随着城市化和工业化的快速发展，砷(As)和铅(Pb)的复合污染严重破坏了水体生态系统的正常功能，并通过食物链传递对人类健康造成巨大威胁。微藻是水环境的初级生产者，对元素生物地球化学循环有重要影响。然而，目前关于As和Pb对微藻的单一和联合毒性效应、以及微藻在两者复合胁迫下的响应机制仍不清楚。本文采用模式生物--莱茵衣藻，探究了 As、Pb单一和复合处理对藻细胞产生的毒性效应，分析了藻细胞在不同处理下的响应机制。主要研究结果如下：

      1. 构建了As和Pb混合体系毒性效应预测模型。选用Logit函数进行拟合浓度-效应之间的关系，得到单一As和Pb体系的EC50值分别为374.97 μg/L和19988.75 μg/L。通过浓度加和模型(CA)和独立作用模型(IA)评估As/Pb混合物毒性作用，同时进行实验验证。结果显示，随着As/Pb混合物浓度比逐渐增大，混合物的毒性效应由加和作用转变为协同作用。

       2. 探究了As对莱茵衣藻的毒性效应和细胞对As的解毒机制。研究结果表明，在低浓度As (0-20 μg/L)下，莱茵衣藻胞内淀粉含量增加，并产生更多的胞外多糖。在50 μg/L As处理下，藻细胞中的响应机制从淀粉积累逐渐转移到多糖分解。As暴露能够显著降低莱茵衣藻叶绿素荧光参数，表明As降低了类囊体膜上的电子传递链中PSⅡ系统的光合效率。在较高As浓度(500 μg/L)下，胞内As(Ⅴ)和As(Ⅲ)含量显著增加，分别达到213.94 μg/1012 cells和39.14 μg/1012 cells，从而增加了As在胞内的积累水平，造成细胞内稳态紊乱，加重了细胞受到的氧化损伤，最终在透射电镜(TEM)下观察到明显的细胞膜破裂和细胞裂解死亡。脂肪酸代谢途径中差异表达的蛋白质显著增多，表明细胞改变了碳代谢流向，将脂类转化为多糖。这一研究结果表明，多糖为细胞内的各种代谢过程提供能量，在莱茵衣藻对As的抗性中起关键作用。

      3. 探究了Pb对莱茵衣藻的毒性效应和细胞对Pb的解毒机制。结果表明，在本研究中Pb对细胞的毒性效应弱于As，Pb浓度达到5000 μg/L时，细胞光合系统中光能的转化和ATP的合成未受阻碍。SEM和TEM没有观察到明显的细胞结构破损和裂解死亡。但与对照组相比，Pb刺激引起细胞产生明显的能量代谢反应，多糖分泌量和呼吸速率显著增加，转录组分析结果显示，与磷脂合成和分泌相关的基因(PECT1、CAC3、captC、Pigo、yuiD、At1g和SFH2等)表达水平发生显著变化。细胞碳水化合物和酶的合成增加了Pb的吸附，并减少了其对细胞亚结构造成的损伤。

       4. 探究了不同浓度的As和Pb混合体系对莱茵衣藻的毒性效应。高浓度混合体系下的重金属毒性比单一体系更加明显，在TEM下观察到As(500 μg/L)+Pb(5000 μg/L)浓度下大量细胞裂解死亡，该浓度下的细胞数目仅为对照组的40%。As(100 μg/L)+Pb(1000 μg/L)处理显著抑制了藻细胞对P的吸收，在As(500 μg/L)+Pb(5000 μg/L)浓度时藻细胞对P的利用率下降了16%，同时总叶绿素含量下降了近70%。高浓度共存体系致使藻细胞合成大量碳水化合物以应对重金属胁迫从而维持正常的生长，当胞内As(Ⅴ)/As(Ⅲ)比值降到0.7，胞内大量积累毒性更强的As(Ⅲ)，细胞加速死亡。在这些因素共同作用下，衣藻细胞的能量代谢变化更加活跃，通过加速能量代谢消耗的方式来抵御强毒性混合体系。

       5.揭示了As和Pb对莱茵衣藻毒性的相互作用关系及联合毒性机理。通过皮尔逊相关性分析，验证了As与Pb联合毒性效应关系为低浓度加和作用和高浓度协同作用。产生上述作用的原因有：（1）对于膜表面的生物大分子物质（如磷脂等)，Pb比As具有更强的亲和性，这导致其优先与Pb结合，导致As不能及时排出而在胞内大量积累，加速细胞的死亡；（2）As和Pb对谷胱甘肽(GSH，合成植物螯合素的重要前体)具有竞争作用，由于Pb浓度远大于As，因此Pb在GSH竞争中取得优势，减少了As与GSH的结合，造成细胞更严重的氧化损伤；（3）与单一As体系相比，As(500 μg/L)+Pb(5000 μg/L)共存下的总叶绿素含量、饱和光电子传递速率Fv/Fm和实际光电子传递效率Y(Ⅱ)显著下降明显下降，表明复合体系阻碍了类囊体膜结构的光电子传递，破坏了光合系统；（4）蛋白质组学结果显示，在As(50 μg/L)+Pb(500 μg/L)处理中发现了71个不同表达的蛋白质(DEPs)，在As(500 μg/L)+Pb(5000 μg/L)的处理中发现了167个DEPs。这些蛋白主要参与了能量代谢、光合作用碳固定、活性氧清除和防御以及氨基酸合成等调控。

       综上，本文对As、Pb单一及复合处理对莱茵衣藻的毒性效应进行了系统研究，发现了As 对衣藻具有更强的毒性，并证实了Pb影响了微藻对As的积累和外排，并在一定程度上增强了As的毒性。同时，本文还分析了As、Pb单一和混合体系下微藻的解毒机制。研究发现，转录和蛋白层面的微观调控在耐受两种重金属的毒性和胞内膜结构、胞内碳水化合物代谢、叶绿体结构的损伤修复等方面发挥重要作用。

        With rapid urbanization and industrialization, combined pollution of arsenic (As) and lead (Pb) has seriously damaged the function of the water ecosystem, posing a significant threat to human health through food chain transmission. Microalgae are primary producers of the water environment, influencing the biogeochemical cycle of elements. However, up to now, the single and combined toxic effects of As and Pb on microalgae and mechanisms of microalgae response to their combined stress are still unclear. In this paper, Chlamydomonas reinhardtii, a model microalga, was used to explore the toxicity of single and combined treatment of As and Pb on algal cells, and the mechanisms of algal resistance to As and Pb with different concentrations was analyzed. The main results were as follows:

        1. The toxicity prediction model for the combined system of As and Pb was constructed. The dose-response relationship was fitted by the Logit function, where the EC50 values of sole As and Pb systems were 374.97 and 19988.75 μg/L, respectively. The interaction of As/Pb was evaluated by the concentration addition model (CA) and independent interaction model (IA). Meanwhile, the experiments were carried out to validate the modeling results. The results showed that with the increase of the As:Pb concentration ratio, the toxicity changed gradually from additive effect to synergistic effect.

2. The cellular toxicity of As and the detoxification mechanism of C. reinhardtii were explored. The results showed that at the low concentration of As (0-20 μg/L), cells accumulated starch and produced more extracellular polysaccharides. With 50 μg/L As, the response of algae cells gradually shifted from starch accumulation to polysaccharide decomposition. The As exposure significantly decreased the chlorophyll fluorescence parameters of algae cells, indicating that As reduced the efficiency of the photosynthetic electron transport chain on the thylakoid membrane. At the higher As concentration (500 μg/L), the intracellular As(Ⅴ) and As(Ⅲ) contents were significantly increased to 213.94 μg/1012 cells and 39.14 μg/1012 cells, respectively. Thus, the accumulation of As in the cell was enhanced, the intracellular homeostasis disorder was induced, and the oxidative damage of the cell was aggravated. Finally, obvious membrane and cell lysis were observed under transmission electron microscope (TEM). Moreover, the number of differentially expressed proteins in the fatty acid metabolism pathway increased significantly, indicating that the cells changed the carbon flow and converted lipids into polysaccharides. The results showed that polysaccharides could provide energy for various metabolic processes in cells, playing a vital role in the resistance of C. reinhardtii to As.

  3. The toxic mechanism of Pb to C. reinhardtii and the detoxification mechanism of microalgae to Pb were explored. The results showed that the toxicity of Pb on cells was weaker than that of As. When the Pb concentration reached 5000 μg/L, the conversion of light energy and the synthesis of ATP in the photosynthetic system of cells were not hindered. No apparent cell structure damage and cleavage death were observed according to the results of SEM and TEM. However, Pb stimulated obvious energy metabolism reactions, as polysaccharides secretion and respiration rate were significantly increased compared with the control group. Transcriptome analysis showed the expression of many genes related to phospholipid synthesis and secretion (PECT1, CAC3, captC, Pigo, yuiD, At1g and SFH2, etc.) were significantly changed. Furthermore, the enhanced synthesis of carbohydrates and enzymes increased the adsorption of Pb and reduced the damage to the cell substructure.

        4. The toxic effects of the combined system of As and Pb in C. reinhardtii were studied. It was observed that the toxicity of the high-concentration combined system was greater than that of the single system. Under TEM, a large number of cells died at the concentration of As (500 μg/L)) + Pb (5000 μg/L)), and the number of cells at this concentration was only 40% of that of the control group. The addition of As (100 μg/L)) + Pb (1000 μg/L)) significantly inhibited the absorption of P by algae cells. At the concentration of As (500 μg/L)) + Pb (5000 μg/L)), the utilization rate of P by algae cells decreased by 16%, and the total chlorophyll content decreased by nearly 70%. The high-concentration coexistence system caused algae cells to synthesize massive carbohydrates to cope with metal stress and maintain cell growth. When the intracellular As(V)/As(III) ratio decreased to 0.7, cell death was enhanced due to the intracellular accumulation of As(Ⅲ). Under the integrated effects of these factors, the change in the energy metabolism of algae cells was more active. It shows that C. reinhardtii could resist the highly toxic combined system by accelerating energy consumption.

        5. The interaction and mechanism of toxicity of As and Pb to C. reinhardtii were revealed. Pearson correlation analysis was used to verify the joint toxic effect relationship between As and Pb, which showed the additive effect at the low concentration and synergistic effect at the high concentration. The causes were systematically analyzed: (1) For the biological macromolecules on the membrane surface (such as phospholipids), Pb had a higher affinity than As, which led to its preferential binding to Pb and consumption. The binding to the membrane led to the As transporter channel activity change. Hence, As(Ⅲ) could not be excreted in time and accumulated in the cell, resulting in more cell death; (2) As and Pb could compete for glutathione (GSH, an essential precursor of artificial plant chelating elements). Because the concentration of Pb was much higher than that of As, Pb had the advantage in the competition for GSH, which made more ROS released by As and induced more oxidative damage; (3) Compared with the single As a system, the total chlorophyll content, saturated photoelectron transport rate Fv/Fm, and actual photoelectron transport efficiency Y(Ⅱ) were significantly decreased under the coexistence of As (500 μg/L) + Pb (5000 μg/L), indicating that the complex system hindered the photoelectron transport of thylakoid membrane structure and destroyed the photosynthetic system; (4) Proteomic results showed that 71 differently expressed proteins (DEPs) were found in the treatment of As (50 μg/L) + Pb (500 μg/L), and 167 DEPs were found in the treatment of As (500 μg/L) + Pb (5000 μg/L). These proteins were mainly involved in regulating energy metabolism, carbon fixation in photosynthesis, scavenging and defense of reactive oxygen species and amino acid synthesis.

       In summary, the toxic effects and mechanisms of single and combined  As and Pb treatments to C. reinhardtii were revealed. It was found that As was more toxic to algae than Pb, and it was confirmed that Pb affected the accumulation and efflux of As by microalgae, and Pb enhanced the toxicity of As to some extent. Meanwhile, the detoxification mechanisms of microalgae under single and combined treatments of As and Pb were systematically analyzed. It was found that regulations at transcriptomic and proteomic-levels played an important role in the toleranceof As and Pb toxicity, through maintaining intracellular membrane structure, regulating intracellular carbohydrate metabolism, and repairing chloroplast structure damage.

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