河南内乡与四川会理烤烟后期生长发育的分子特征研究
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摘要
本论文大田试验以河南内乡与四川会理大田生产的烤烟(Nicotiana tabacum L.)品种云烟85的上部烟叶为材料,从移栽后60 d开始取样,采集第18片叶同一部位分别:1,检测相关基因的表达量;2,检测化学成分和相关酶活性;3,取烟叶中间部位主脉两侧,用于组织切片的观察。以后每隔10 d取样一次。首先,本文研究了河南内乡与四川会理的烟草在移栽后60-100 d期间烤烟的化学成分及致香成分含量的差异,并分为碳代谢、氮代谢和次生代谢三种代谢相关的物质进行分析,通过分析与三种代谢相关的酶活性及相关基因表达的差异,从分子水平分析这些差异的原因。其次,本文研究了河南内乡与四川会理烟草烟叶中蔗糖和类胡萝卜素合成相关的关键基因(蔗糖磷酸合成酶、蔗糖合酶和番茄红素β-环化酶)的变化,以及这些基因与温度、光照和温度与光照交互作用之间的关系,为调控这些基因的表达奠定理论和技术基础。
再者,本文建立了烤烟DNA甲基化水平的HPLC检测方法,并以确山烟叶生长期间主要的四个气象因子温度、日照、降水、湿度和烟叶DNA甲基化水平之间建立了Logistic方程的数学模型,研究烟草生长过程中气象因子与DNA甲基化之间的交互作用,从分子水平上探讨气象因子对烟叶生理生化的影响,为深入研究特色烟叶的形成与气象因子的关系提供依据。
本论文通过相关的试验,得出的主要结论如下:
1,河南内乡与四川会理烤烟的生长期不太一致,河南内乡的烟叶前期生长快,后期生长慢;会理的烟叶表现前期慢,后期生长较快。河南内乡与四川会理烟叶可溶性总糖、类胡萝卜素及其降解产物含量的差异较大。河南内乡与四川会理烟叶在移栽后60-100 d期间,烟叶的可溶性总糖的含量变化均为单峰曲线,在移栽后70 d达到高点,随后可溶性总糖的含量逐渐下降,到移栽后100 d时,烟叶中可溶性总糖的含量达到最低。烤后会理烟叶中可溶性总糖比内乡烟叶的要高1.5倍。
2,在烟草移栽后60-80 d烟叶中蔗糖磷酸合成酶(SPS)的活性会理高于内乡烟叶,而到移栽90 d后,会理烟叶中SPS酶活性迅速降低,并且SPS酶活性低于内乡的。在烟草移栽后60-90 d烟叶中蔗糖合酶(SS)的活性会理低于内乡烟叶,而到移栽90 d后,内乡烟叶中SS酶活性降低的较快,并且会理SS酶活性高于内乡的。
3,对烤烟叶圆片进行的温度和光照正交试验的结果表明,温度和光照处理能够下调或上调SPS和SS基因的表达,只有在适当的温度和光照条件下,才有利于SPS基因的表达。在较高温度和光照强度条件下,有利于SS基因的表达。极差的结果表明,三个因子对SPS和SS基因表达的影响作用的大小顺序一样,即(温度×光照强度)>光照强度=温度。Duncan’s测验比较结果表明:温度×光照强度分别对SPS和SS基因的表达影响较大,且达到差异显著水平(P<0.05)。
4,SPS和SS基因的表达量和Northern杂交的结果表明,四川会理烟叶中SPS基因表达水平明显高于河南内乡烟叶。四川会理烟叶中SS基因的杂交带亮度稍高于河南内乡烟叶的条带亮度,表明四川会理烟叶中SS基因表达水平稍高于河南内乡烟叶。现蕾期四川会理烟叶SPS酶活性比河南内乡烟叶提高有利,而两地区烟叶SS酶活性的差异不大。四川会理烟叶的可溶性总糖的含量高于河南内乡的原因是:四川会理烟叶中SPS酶活性和SPSA基因的表达高于内乡烟叶,而四川会理烟叶中SS酶活性和SS基因的表达与河南内乡烟叶的差异不大,使得糖分在烟叶中积累。
5,两地区烟叶中烟碱含量随着烟叶的成熟有逐渐增加的趋势,而烟叶中总氮和蛋白质的含量却随着烟叶的成熟逐渐下降。游离氨基酸的变化为,内乡烟叶在移栽后70 d达到高峰,随后逐渐下降。在移栽后60-100 d期间,NR酶活性和基因表现为逐渐下降,且会理烟叶中NR酶活性一直高于内乡烟叶;河南内乡与四川会理烟叶中GS基因的表达在此期间均表现逐渐下降,GS酶活性也表现为会理烟叶大于内乡;而河南内乡与四川会理烟叶中GDH基因的表达表现为逐渐升高的趋势,且四川会理烟叶中GDH基因的表达比河南内乡烟叶中该基因表达要高。
6,在现蕾期四川会理烤烟上部叶中类胡萝卜素的含量显著高于河南内乡的,基因Lyc-β的半定量RT-PCR和Northern杂交的结果表明,四川会理烟叶中Lyc-β基因表达水平明显高于河南内乡烟叶。Lyc-β基因的表达受到温度和光照的调节,对Lyc-β基因的表达影响的顺序是,光照强度>温度>温度×光照强度,且三因素的三个水平之间没有显著差异(P<0.05)。较高光照的条件下,Lyc-β基因的表达是较低光照的1.60倍;温度为30℃时,Lyc-β基因的表达最高,温度在25℃时次之;温度在35℃时最小。在光照时间为2-6 h期间,Lyc-β基因随着光照时间的延长表达量增加,但光照处理超过8 h之后,Lyc-β基因随着光照时间的延长表达量下调。总之,光照和温度可能是通过调节lyc-β基因的表达造成不同生态区烟叶中类胡萝卜素差异的原因。
7,气象因子与DNA甲基化之间Logistic模型的单因子效应分析表明,平均最高温度的升高对DNA甲基化水平的升高影响最大;总日照长度的延长也会提高烟叶DNA甲基化水平;平均相对湿度对DNA甲基化的影响不大;总降雨量的增加能够降低总DNA甲基化水平,虽然其系数绝对值较小,但总降雨量的数值较大,同样对DNA去甲基化水平的影响幅度也大。总之,平均最高温度、总日照时数、平均相对湿度与DNA甲基化水平正相关;总降雨量与DNA甲基化水平负相关。两因素交互效应分析表明,平均最高温度和平均相对湿度、平均最高温度和总日照时数之间对烟叶的DNA甲基化具有协同效应;总降雨量和平均最高温度、总降雨量和总日照时数、总降雨量和平均相对湿度之间对烟叶的DNA甲基化有拮抗作用;而平均相对湿度与总日照时数之间对烟叶的DNA甲基化水平没有明显的协同效应或拮抗作用。
This paper was designed with the top leaves (the 18th leaf) of flue-cured tobacco (Nicotiana tabacum L.) between Henan and Sichuan for the materials. Sixty days after transplanting, we investigated the tobacco agronomic characters and collected the same position of the 18th leaves in order to detect chemical composition, related enzyme activity, related genes expression, and observations of tissue slice. We sampled after that every 10 days.
Firstly, this paper discussed the differences in the three physicochemical properties: carbon metabolism, nitrogen metabolism and secondary metabolism of tobacco between Henan and Sichuan during 60-100 d after transplanting. Moreover, analyzed the differences of enzyme activity, genes expression related to the three metabolic, and analyzed the causes of the differences from molecular level. Secondly, this paper studied the differences on key genes (SPS, SS, lyc-β) related to sucrose and carotenoids synthetic of tobacco between Henan and Sichuan. This paper also studied the relation between these genes, temperature, and light to build a theoretical and technical basis to regulate these genes expression.
In addition, this paper established Logistic model between tobacco DNA methylation levels and four meteorological factors (temperature, sunshine, rainfall, humidity) during the growth of Queshan tobacco, and researched the interact between meteorological factors during tobacco growth and DNA methylation, and explored meteorological factors’influence to tobacco physiology and biochemistry at the molecular level. In addition, this provides basis to further research the relation between the formation of characteristic leaf tobacco and meteorological factors. The main conclusions of this experiment were as follows:
1. Total thickness of tobacco leaves both in Henan and Sichuan continuously increased after transplanting. The Neixiang tobacco growed fast in the early and slow in late. However, Huili’s growed slow at the earlier time and fast later. During 60-100 d after transplanting, both in Henan and Sichuan, soluble sugar content of tobacco presented single apex curve, 70 d after transplanting was the time that the peak value show, then 100 d after transplanting total soluble sugar content decreased to the lowest. Soluble sugar content of Huili tobacco was 1.5 times higher than Neixiang’s after cured.
2. Sixty-eighty days after transplanting, SPS activity of tobacco in Huili is higher than that of tobacco in Neixiang, however, at 90 d after transplanting, SPS activity of tobacco in Huili decreased rapidly to lower than that in Neixiang. 60-90 d after transplanting, the SS activity of tobacco in Huili is lower than that of Neixiang, and the SS activity in Neixiang decreased rapidly to lower than that of Huili at 90 d after transplanting. Between 60-100 d after transplanting, the activity of sucrose invertase in Huili is higher than Neixiang. During the period 60-100 d after tobacco transplanting, SPS expression of tobacco in Sichuan showed a gradual decline but the level is higher than that in Henan. During the same period, SS expression of tobacco in Sichuan is biased single peak changes, which at the 90 d reached a peak. The genes expression of tobacco had no significant difference in Henan and Sichuan.
3. The leaf discs for temperature and light orthogonal experiment results in laboratory showed that the temperature and light treatment could decrease or raise the SPS and SS genes expression: only a proper temperature and light condition was conducive to the SPS expression. High temperature and light intensity conditions will be helpful for SS gene expression. Influence of the three factors on SPS and SS genes expression is in the order that (temperature×light intensity)> light intensity = temperature. Duncan's test comparison results showed that the condition that the temperature interact light intensity achieves great influence on SPS and SS genes expression, which achieves significant levels (P < 0.05).
4. SPS gene expression and Northern blot results showed that SPS gene expression levels of the tobacco in Sichuan were significantly higher than that of Henan. The result that SS gene hybridization bands brightness of Sichuan Tobacco was slightly higher than that of Henan revealed that SS gene expression in Sichuan tobacco leaves was slightly higher than that in Henan. In budding stage, SPS activity in Huili, Sichuan tobacco leaves is higher than that in Henan, however, SS activity in both regions had no much difference. In short, light and temperature may regulate the expression of lyc-βgene in tobacco, and cause the reasons for carotenoids contents in different ecological areas. Total soluble sugar content of Sichuan Huili tobacco was higher than in Henan Neixiang. The reason lies in the fact that SPS activity and gene expression of Sichuan Huili tobacco was higher than Hernan. However, SS activity and gene expression of tobacco were little difference between Sichuan and Henan.
5. As tobacco was mature, nicotine content in two areas’tobacco both showed an increasing and content of total nitrogen and protein was decreasing. Variation of free amino acids was that it reached a peak on 70 d after transplanting, and then decreased gradually. During the periods 60-100 d after transplanting, NR activity and gene expression gradually decreased, and Huili's always had higher levels than Neixiang's. During the same period, GS gene expression of tobacco both Huili and Neixiang showed a gradual decline, however, Huili's was higher than Neixiang's. GDH gene expression in tobacco of the two areas showed a gradual increase tendency, moreover, Huili's was higher than Neixiang's. 6. At the budding age, in the cured upper leaves, the content of carotenoids in Sichuan tobacco was significantly higher than that in Henan. Results of semi-quantitative RT-PCR of Lyc-βgene and Northern blot show that the Lyc-βgene expression of Sichuan tobacco was significantly higher than Henan tobacco. Expression of Lyc-βgene was regulated by temperature and light and the influence effect is in the order that the light intensity> Temperature> Temperature×light intensity, furthermore, levels among the three factors had no significant difference (P < 0.05). Expression of Lyc-βgene under high light conditions is 1.60 times higher than that in low light. At the temperature of 30℃, its expression reaches the highest, followed by 25℃, the last 35℃. In the range of light time from 2 to 6 h, the expression extension of the gene increased with the extend of the light time, but would reduce when the light time exceeded 8 h.
7. Single-factor effects analysis of Logistic model between meteorological factors and DNA methylation showed that the rise of the average maximum temperature would cause the greatest effect on DNA methylation increase. The extended total sunshine would cause the extension of DNA methylation in tobacco. DNA methylation relies little on mean relative humidity. The increase of total rainfall could reduce the total level of DNA methylation. Although the absolute value of the coefficient was relatively smaller, the value of total rainfall was large enough, which would also get great influence to DNA Methylation. Interaction analysis of two-factor showed that the average maximum temperature and average relative humidity, mean maximum temperature and total sunshine hours would cause a synergistic effect on tobacco DNA methylation; meanwhile, the total rainfall and average maximum temperature, total precipitation and total sunshine Hours, the total rainfall and average relative humidity would lead to antagonism to the DNA methylation. The mean relative humidity and total sunshine hours showed no apparent synergistic effect or antagonistic effect on the level of DNA methylation.
再者,本文建立了烤烟DNA甲基化水平的HPLC检测方法,并以确山烟叶生长期间主要的四个气象因子温度、日照、降水、湿度和烟叶DNA甲基化水平之间建立了Logistic方程的数学模型,研究烟草生长过程中气象因子与DNA甲基化之间的交互作用,从分子水平上探讨气象因子对烟叶生理生化的影响,为深入研究特色烟叶的形成与气象因子的关系提供依据。
本论文通过相关的试验,得出的主要结论如下:
1,河南内乡与四川会理烤烟的生长期不太一致,河南内乡的烟叶前期生长快,后期生长慢;会理的烟叶表现前期慢,后期生长较快。河南内乡与四川会理烟叶可溶性总糖、类胡萝卜素及其降解产物含量的差异较大。河南内乡与四川会理烟叶在移栽后60-100 d期间,烟叶的可溶性总糖的含量变化均为单峰曲线,在移栽后70 d达到高点,随后可溶性总糖的含量逐渐下降,到移栽后100 d时,烟叶中可溶性总糖的含量达到最低。烤后会理烟叶中可溶性总糖比内乡烟叶的要高1.5倍。
2,在烟草移栽后60-80 d烟叶中蔗糖磷酸合成酶(SPS)的活性会理高于内乡烟叶,而到移栽90 d后,会理烟叶中SPS酶活性迅速降低,并且SPS酶活性低于内乡的。在烟草移栽后60-90 d烟叶中蔗糖合酶(SS)的活性会理低于内乡烟叶,而到移栽90 d后,内乡烟叶中SS酶活性降低的较快,并且会理SS酶活性高于内乡的。
3,对烤烟叶圆片进行的温度和光照正交试验的结果表明,温度和光照处理能够下调或上调SPS和SS基因的表达,只有在适当的温度和光照条件下,才有利于SPS基因的表达。在较高温度和光照强度条件下,有利于SS基因的表达。极差的结果表明,三个因子对SPS和SS基因表达的影响作用的大小顺序一样,即(温度×光照强度)>光照强度=温度。Duncan’s测验比较结果表明:温度×光照强度分别对SPS和SS基因的表达影响较大,且达到差异显著水平(P<0.05)。
4,SPS和SS基因的表达量和Northern杂交的结果表明,四川会理烟叶中SPS基因表达水平明显高于河南内乡烟叶。四川会理烟叶中SS基因的杂交带亮度稍高于河南内乡烟叶的条带亮度,表明四川会理烟叶中SS基因表达水平稍高于河南内乡烟叶。现蕾期四川会理烟叶SPS酶活性比河南内乡烟叶提高有利,而两地区烟叶SS酶活性的差异不大。四川会理烟叶的可溶性总糖的含量高于河南内乡的原因是:四川会理烟叶中SPS酶活性和SPSA基因的表达高于内乡烟叶,而四川会理烟叶中SS酶活性和SS基因的表达与河南内乡烟叶的差异不大,使得糖分在烟叶中积累。
5,两地区烟叶中烟碱含量随着烟叶的成熟有逐渐增加的趋势,而烟叶中总氮和蛋白质的含量却随着烟叶的成熟逐渐下降。游离氨基酸的变化为,内乡烟叶在移栽后70 d达到高峰,随后逐渐下降。在移栽后60-100 d期间,NR酶活性和基因表现为逐渐下降,且会理烟叶中NR酶活性一直高于内乡烟叶;河南内乡与四川会理烟叶中GS基因的表达在此期间均表现逐渐下降,GS酶活性也表现为会理烟叶大于内乡;而河南内乡与四川会理烟叶中GDH基因的表达表现为逐渐升高的趋势,且四川会理烟叶中GDH基因的表达比河南内乡烟叶中该基因表达要高。
6,在现蕾期四川会理烤烟上部叶中类胡萝卜素的含量显著高于河南内乡的,基因Lyc-β的半定量RT-PCR和Northern杂交的结果表明,四川会理烟叶中Lyc-β基因表达水平明显高于河南内乡烟叶。Lyc-β基因的表达受到温度和光照的调节,对Lyc-β基因的表达影响的顺序是,光照强度>温度>温度×光照强度,且三因素的三个水平之间没有显著差异(P<0.05)。较高光照的条件下,Lyc-β基因的表达是较低光照的1.60倍;温度为30℃时,Lyc-β基因的表达最高,温度在25℃时次之;温度在35℃时最小。在光照时间为2-6 h期间,Lyc-β基因随着光照时间的延长表达量增加,但光照处理超过8 h之后,Lyc-β基因随着光照时间的延长表达量下调。总之,光照和温度可能是通过调节lyc-β基因的表达造成不同生态区烟叶中类胡萝卜素差异的原因。
7,气象因子与DNA甲基化之间Logistic模型的单因子效应分析表明,平均最高温度的升高对DNA甲基化水平的升高影响最大;总日照长度的延长也会提高烟叶DNA甲基化水平;平均相对湿度对DNA甲基化的影响不大;总降雨量的增加能够降低总DNA甲基化水平,虽然其系数绝对值较小,但总降雨量的数值较大,同样对DNA去甲基化水平的影响幅度也大。总之,平均最高温度、总日照时数、平均相对湿度与DNA甲基化水平正相关;总降雨量与DNA甲基化水平负相关。两因素交互效应分析表明,平均最高温度和平均相对湿度、平均最高温度和总日照时数之间对烟叶的DNA甲基化具有协同效应;总降雨量和平均最高温度、总降雨量和总日照时数、总降雨量和平均相对湿度之间对烟叶的DNA甲基化有拮抗作用;而平均相对湿度与总日照时数之间对烟叶的DNA甲基化水平没有明显的协同效应或拮抗作用。
This paper was designed with the top leaves (the 18th leaf) of flue-cured tobacco (Nicotiana tabacum L.) between Henan and Sichuan for the materials. Sixty days after transplanting, we investigated the tobacco agronomic characters and collected the same position of the 18th leaves in order to detect chemical composition, related enzyme activity, related genes expression, and observations of tissue slice. We sampled after that every 10 days.
Firstly, this paper discussed the differences in the three physicochemical properties: carbon metabolism, nitrogen metabolism and secondary metabolism of tobacco between Henan and Sichuan during 60-100 d after transplanting. Moreover, analyzed the differences of enzyme activity, genes expression related to the three metabolic, and analyzed the causes of the differences from molecular level. Secondly, this paper studied the differences on key genes (SPS, SS, lyc-β) related to sucrose and carotenoids synthetic of tobacco between Henan and Sichuan. This paper also studied the relation between these genes, temperature, and light to build a theoretical and technical basis to regulate these genes expression.
In addition, this paper established Logistic model between tobacco DNA methylation levels and four meteorological factors (temperature, sunshine, rainfall, humidity) during the growth of Queshan tobacco, and researched the interact between meteorological factors during tobacco growth and DNA methylation, and explored meteorological factors’influence to tobacco physiology and biochemistry at the molecular level. In addition, this provides basis to further research the relation between the formation of characteristic leaf tobacco and meteorological factors. The main conclusions of this experiment were as follows:
1. Total thickness of tobacco leaves both in Henan and Sichuan continuously increased after transplanting. The Neixiang tobacco growed fast in the early and slow in late. However, Huili’s growed slow at the earlier time and fast later. During 60-100 d after transplanting, both in Henan and Sichuan, soluble sugar content of tobacco presented single apex curve, 70 d after transplanting was the time that the peak value show, then 100 d after transplanting total soluble sugar content decreased to the lowest. Soluble sugar content of Huili tobacco was 1.5 times higher than Neixiang’s after cured.
2. Sixty-eighty days after transplanting, SPS activity of tobacco in Huili is higher than that of tobacco in Neixiang, however, at 90 d after transplanting, SPS activity of tobacco in Huili decreased rapidly to lower than that in Neixiang. 60-90 d after transplanting, the SS activity of tobacco in Huili is lower than that of Neixiang, and the SS activity in Neixiang decreased rapidly to lower than that of Huili at 90 d after transplanting. Between 60-100 d after transplanting, the activity of sucrose invertase in Huili is higher than Neixiang. During the period 60-100 d after tobacco transplanting, SPS expression of tobacco in Sichuan showed a gradual decline but the level is higher than that in Henan. During the same period, SS expression of tobacco in Sichuan is biased single peak changes, which at the 90 d reached a peak. The genes expression of tobacco had no significant difference in Henan and Sichuan.
3. The leaf discs for temperature and light orthogonal experiment results in laboratory showed that the temperature and light treatment could decrease or raise the SPS and SS genes expression: only a proper temperature and light condition was conducive to the SPS expression. High temperature and light intensity conditions will be helpful for SS gene expression. Influence of the three factors on SPS and SS genes expression is in the order that (temperature×light intensity)> light intensity = temperature. Duncan's test comparison results showed that the condition that the temperature interact light intensity achieves great influence on SPS and SS genes expression, which achieves significant levels (P < 0.05).
4. SPS gene expression and Northern blot results showed that SPS gene expression levels of the tobacco in Sichuan were significantly higher than that of Henan. The result that SS gene hybridization bands brightness of Sichuan Tobacco was slightly higher than that of Henan revealed that SS gene expression in Sichuan tobacco leaves was slightly higher than that in Henan. In budding stage, SPS activity in Huili, Sichuan tobacco leaves is higher than that in Henan, however, SS activity in both regions had no much difference. In short, light and temperature may regulate the expression of lyc-βgene in tobacco, and cause the reasons for carotenoids contents in different ecological areas. Total soluble sugar content of Sichuan Huili tobacco was higher than in Henan Neixiang. The reason lies in the fact that SPS activity and gene expression of Sichuan Huili tobacco was higher than Hernan. However, SS activity and gene expression of tobacco were little difference between Sichuan and Henan.
5. As tobacco was mature, nicotine content in two areas’tobacco both showed an increasing and content of total nitrogen and protein was decreasing. Variation of free amino acids was that it reached a peak on 70 d after transplanting, and then decreased gradually. During the periods 60-100 d after transplanting, NR activity and gene expression gradually decreased, and Huili's always had higher levels than Neixiang's. During the same period, GS gene expression of tobacco both Huili and Neixiang showed a gradual decline, however, Huili's was higher than Neixiang's. GDH gene expression in tobacco of the two areas showed a gradual increase tendency, moreover, Huili's was higher than Neixiang's. 6. At the budding age, in the cured upper leaves, the content of carotenoids in Sichuan tobacco was significantly higher than that in Henan. Results of semi-quantitative RT-PCR of Lyc-βgene and Northern blot show that the Lyc-βgene expression of Sichuan tobacco was significantly higher than Henan tobacco. Expression of Lyc-βgene was regulated by temperature and light and the influence effect is in the order that the light intensity> Temperature> Temperature×light intensity, furthermore, levels among the three factors had no significant difference (P < 0.05). Expression of Lyc-βgene under high light conditions is 1.60 times higher than that in low light. At the temperature of 30℃, its expression reaches the highest, followed by 25℃, the last 35℃. In the range of light time from 2 to 6 h, the expression extension of the gene increased with the extend of the light time, but would reduce when the light time exceeded 8 h.
7. Single-factor effects analysis of Logistic model between meteorological factors and DNA methylation showed that the rise of the average maximum temperature would cause the greatest effect on DNA methylation increase. The extended total sunshine would cause the extension of DNA methylation in tobacco. DNA methylation relies little on mean relative humidity. The increase of total rainfall could reduce the total level of DNA methylation. Although the absolute value of the coefficient was relatively smaller, the value of total rainfall was large enough, which would also get great influence to DNA Methylation. Interaction analysis of two-factor showed that the average maximum temperature and average relative humidity, mean maximum temperature and total sunshine hours would cause a synergistic effect on tobacco DNA methylation; meanwhile, the total rainfall and average maximum temperature, total precipitation and total sunshine Hours, the total rainfall and average relative humidity would lead to antagonism to the DNA methylation. The mean relative humidity and total sunshine hours showed no apparent synergistic effect or antagonistic effect on the level of DNA methylation.
引文
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