喹赛多毒理性质的代谢组学研究
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摘要
作为一种抗菌促生长剂,喹赛多有很大的潜力被添加到饲料中用于促进畜禽类动物的生长,但要成为饲料添加剂必须通过全面严格的食品安全评估。迄今为止,关于喹赛多毒作用性质的传统毒理学研究相对较多,但是喹赛多对机体内源性代谢的影响还没有相关的报道。因此,本论文用基于核磁共振波谱的代谢组学方法研究了喹赛多对Kunming小鼠内源性代谢产物的影响；为了进一步研究喹赛多对机体内源性代谢物影响的种属差异,我们又设计了喹赛多对Wistar大鼠的急性毒理实验；在这两个实验的基础上,为了再进一步探讨长期喹赛多暴露对机体内源性代谢的影响,我们还设计了喹赛多对Wistar大鼠的亚慢性毒理实验。在亚慢性毒理实验中,除了传统的代谢组学方法,我们还采用了大鼠肝脏表达谱全基因组测序以及荧光定量PCR检测的方法,研究了长期的喹赛多暴露对大鼠肝脏的转录组的影响。
喹赛多对Kunming小鼠的急性毒理实验一共设计了三个剂量组(100,650,4000mg/kg body weight)和一个对照组。实验采取的是一次性灌胃给药的方式,研究了给药后小鼠尿样、血样、肝脏以及肾脏中代谢物水平的变化。在整个实验期间,代谢物的变化具有剂量依赖性,只有低剂量和中剂量组的Kunming小鼠的代谢水平恢复到了正常。喹赛多扰乱了Kunming小鼠肠道菌群的正常生命活动,这表现在一系列与肠道菌群代谢相关的物质如马尿酸、氧化三甲胺、三甲胺以及二甲胺等代谢物水平的显著下降。除此之外,高剂量的给药也导致了小鼠肝脏中氨基酸的聚集以及核苷酸代谢水平的下降。而在肾脏中,核苷酸以及一系列的渗透因子比如肌醇、胆碱以及甘油脂酰胆碱的含量明显增加,而氨基酸的含量则出现了显著性的下降。这些结果表明,喹赛多抑制了肝脏中的氨基酸代谢,扰乱了肠道菌群的生命活动,影响了肝脏和肾脏的渗透压平衡以及核苷酸代谢。
喹赛多在Wistar大鼠体内的急性毒理实验同样也设计了三个剂量组(50,325,2000mg/kg body weight)和一个对照组,实验的方式以及检测分析的样品和小鼠急性毒理实验是一样的。总体上来说,大鼠对喹赛多暴露的反应比小鼠要敏感。这表现在给药后大鼠血液的内平衡被打破,出现了一系列显著性变化的物质,如中剂量组大鼠血样中代谢水平显著性增加的异亮氨酸、缬氨酸、甲硫氨酸、天冬酰胺和肌酸以及高剂量组中代谢水平显著性增加的丙氨酸、琥珀酸、甲硫氨酸、葡萄糖、乳酸和代谢水平显著性下降的脂类。另外,与小鼠反应不一样的还有大鼠肝脏中的氨基酸代谢并没有受到明显的抑制、大鼠体内的能量代谢受抑制的情况要比小鼠严重并且没有发现大鼠肾脏代谢水平的显著性变化；而和小鼠一样的是,其体内核酸代谢也受到了明显的抑制以及肠道菌群的正常生命活动也受到了明显的扰动。
喹赛多在Wistar大鼠体内的亚慢性毒理实验也设计了三个剂量组(50,150,2500mg/kg feed)和一个对照组,经过三周的适应期后,给药12周,恢复两周,在给药后第6周、12周以及恢复两周后的14周处死大鼠。代谢组学的分析结果表明,对于尿样和肾脏来说,低剂量和中剂量组大鼠对药物的反应时而显著时而不显著,呈现逐渐适应的趋势,高剂量组的大鼠代谢物受到的扰动比较大,但经过两周的恢复期以后,剂量组均恢复到了正常的状态。对于血样和肝脏,低剂量组和中剂量组的血样仅出现一些血生化指标的变化,并未检测到显著性变化的代谢物。高剂量组中无论是血生化指标还是代谢物的代谢水平都受到了很大的扰动。而在肝脏中,药物的影响似乎有一种累积效应。随着时间的变化,药物对代谢物的扰动甚至出现在低剂量组中。而病理切片检测也发现了中剂量和高剂量大鼠肝脏受损的现象。对给药12周后高剂量组大鼠肝脏的基因表达结果进行转录组学分析,结果发现很多与药物代谢途径相关的基因如Cypla、Hsdllbl和Cyp2a2,与脂类代谢和转运途径相关的基因如Scd、 Acotl/2/4、Acsm2和Pexlla、与糖酵解和TCA循环途径相关的基因如Slc34a2和Pfkfb1与氨基酸代谢途径相关的基因如Daao、与转录调控相关的基因如Nfe2、 Pir和Scyllbp1、与炎性反应途径相关的基因如Cxcll3、 TmemSSa、Rtl-N和Icaml以及与神经系统途径相关的基因如Faml34b、Egrl、Scyllbpl、Colq、Gabbprl、Trim2、Epha4、Rtn4rll和Hspbl的表达都发生了显著性的变化,而代谢组学分析的结果表明发生显著性变化的代谢物所涉及到的代谢途径大多数与此也是一致的。
本论文从转录组和代谢组水平探讨了喹赛多的作用机制。机体吸收喹赛多以后,主要的代谢器官是肝脏。在药物代谢的过程中,会释放一定量的活性氧自由基(ROS),活性氧自由基的长期刺激极有可能会引发机体的脂质过氧化,从而导致机体内一系列的代谢途径如脂类代谢、氨基酸代谢、TCA循环以及核苷酸代谢等的异常。喹赛多对机体的损伤还表现在肝脏的病理学检测上,我们发现喹赛多处理以后,肝脏中出现了细胞水样变性、点状坏死以及脂肪变性等异常。除此之外,作为一种广谱抗菌剂,喹赛多也显著地抑制了机体肠道菌群的生命活动。总体上来说,喹赛多对实验动物的影响表现出了明显的剂量依赖效应,不同种属的动物对喹赛多的反应有相似也有不同,Wistar大鼠对喹赛多的反应比Kunming小鼠要敏感。50mg/kg feed是喹赛多用作饲料添加剂的推荐剂量,在亚慢性毒理实验中,我们发现此剂量无论是对大鼠的尿样还是肾脏代谢组的影响都是非常轻微并且可以恢复的,没有发现血样的异常。但是长期喂饲此剂量的喹赛多,对肝脏还是有可能造成一定程度的影响。总之,在喹赛多的使用过程中,严格地控制其添加剂量并给予一定的恢复期,从代谢组的水平上来说是可行的。
综上所述,本论文利用代谢组学和转录组学的研究方法系统地研究了喹赛多的毒理学性质。为喹赛多的食品安全评估提供了重要的参考。
As an antimicrobial growth promoter, cyadox has great potential to be used as additive in the feedstuff for livestock and poultry. However, a comprehensive food safety assessment should be performed before becoming feed additives. So far, there is no a report on how cyadox impacts on endogenous metabolites of animals, though many traditional toxicological studies of cyadox have been conducted. Therefore, in this paper, NMR-based metabonomic techniques were used to study the impact of cyadox on the endogenous metabolisms of Kunming mice. To further study metabolic responses of different species, endogenous metabolome changes induced by cyadox in Wistar rats were also investigated. On the basis of these two experiments, we also have designed a sub-chronic toxicological experiment on Wistar rats, in order to explore how endogenous metabolome changes under long-term cyadox exposure. In addition, transcriptomics profile was also used to investigate the variance of liver on transcriptome level.
In the study of metabolic influence of acute cyadox exposure on Kunming mice, three groups of mice were respectively given a single dose of cyadox at three different concentrations (100,650and4000mg/kg body weight) via gavage. We present here the metabolic alterations of urine, plasma, liver and renal medulla extracts induced by cyadox exposure. The metabolic alterations induced by cyadox exposure are dose dependent and metabolic recovery is only achieved for low and moderate levels of cyadox exposure during experimental period. Cyadox exposure resulted in a disturbance of gut microbiota, which is manifested in the depleted levels of urinary hippurate, trimethylamine-N-oxide, dimethylamine and trimethylamine. In addition, cyadox exposure on high levels caused accumulations of amino acids and depletions of nucleotides in the liver. Furthermore, marked elevations of nucleotides and a range of organic osmolytes, such as myo-inositol, choline and glycerophosphocholine and decreased levels of amino acids are observed in the renal medulla of cyadox exposed mice. These results suggest that cyadox exposure causes inhibition of amino acids metabolism in the liver and disturbance of gut microbiota community, influences osmolytic homeostasis and nucleic acids synthesis in both the liver and the kidney.
In the study of metabolic influence of acute cyadox exposure on Wistar rats, three groups of Wistar rats were respectively given a single dose of cyadox at three different concentrations (50,325and2000mg/kg body weight) via gavage. Compared to the mice, rats were more sensitive to cyadox mainly represented in the changes on the plasma profiles, an indication of inbalanced organism homeostasis. A series of significant changes including increased leucine, valine, methionine, asparagine, and creatine in moderate dose group and the elevated level of alanine, succinic acid, methionine, glucose and lactic acid and decreased lipid in the high-dose group were observed. In addition, no significant inhibition of amino acids metabolism in rat liver and no disturbance in the rat kidney were observed. However, disturbances of nucleic acid metabolism in the liver and gut microbiota were also observed in the rats.
In the study of metabolic and transcriptome influences of sub-chronic cyadox exposure on Wistar rats, three groups of Wistar rats were respectively fed with foods containing three different concentrations of cyadox (50,150,2500mg/kg feed). Two weeks recovery were conducted before experiment ended. Results of metabonomic analysis showed that metabolites in urine of rats in low-and moderate-dose changed freely and were not in some rule. Kidney showed variance on the metabolism of amino acids and nucleic acids but no change was observed after two weeks recovery. Variations in the profiles of plasma were only observed in the high dose group during the entire experiment time. Cyadox seemed to have a cumulative effect on the profiles liver, The metabolic displacement increased as the dosage becomes high level. No obvious recovery was observed for rats exposed to moderate and high levels of cyadox. Moreover, disturbance even appeared in the rats of the low dose group, after two weeks recovery time, though no abnormal changes were observed during the dosed time. We further investigated gene expressions of the liver of high-dose group after12weeks cyadox administration. Genes involved drug metabolism such as Cyplal, Hsdllb1and Cyp2a2, involved lipid metabolism such as Scd, Acotl/2/4, Acsm2and Pexlla, involved glycolytic pathway and TCA cycle pathway such as Slc34a2and Pfkfbl, involved amino acid metabolic pathway such as Daao, involved transcriptional regulatory pathways such as Nfe2, Pir and Scyllbpl, involved inflammatory reaction pathway such Cxcll3, Tmem55a, Rtl-N and Icaml and involved nervous system pathway such as Fam134b, Scyllbpl, Colq, Gabbprl, Trim2, Epha4, Rtn4rll and Hspbl all changed significantly, which were mostly consistent with the metabolic changes.
Mechanisms of cyadox action on organism were investigated in this thesis from transcriptome and metabolome aspects. Liver is the major metabolic organ for cyadox. A certain amount of reactive oxygen species (ROS) will be released during the process of cyadox metabolism and long-term stimulation of reactive oxygen species might cause lipid peroxidation, resulting disturbances in a series of metabolic pathways, such as in the lipid metabolism, amino acid metabolism, TCA cycle, nucleotide metabolism and so on. In addition, hydropic degeneration, punctate necrosis and fatty degeneration of hepatic cells were also observed in liver pathology. Moreover, as a broad-spectrum antibacterial agent, cyadox also suppressed the activity of the gut microbiota significantly. Overall, dose-dependent effect and different actions on different species were observed. Wistar rats were more sensitive to cyadox than Kunming mice. The dose of cyadox with50mg/kg feed is the recommended dose used as feed additives. In the sub-chronic toxicological experiments, rats'responses on this dosage on either urine or kidneys are very slight and can be restored and no abnormalities were observed on the plasma. However, long-term feeding of cyadox on this dose may cause a certain degree of influence on the liver. In short, it's practicable on the metabolic aspect for cyadox's used in livestock farming when the dosage was strictly controlled and some recovery time was given.
In summary, this thesis provides a comprehensive view of the toxicological effects of cyadox with the help of metabolomics and transcriptomics profile, which is important in animal and human food safety.
喹赛多对Kunming小鼠的急性毒理实验一共设计了三个剂量组(100,650,4000mg/kg body weight)和一个对照组。实验采取的是一次性灌胃给药的方式,研究了给药后小鼠尿样、血样、肝脏以及肾脏中代谢物水平的变化。在整个实验期间,代谢物的变化具有剂量依赖性,只有低剂量和中剂量组的Kunming小鼠的代谢水平恢复到了正常。喹赛多扰乱了Kunming小鼠肠道菌群的正常生命活动,这表现在一系列与肠道菌群代谢相关的物质如马尿酸、氧化三甲胺、三甲胺以及二甲胺等代谢物水平的显著下降。除此之外,高剂量的给药也导致了小鼠肝脏中氨基酸的聚集以及核苷酸代谢水平的下降。而在肾脏中,核苷酸以及一系列的渗透因子比如肌醇、胆碱以及甘油脂酰胆碱的含量明显增加,而氨基酸的含量则出现了显著性的下降。这些结果表明,喹赛多抑制了肝脏中的氨基酸代谢,扰乱了肠道菌群的生命活动,影响了肝脏和肾脏的渗透压平衡以及核苷酸代谢。
喹赛多在Wistar大鼠体内的急性毒理实验同样也设计了三个剂量组(50,325,2000mg/kg body weight)和一个对照组,实验的方式以及检测分析的样品和小鼠急性毒理实验是一样的。总体上来说,大鼠对喹赛多暴露的反应比小鼠要敏感。这表现在给药后大鼠血液的内平衡被打破,出现了一系列显著性变化的物质,如中剂量组大鼠血样中代谢水平显著性增加的异亮氨酸、缬氨酸、甲硫氨酸、天冬酰胺和肌酸以及高剂量组中代谢水平显著性增加的丙氨酸、琥珀酸、甲硫氨酸、葡萄糖、乳酸和代谢水平显著性下降的脂类。另外,与小鼠反应不一样的还有大鼠肝脏中的氨基酸代谢并没有受到明显的抑制、大鼠体内的能量代谢受抑制的情况要比小鼠严重并且没有发现大鼠肾脏代谢水平的显著性变化；而和小鼠一样的是,其体内核酸代谢也受到了明显的抑制以及肠道菌群的正常生命活动也受到了明显的扰动。
喹赛多在Wistar大鼠体内的亚慢性毒理实验也设计了三个剂量组(50,150,2500mg/kg feed)和一个对照组,经过三周的适应期后,给药12周,恢复两周,在给药后第6周、12周以及恢复两周后的14周处死大鼠。代谢组学的分析结果表明,对于尿样和肾脏来说,低剂量和中剂量组大鼠对药物的反应时而显著时而不显著,呈现逐渐适应的趋势,高剂量组的大鼠代谢物受到的扰动比较大,但经过两周的恢复期以后,剂量组均恢复到了正常的状态。对于血样和肝脏,低剂量组和中剂量组的血样仅出现一些血生化指标的变化,并未检测到显著性变化的代谢物。高剂量组中无论是血生化指标还是代谢物的代谢水平都受到了很大的扰动。而在肝脏中,药物的影响似乎有一种累积效应。随着时间的变化,药物对代谢物的扰动甚至出现在低剂量组中。而病理切片检测也发现了中剂量和高剂量大鼠肝脏受损的现象。对给药12周后高剂量组大鼠肝脏的基因表达结果进行转录组学分析,结果发现很多与药物代谢途径相关的基因如Cypla、Hsdllbl和Cyp2a2,与脂类代谢和转运途径相关的基因如Scd、 Acotl/2/4、Acsm2和Pexlla、与糖酵解和TCA循环途径相关的基因如Slc34a2和Pfkfb1与氨基酸代谢途径相关的基因如Daao、与转录调控相关的基因如Nfe2、 Pir和Scyllbp1、与炎性反应途径相关的基因如Cxcll3、 TmemSSa、Rtl-N和Icaml以及与神经系统途径相关的基因如Faml34b、Egrl、Scyllbpl、Colq、Gabbprl、Trim2、Epha4、Rtn4rll和Hspbl的表达都发生了显著性的变化,而代谢组学分析的结果表明发生显著性变化的代谢物所涉及到的代谢途径大多数与此也是一致的。
本论文从转录组和代谢组水平探讨了喹赛多的作用机制。机体吸收喹赛多以后,主要的代谢器官是肝脏。在药物代谢的过程中,会释放一定量的活性氧自由基(ROS),活性氧自由基的长期刺激极有可能会引发机体的脂质过氧化,从而导致机体内一系列的代谢途径如脂类代谢、氨基酸代谢、TCA循环以及核苷酸代谢等的异常。喹赛多对机体的损伤还表现在肝脏的病理学检测上,我们发现喹赛多处理以后,肝脏中出现了细胞水样变性、点状坏死以及脂肪变性等异常。除此之外,作为一种广谱抗菌剂,喹赛多也显著地抑制了机体肠道菌群的生命活动。总体上来说,喹赛多对实验动物的影响表现出了明显的剂量依赖效应,不同种属的动物对喹赛多的反应有相似也有不同,Wistar大鼠对喹赛多的反应比Kunming小鼠要敏感。50mg/kg feed是喹赛多用作饲料添加剂的推荐剂量,在亚慢性毒理实验中,我们发现此剂量无论是对大鼠的尿样还是肾脏代谢组的影响都是非常轻微并且可以恢复的,没有发现血样的异常。但是长期喂饲此剂量的喹赛多,对肝脏还是有可能造成一定程度的影响。总之,在喹赛多的使用过程中,严格地控制其添加剂量并给予一定的恢复期,从代谢组的水平上来说是可行的。
综上所述,本论文利用代谢组学和转录组学的研究方法系统地研究了喹赛多的毒理学性质。为喹赛多的食品安全评估提供了重要的参考。
As an antimicrobial growth promoter, cyadox has great potential to be used as additive in the feedstuff for livestock and poultry. However, a comprehensive food safety assessment should be performed before becoming feed additives. So far, there is no a report on how cyadox impacts on endogenous metabolites of animals, though many traditional toxicological studies of cyadox have been conducted. Therefore, in this paper, NMR-based metabonomic techniques were used to study the impact of cyadox on the endogenous metabolisms of Kunming mice. To further study metabolic responses of different species, endogenous metabolome changes induced by cyadox in Wistar rats were also investigated. On the basis of these two experiments, we also have designed a sub-chronic toxicological experiment on Wistar rats, in order to explore how endogenous metabolome changes under long-term cyadox exposure. In addition, transcriptomics profile was also used to investigate the variance of liver on transcriptome level.
In the study of metabolic influence of acute cyadox exposure on Kunming mice, three groups of mice were respectively given a single dose of cyadox at three different concentrations (100,650and4000mg/kg body weight) via gavage. We present here the metabolic alterations of urine, plasma, liver and renal medulla extracts induced by cyadox exposure. The metabolic alterations induced by cyadox exposure are dose dependent and metabolic recovery is only achieved for low and moderate levels of cyadox exposure during experimental period. Cyadox exposure resulted in a disturbance of gut microbiota, which is manifested in the depleted levels of urinary hippurate, trimethylamine-N-oxide, dimethylamine and trimethylamine. In addition, cyadox exposure on high levels caused accumulations of amino acids and depletions of nucleotides in the liver. Furthermore, marked elevations of nucleotides and a range of organic osmolytes, such as myo-inositol, choline and glycerophosphocholine and decreased levels of amino acids are observed in the renal medulla of cyadox exposed mice. These results suggest that cyadox exposure causes inhibition of amino acids metabolism in the liver and disturbance of gut microbiota community, influences osmolytic homeostasis and nucleic acids synthesis in both the liver and the kidney.
In the study of metabolic influence of acute cyadox exposure on Wistar rats, three groups of Wistar rats were respectively given a single dose of cyadox at three different concentrations (50,325and2000mg/kg body weight) via gavage. Compared to the mice, rats were more sensitive to cyadox mainly represented in the changes on the plasma profiles, an indication of inbalanced organism homeostasis. A series of significant changes including increased leucine, valine, methionine, asparagine, and creatine in moderate dose group and the elevated level of alanine, succinic acid, methionine, glucose and lactic acid and decreased lipid in the high-dose group were observed. In addition, no significant inhibition of amino acids metabolism in rat liver and no disturbance in the rat kidney were observed. However, disturbances of nucleic acid metabolism in the liver and gut microbiota were also observed in the rats.
In the study of metabolic and transcriptome influences of sub-chronic cyadox exposure on Wistar rats, three groups of Wistar rats were respectively fed with foods containing three different concentrations of cyadox (50,150,2500mg/kg feed). Two weeks recovery were conducted before experiment ended. Results of metabonomic analysis showed that metabolites in urine of rats in low-and moderate-dose changed freely and were not in some rule. Kidney showed variance on the metabolism of amino acids and nucleic acids but no change was observed after two weeks recovery. Variations in the profiles of plasma were only observed in the high dose group during the entire experiment time. Cyadox seemed to have a cumulative effect on the profiles liver, The metabolic displacement increased as the dosage becomes high level. No obvious recovery was observed for rats exposed to moderate and high levels of cyadox. Moreover, disturbance even appeared in the rats of the low dose group, after two weeks recovery time, though no abnormal changes were observed during the dosed time. We further investigated gene expressions of the liver of high-dose group after12weeks cyadox administration. Genes involved drug metabolism such as Cyplal, Hsdllb1and Cyp2a2, involved lipid metabolism such as Scd, Acotl/2/4, Acsm2and Pexlla, involved glycolytic pathway and TCA cycle pathway such as Slc34a2and Pfkfbl, involved amino acid metabolic pathway such as Daao, involved transcriptional regulatory pathways such as Nfe2, Pir and Scyllbpl, involved inflammatory reaction pathway such Cxcll3, Tmem55a, Rtl-N and Icaml and involved nervous system pathway such as Fam134b, Scyllbpl, Colq, Gabbprl, Trim2, Epha4, Rtn4rll and Hspbl all changed significantly, which were mostly consistent with the metabolic changes.
Mechanisms of cyadox action on organism were investigated in this thesis from transcriptome and metabolome aspects. Liver is the major metabolic organ for cyadox. A certain amount of reactive oxygen species (ROS) will be released during the process of cyadox metabolism and long-term stimulation of reactive oxygen species might cause lipid peroxidation, resulting disturbances in a series of metabolic pathways, such as in the lipid metabolism, amino acid metabolism, TCA cycle, nucleotide metabolism and so on. In addition, hydropic degeneration, punctate necrosis and fatty degeneration of hepatic cells were also observed in liver pathology. Moreover, as a broad-spectrum antibacterial agent, cyadox also suppressed the activity of the gut microbiota significantly. Overall, dose-dependent effect and different actions on different species were observed. Wistar rats were more sensitive to cyadox than Kunming mice. The dose of cyadox with50mg/kg feed is the recommended dose used as feed additives. In the sub-chronic toxicological experiments, rats'responses on this dosage on either urine or kidneys are very slight and can be restored and no abnormalities were observed on the plasma. However, long-term feeding of cyadox on this dose may cause a certain degree of influence on the liver. In short, it's practicable on the metabolic aspect for cyadox's used in livestock farming when the dosage was strictly controlled and some recovery time was given.
In summary, this thesis provides a comprehensive view of the toxicological effects of cyadox with the help of metabolomics and transcriptomics profile, which is important in animal and human food safety.
引文
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