叶鬼针草对Cd、Pb的富集能力与转运能力基本是随生命进程的发展呈现先升后降趋势。三叶鬼针草生命周期一般为180d,利用三叶鬼针草修复重金属污染土壤,最佳采收时期应在植物从营养生长向生殖生长过渡时期,即90-120d内。
7.三叶鬼针草毛状根对Cd、Pb污染水体的修复。低浓度Cd、Pb促进毛状根生物量积累,高浓度则抑制积累。Cd-Pb复合胁迫,低、中浓度Pb促进三叶鬼针草毛状根生物量积累；高浓度Pb与低浓度Cd、低中高浓度Pb与高浓度Cd复合胁迫均抑制生物量积累。胁迫浓度与胁迫时间协同作用促进毛状根对Cd、Pb的生物富集。毛状根在不同的胁迫浓度下均有不同的最大吸收范围。短期胁迫,高胁迫浓度有利于毛状根对重金属的富集,当达到最大吸收值后,随胁迫时间延长,富集能力逐渐减弱；低、中胁迫浓度下,随胁迫时间延长,逐渐达到吸收最大范围并维持到胁迫期末。Cd-Pb复合胁迫,低、中Pb促进毛状根对Cd富集,高浓度Pb则抑制Cd的富集;低浓度Cd仅对低浓度Pb有微弱促进积累效应,其余均抑制Pb富集。单一Cd、Pb胁迫,三叶鬼针草毛状根体内最大Cd含量2532mg/kg,最大Pb含量2011mg/kg; Cd-Pb复合胁迫,Cd的最大含量为2223mg/kg, Pb最大含量为1960mg/kg。Cd-Pb复合,Pb对Cd有促进效应,Cd对Pb为抑制效应。细胞壁固定作用和液泡区室化是三叶鬼针草毛状根忍耐重金属Cd、Pb胁迫的机制之一。
With the expansion and further enhancement of industrialization and urbanlization of human society, material civilization has reached its highest peak, thus it can provide human beings with more convenience and affluence that has never seen before. Meanwhile, more and more emergent situations were produced in the ecological environment, such as heavy metal pollution, persistent organic pollutants and desertification. With its ecological toxicology characteristic of strong concealment, long half-life, immense toxicity, it's hard to be eliminated and easy to be biological signifying and inyade the food chain, heavy metal pollution has become one of the biggest threat to ecological environment in the world. There are many ways to removal heavy metals from the contaminated environment, which mainly based on three kinds of remediation technologies including physical, chemical and bioremediation. Currently, phytoremediation is regarded as the most promising method because of its low cost, environmental friendly, enduring restoration process and its suitability for the large scale of heavy metal pollution in current situation. The key of phytoremediation is hyperaccumulator plants. Taking B. pilosa L. as research object, we studied the responding mechanisms of photosynthetic physiology, plasma membrane peroxidation, organic acid and chemical forms, osmotic regulation substances and antioxidant enzymes, organic acids and nutrient elements in the rhizosphere soil to B. pilosa under the stresses caused by cadmium(Cd)and lead(Pb)in order to estimate the phytoremediation's potential abilities to restore Cd, Pb pollution. We still explore its restoration to Cd and Pb in polluted water by using the induced hairy roots from leaves of B. pilosa. The followings are the main conclusions of this paper.
1. The response of photosynthesis by Cd and Pb. Content of chlorophyll, net photosynthesis, stomatal conductance and water vapor rate of B. pilosa has the trend of increase at the beginning and then decreas with the increase of the density of the stress of Cd or Pb, and the intercellular CO2concentration keeps weak increase. The compound stress of Cd-Pb shows addictive effect, in other words, low compound density of stress of Cd and Pb promotes photosynthesis, and high compound density restrains photosynthesis. Cd, Pb and Cd-Pb stresses destroy the compintegrality of photosynthetic apparatus and activate the self-restoration of photosynthetic apparatus by restraining the light and restraining or promoting the non-radicative energy, the transimiting rate of photosynthetic electricity and the opening of PS Ⅱ. Cd and Pb stress can promote or restrain photosynthesis mainly because of their promoting or restraining the composition of chlorophyll, thus result in the increase or decline of the content of chlorophyll.
2. Cd, Pb and Cd-Pb stresses all can lead to the accumulation of superoxide radicals in B. pilosa. By increase the density of single Cd or Pb stress, the content of superoxide radicals increases at the beginning and then decreases; low and middle concentration of Pb stress promotes the accumulation of superoxide radicals in Cd-Pb stressed plants and high concentration of Pb restrains the acuumulation. Cd stress has no influence on the accumulation of superoxide radicals in Cd-Pb stressed plants. The variation of the amounts of malondialdehyde in plants primarily is the same as that of superoxide radicals. The activity of lipoxygenase increases with the increase of heavy metal stress but different among organs of the plants. With the increase of Cd or Pb, unsaturated fatty acids and long chain saturated fatty acids increase and keep in high level with high metal stress. Both Double band index(DBI)and Unsaturated fatty acids to percentage(UFA%)keep in high level with high density stress too. That means in the station of Cd or Pb stress, B. pilosa can enhance the fluidity of membrane by increase the unsaturated degree of fatty acids and maintain the stableness of the membrane structure by increase the content of the long chain saturated fatty acids. So as to improve their ability of endure the oxygen destruction caused by heavy metal stress.
3. Cd, Pb and Cd-Pb stresses can lead to the accumulation of soluble sugar and proline in the leaves of the B. pilosa. The degree of the accumulation differs from one to another due to different kinds of heavy metal stress and the plant's senstineness to the concentration. With the increase of Cd concentration stress, soluble sugar in leave of B. pilosa first increases and then decreases. The amount increase continuingly with the increase of Pb. Soluble proteins in B. pilosa increase at the beginning and then decrease with the increase of Cd or Pb stress. The Cd-Pb compound stress can promote the contents decreased of soluble protein and soluble protein in B. pilosa. With the increase of Cd. Pb or Cd-Pb stress, the activity of main antioxidant enzyme SOD. CAT and POD do not show any uniformity. With the increase of Cd. Pb or Cd-Pb compound stress, the activity of SOD in leaves of B. pilosa gradually increases. With the increase of Cd stress, the activity of CAT in B. pilosa increases first and then decreases. With the increase of Pb stress, the activity of CAT in B. pilosa continuingly increases. Under the condition of Cd-Pb compound stress, high concentration of stress promotes the increase of the activity of CAT in B. pilosa. With the increase of Cd stress, the activity of POD in B. pilosa increases at first and then decreases. And the activity of POD increases gradually in leaves of the paint with increase of Pb stress. Cd-Pb compound stress can slightly promote the activity of POD in leaves of plant and it relies on the concentration of Pb. Under the stress of Cd, Pb or Cd-Pb, B. pilosa can maintain the osmosis balance mainly by adjusting the content of proline, soluble sugar and soluble protein in the leaves and adjust the activity of SOD. CAT and POD to improve their ability in order to endure the antioxidant injury by heavy metal. As a result, they can enhance their ability to endure Cd and Pb stress.
4. Cd, Pb stresses promote the accumulation of oxalate in B. pilosa. They also chelate the absorbed heavy metal by adjusting the composition of malic acid, citric acid, acetic acid and amber acid in the roots and leaves of plants to reduce the toxicity of the heavy metal. In plants' bodies. Cd mainly exists in B. pilosa with the form of Fr3, which combined with protein and pectin. In this situation, low stress is beneficial to the transmission of Cd from roots to other positions and high stress restian the transport of Cd to leaves; Pb mainly exists in form of Fr5, a kind of oxalate, and high stress promotes the accumulation of Pb in the leaves. In compound stress conditions. Pb promotes Fr3turning to Fr4or Fr5to endure the contamination. Compared with Fr3, Fr4or Fr5is inactive. But Cd promotes Fr5to turn to active forms of Fr3, Fr4, Fr2and Fr1resulting to more toxicity effects. Cd and Pb mainly distribute in the subcellulars in form of cytoderm(F1)and dissoluvable parts(F4), but the content of chloroplast or plastid(F2)and other organelles is sparse. Pb can promote the increase of the content and proportion of low density Cd stressed plants' cytoderm and dissoluvable parts in case of Cd-Pb compound stress Cd can promote the increase of the proportion and content of Pb in chloroplasts or plastids, but reduce the content and proportion of cytoderms and dissoluvable parts. The plants enhance their ability of chelating heavy metals and reduce the toxicity of absorbed Cd and Pb by strategies of increase oxalate, malic acid and citric acid. They also increase the inert chemical forms of heavy metal in the subcellular distribution in order to enhance their ability to endure Cd and Pb stress.
5. In the case of Cd, Pb and Cd-Pb stresses, B. pilosa can accumulate large amount of axalate in rhizosphere. With the increase of Cd stress, the content of oxalate increases first and then decreases, while its content continuingly increases with increase of the density of Pb stress. Under the condition of Cd-Pb compound stress. The content of oxalate raises with the increase of Pb in low density Cd-Pb compound stress but increases first and then decreases in high density Cd-Pb compound stress. Cd exists mainly in the form of residual and its content increases with the increase of Cd stress. Pb mainly exists in the form of carbonate combination and Fe-Mn oxygenized combination take second place. Under the condition of Cd-Pb compound stress, increase of Pb can promote the Cd increase in changeable and carbonate combination form. The promotion is most obvious when adding Pb is1000mg/kg. Cd can promote residual Pb increase and M3of Pb decrease. Pb promote Cd transforming from less active form(M5)to active form(M3)but Cd promote Pb transforming from active form(M2)to less active form(M5). Cd or Pb has no obvious influence on the rhizosphere nutrition elements but K in the plant. But Cd-Pb compound stress decrease the accumulation of N, P and K. B. pilosa can enhance secreting oxalic acid to combinate heavy metal in order to affect the bioactivity of heavy metal in roots. As a result, they adjust the activity ratio and mobility ratio of the heavy metal and control the amount and time of the absorbed heavy metal, so that plants can enhance their ability to endure heavy metal.
6. Within a life period of B. pilosa, content of heavy metal in upper ground parts and roots gradually increases with the growing of plants, which consistent with biomass. After90days' stressing, when growth of the plants reaches their climax, the content of heavy metal and biomass in plants achieve the largest scale. The amount of heavy metal maintains in their maximum or slightly declines with life period changes from nutritive growth to reproductive growth. Under the condition of single heavy metal stress, the content of heavy metal in plants increases with the increase of the stress. Generally, in Cd-Pb compound situation, low and moderate density of Pb can promote the absorbing and transporting of low density Cd, but high density Pb seriously restrains the absorbing of Cd. In high density Pb-Cd compound stress, Pb restrains the plants to transmit Cd to upper ground parts but Cd shows no obvious influence on the absorbing and transporting of Pb. Generally, in a test group, when the content of Cd is large, the content of Pb is relatively small and if the content of Cd is small, the content of Pb is large, but the total amount of heavy metal remains invariablenes. Within life period of the plants, they show ability to accumulate and transport heavy metal, which increase at the beginning and then decreasing with the growth of the plants. In generally, the life span of B. pilosa is180days. So the best time to collect plants is90-120days for B. pilosa to phytoextract the heavy metal in polluted soil in excellent performance.
7. Low density of Cd or Pb promotes the accumulation of biomass in hairy roots of B. pilosa but high density restrains. In case of Cd-Pb compound stress, low and moderate density of Pb stresses promote accumulation of biomass; while other density Pb compound to high Cd stress restrain the accumulation of biomass. Under the condition of Cd or Pb stress, the hairy roots' accumulation of Cd and Pb is the additive result of the density and time of stressing. Hairy roots have different maximum absorbing amount in different density of stresses. In short period stresses, high density stress avails the hairy roots to accumulate heavy metals. Under the condition of low and high density stress, the absorbing of heavy metal gradually approaches peak and then maintains to the end of the stress. In case of Cd-Pb compound stress, low and moderate density of Pb promotes the hairy roots to accumulate Cd, while high density of Pb restrains the accumulation; low density of Pb stress has weak promotion merely to the accumulation of low density of Pb but it restrains the accumulation of Pb in the case of other varieties of density. Under the condition of Cd or Pb stress, the maximum contents of Cd or Pb in the hairy roots are2532mg/kg and2011mg/kg respectively. Under the condition of Cd-Pb compound stress, the maximum contents of Cd and Pb are2223mg/kg and1960mg/kg respectively. Pb stress has the effect of promoting the absorbing of Cd while Cd has the effect of restraining the absorbing of Pb. Compartmentalization and fixed holding of detoxification mechanisms of heavy metal by cell wall vacuole are the strategies of the hairy roots' endurance to Cd and Pb stress.
引文
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