烤烟（Nicotiana tabacum L.）对增强紫外线-B辐射的生理响应与适应性研究
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摘要
大气臭氧层的衰减,会导致地球表面具有生物学效应的日光UV-B辐射逐渐增强。本研究以云南两个烤烟主栽品种“云烟87”和“红花大金元”为试验材料,模拟昆明地区在大气臭氧层衰减24.65%、39.53%条件下,UV-B辐射将分别增强为：5.30kJ·m-2·d-1(T1)、8.50kJ·m-2·d-1(T2)。通过整个烤烟大田条件下模拟增强UV-B辐射,研究了两个烤烟品种的生长发育、细胞抗氧化能力、光合作用和烤后烟叶化学成分受到的影响。并希望通过研究揭示大气臭氧层衰减后,UV-B辐射增强对烤烟生长和烟叶质量变化形成机理。结果表明：增加低强度UV-B辐射可明显促进了两个烤烟品种的生长,也促进了烤烟的发育进程,使烤烟现蕾期、始花期和盛花期提前。两个烤烟品种的株高和茎粗也是以低UV-B辐射最大,其次是对照,高剂量的UV-B辐射处理上述指标最小。高强度的UV-B辐射可使云烟87品种的节距、叶面积的生长受到抑制。同时增加低强度UV-B辐射可改善了两个烤烟品种叶片的组织结构。
增强UV-B辐射可显著提高两个烤烟品种叶片膜脂过氧化的程度。随着UV-B辐射的增强,两个品种的SOD、POD、CAT活性均表现显著增加,可溶性蛋白质含量也显著增加,脯氨酸含量显著下降。低强度UV-B辐射促进了两个烤烟品种多酚氧化酶活性的升高和其它可吸收UV-B物质的含量增加,但高强度UV-B辐射可使多酚氧化酶活性和其它可吸收V-B物质含量受到一定的抑制。低强度UV-B辐射促进了两个烤烟品种叶绿素a、叶绿素b和总叶绿素含量的增加,高强度UV-B辐射使叶绿素结构受到破环,叶绿素a、叶绿素b和总叶绿素含量下降。增强UV-B辐射对两个品种的类胡萝卜素含量影响不一致,红大品种的类胡萝卜素含量以低强度UV-B辐射的最高,随着UV-B辐射的进一步增强其含量又降低,而云烟87的类胡萝卜素含量在增强UV-B辐射和对照之间无差别。两个烤烟品种的光合速率、气孔导度和蒸腾速率对增强UV-B辐射的响应不同。红大品种的光合速率、气孔导度和蒸腾速率随着UV-B辐射的增强而增大；而云烟87的光合速率、气孔导度和蒸腾速率均以低强度UV-B辐射下达最高,高强度UV-B辐射下又降低。增强UV-B辐射对两个烤烟品种胞间CO2浓度的响应基本一致,都随着UV-B辐射的增强先减小,高强度UV-B辐射下又增加。
增强UV-B辐射使两个烤烟品种的水溶性糖含量降低,总氮含量提高,增强UV-B辐射使云烟87水溶性糖含量和总氮含量的变化幅度比红大的高,说明云烟87较红大品种对UV-B辐射增强更敏感。增强UV-B辐射可提高两个烤烟品种的全钾、游离氨基酸、质体色素和类黄酮含量。增强UV-B辐射对红大和云烟87品种的烟碱含量影响不同,云烟87的烟碱含量随UV-B辐射增大而增加,而红大品种烟碱的含量却表现出相反的变化。UV-B辐射增强可使两个烤烟品种邻苯二酚的含量降低。
通过对22个烤烟品种苗期增强UV-B辐射试验,增强UV-B辐射对大多烤烟品种幼苗株高、茎围、最大叶长、最大叶宽、地上部干重、地下部干重及鲜重都有明显的抑制作用,且比相同品种大田期的抑制程度要高。在UV-B辐射下,大多数烤烟品种叶片中膜脂过氧化程度明显提高,且超氧化物歧化酶和多酚氧化酶的活性受到的影响更大,脯氨酸和其它紫外吸收物质含量也明显增加。从增强UV-B辐射对烤烟光合色素的影响分析,UV-B辐射对大多数烤烟品种的叶绿素a和类胡萝卜素含量具有明显的抑制作用。这也是在苗期大多数烤烟品种生长受抑制的主要原因之一通过烤烟幼苗UV-B生长响应指数和总叶绿素含量变化值综合评价烤烟品种对UV-B的敏感性,结果显示云烟87和云烟85在幼苗期对UV-B耐受性相对较强,而KRK26和红大的耐受性相对较弱,这和大田期评价UV-B敏感性的研究结果差异较大。
通过在大田期长期增强UV-B辐射对不同烤烟品种生长和光合作用的试验表明,增强UV-B辐射对烤烟的生长和光合作用有显著的作用,UV-B辐射对一些品种生长有刺激作用,但对多数品种生长具有抑制作用。增强UV-B辐射对大多数品种的最大叶长、最大叶宽和茎围影响不大,具有刺激作用的品种较具有抑制作用的品种多。株高和节距对UV-B辐射的响应基本一致,表现出抑制作用的品种多于有促进作用品种。增强UV-B辐射对大多数品种的叶绿素a、叶绿素b、总叶绿素、光合速率、气孔导度、蒸腾速率和最低叶绿素荧光具有抑制作用,对叶绿素a和叶绿素b的比值、类胡萝卜素、类胡萝卜素和总叶绿索的比值、胞间CO2浓度、最大叶绿素荧光值和最大量子效率具有促进作用。其中总叶绿素/光合速率、光合速率/气孔导度、光合速率/蒸腾速率、总叶绿素/最低叶绿素荧光、叶绿素a/叶绿素b的比值与最大量子效率、最大叶绿素荧光和最大量子效率等密切相关。通过UV-B响应指数、总叶绿素含量的变化和Fv/Fm的变化值评价烤烟品种对UV-B的敏感性,所得结果不太一致。综合来看,Fv/Fm变化值的评价结果和生产实际更为接近,但与烤烟品种的来源地关系不大
The depletion of the ozone led to the increase of ultraviolet-B(UV-B) with biological effects in the earth's surface. In the first experiment of our study, the effects of enhanced UV-B radiation on growth, development, antioxidant capacity, photosynthesis and chemical components contents were studied in the leaves of two flue-cured tobacco cultivars-"Yunyan87"(Y87) and "Honghuadajinyuan"(HD). Selected cultivars were exposed to UV-B radiation at ambient(control, CK), ambient plus5.30kJ·m-2·d-1(T1) and ambient plus8.50kJ·m-2·d-1(T2) levels that respectively simulated24.65%and39.53%ozone depletions under summer solstice field conditions in Kunming City (located at25°04' N and102°73'E). The study was meant to provide theoretical basis for the formation mechanism of quality of flue-cured tobacco leaves in different ambient UV-B regions. The results were as follows:
Low UV-B radiation significantly promote the growth of two flue-cured tobacco varieties, but also to promote reproductive developmental process of flue-cured tobacco, make squaring period, initial time of flowering and full-bloom stage shorten. The plant height and stem diameter of low UV-B radiation were the largest, whereas high UV-B radiation were the least. Influential tendencies of UV-B radiation on internode and plant height of HD are just the same. And the internode of Y87was decrease with enhanced UV-B radiation. The length, width and leaf area of HD were increase with enhanced UV-B radiation, whereas those of Y87were the largest under the low UV-B radiation condition. The low UV-B radiation promoted the increase of leaf area index, weight per unit leaf area and weight per leaf of both cultivars. Compared to control, the high UV-B radiation decreased weight per unit leaf area and weight per leaf of both cultivars, and increased leaf area index. The thickness of palisade tissues, leaf thickness, the ratio of palisade tissues thickness to spongy and to leaf thickness of low UV-B radiation were the largest, whereas high UV-B radiation were the least. Change in plant height. internode and leaf area in response to UV-B radiation indicated that Y87might be more sensitive to UV-B stress. The low UV-B radiation can improve the growth and anatomical structure of tobacco leaves. UV-B radiation remarkably enhances MDA content level of two flue-cured tobacco varieties leaves. With the enhancement of UV-B radiation, two flue-cured tobacco varieties leaves of SOD and POD. CAT activity increased significantly, and soluble protein content also increased significantly, and proline content decreased significantly. Low UV-B radiation promoted polyphenol oxidase activity and UV-B absorbing substances of two flue-cured tobacco varieties increased, but high UV-B radiation made polyphenol oxidase activity and UV-B absorption material by certain inhibition. Low UV-B radiation also promoted chlorophyll a, chlorophyll b and total chlorophyll content of two flue-cured tobacco varieties to increase, while high UV-B radiation made chlorophyll a, chlorophyll b and total chlorophyll content decreased due to the broken chlorophyll structure by high UV-B radiation. Effect of enhanced UV-B radiation on the carotenoid content of two varieties was inconsistent, the carotenoid content of HD under low UV-B radiation condition was the highest, with the further enhancement of the UV-B radiation and its content decreased, while the carotenoid of Y87is no difference between the enhanced UV-B radiation and control. Responses of two flue-cured tobacco varieties of photosynthetic rate, transpiration rate and stomatal conductance to enhance UV-B radiation were different. Photosynthetic rate, stomatal conductance and transpiration rate of HD increases with the enhancement of UV-B radiation, while photosynthetic rate, stomatal conductance and transpiration rate of Y87are highest under low UV-B radiation condition, those are lower under high UV-B radiation condition. Enhance UV-B radiation on intercellular CO2concentration of two flue-cured tobacco varieties has almost the same response, as low UV-B radiation to reduce, high UV-B radiation to increase. Water soluble total sugar and catechol levels declined markedly in leaves of both cultivars with two doses of UV-B radiation compared with CK. However, total nitrogen, potassium, free amino acids, chromoplast pigment and flavonoid levels increased in leaves of both cultivars. While nicotine level increased in the leaves of Y87, it decreased in the leaves of HD under two doses of UV-B radiation compared with CK. Changes in water soluble total sugar and total nitrogen levels in response to UV-B radiation suggested that Y87was more sensitive to UV-B stress than HD.
Through short-term UV-B radiation experiment of22flue-cured tobacco varieties seedling showed that enhanced UV-B radiation on most of flue-cured tobacco varieties the plant height, stem circumference, maximum leaf length, maximum leaf width, shoot dry weight, underground dry weightand fresh weight has a significant inhibitory effect, and higher than the degree of inhibition of the field period of the same varieties. UV-B radiation made leaf membrane lipid peroxidation of most of flue-cured tobacco varieties increased significantly, mainly to improve the activity of superoxide dismutase and polyphenol oxidase, as well as increasing proline and UV absorbing compounds to protect the flue-cured tobacco leaves from reactive oxygen and membrane lipid peroxidation harm. UV-B radiation significantly inhibited the chlorophyll a and carotenoids of most of the flue-cured tobacco varieties. This is also why the growth of most of the flue-cured tobacco varieties were inhibited. Through the comprehensive evaluation to UV-B sensitivity of growth response index and total chlorophyll content change value of the flue-cured tobacco varieties seedling, the results showed that Y87and Y85in seedling stage to UV-B patience were relatively strong, and the KRK26and HD patience were relatively weak, this result is different from the field period.
Through field experimengt of the effect of long-term enhanced UV-B radiation on growth and photosynthesis of the different flue-cured tobacco varieties showed that enhanced UV-B radiation on growth and photosynthesis of flue-cured tobacco varieties have a remarkable effect, UV-B radiation have a stimulating effect on some varieties, but for most varieties inhibited. Enhanced UV-B radiation has little effect to most of varieties of leaf length, maximum leaf width and stem circumference, has a stimulating effect on the number of varieties than inhibit the number of varieties. The response of plant height and pitch to UV-B radiation is consistent, the inhibition of the number of varieties is higher than the number of varieties of the stimulation. Enhanced UV-B radiation on most varieties of chlorophyll a, chlorophyll b, total chlorophyll, photosynthetic rate, transpiration rate, stomatal conductance and chlorophyll fluorescence has minimum inhibition. The ratio of chlorophyll a and chlorophyll b, carotenoids, carotene and total chlorophyll ratio, the intercellular CO2concentration, the maximum chlorophyll fluorescence and the maximum quantum efficiency had stimulative effect. The total chlorophyll and photosynthetic rate, photosynthetic rate and stomatal conductance, photosynthetic rate and transpiration rate, total chlorophyll and the lowest chlorophyll fluorescence, chlorophyll a and chlorophyll b ratio and the maximum quantum efficiency, the maximum chlorophyll fluorescence and the maximum quantum efficiency are closely related. Three kinds of results, among UV-B response index, total chlorophyll content changes with Fv/Fm change value to evaluate flue-cured tobacco varieties to UV-B sensitivity are less consistent. Taken together, Fv/Fm change value and the results of the evaluation of the actual production are close, but the results had little relation with the origin of flue-cured tobacco varieties.
增强UV-B辐射可显著提高两个烤烟品种叶片膜脂过氧化的程度。随着UV-B辐射的增强,两个品种的SOD、POD、CAT活性均表现显著增加,可溶性蛋白质含量也显著增加,脯氨酸含量显著下降。低强度UV-B辐射促进了两个烤烟品种多酚氧化酶活性的升高和其它可吸收UV-B物质的含量增加,但高强度UV-B辐射可使多酚氧化酶活性和其它可吸收V-B物质含量受到一定的抑制。低强度UV-B辐射促进了两个烤烟品种叶绿素a、叶绿素b和总叶绿素含量的增加,高强度UV-B辐射使叶绿素结构受到破环,叶绿素a、叶绿素b和总叶绿素含量下降。增强UV-B辐射对两个品种的类胡萝卜素含量影响不一致,红大品种的类胡萝卜素含量以低强度UV-B辐射的最高,随着UV-B辐射的进一步增强其含量又降低,而云烟87的类胡萝卜素含量在增强UV-B辐射和对照之间无差别。两个烤烟品种的光合速率、气孔导度和蒸腾速率对增强UV-B辐射的响应不同。红大品种的光合速率、气孔导度和蒸腾速率随着UV-B辐射的增强而增大；而云烟87的光合速率、气孔导度和蒸腾速率均以低强度UV-B辐射下达最高,高强度UV-B辐射下又降低。增强UV-B辐射对两个烤烟品种胞间CO2浓度的响应基本一致,都随着UV-B辐射的增强先减小,高强度UV-B辐射下又增加。
增强UV-B辐射使两个烤烟品种的水溶性糖含量降低,总氮含量提高,增强UV-B辐射使云烟87水溶性糖含量和总氮含量的变化幅度比红大的高,说明云烟87较红大品种对UV-B辐射增强更敏感。增强UV-B辐射可提高两个烤烟品种的全钾、游离氨基酸、质体色素和类黄酮含量。增强UV-B辐射对红大和云烟87品种的烟碱含量影响不同,云烟87的烟碱含量随UV-B辐射增大而增加,而红大品种烟碱的含量却表现出相反的变化。UV-B辐射增强可使两个烤烟品种邻苯二酚的含量降低。
通过对22个烤烟品种苗期增强UV-B辐射试验,增强UV-B辐射对大多烤烟品种幼苗株高、茎围、最大叶长、最大叶宽、地上部干重、地下部干重及鲜重都有明显的抑制作用,且比相同品种大田期的抑制程度要高。在UV-B辐射下,大多数烤烟品种叶片中膜脂过氧化程度明显提高,且超氧化物歧化酶和多酚氧化酶的活性受到的影响更大,脯氨酸和其它紫外吸收物质含量也明显增加。从增强UV-B辐射对烤烟光合色素的影响分析,UV-B辐射对大多数烤烟品种的叶绿素a和类胡萝卜素含量具有明显的抑制作用。这也是在苗期大多数烤烟品种生长受抑制的主要原因之一通过烤烟幼苗UV-B生长响应指数和总叶绿素含量变化值综合评价烤烟品种对UV-B的敏感性,结果显示云烟87和云烟85在幼苗期对UV-B耐受性相对较强,而KRK26和红大的耐受性相对较弱,这和大田期评价UV-B敏感性的研究结果差异较大。
通过在大田期长期增强UV-B辐射对不同烤烟品种生长和光合作用的试验表明,增强UV-B辐射对烤烟的生长和光合作用有显著的作用,UV-B辐射对一些品种生长有刺激作用,但对多数品种生长具有抑制作用。增强UV-B辐射对大多数品种的最大叶长、最大叶宽和茎围影响不大,具有刺激作用的品种较具有抑制作用的品种多。株高和节距对UV-B辐射的响应基本一致,表现出抑制作用的品种多于有促进作用品种。增强UV-B辐射对大多数品种的叶绿素a、叶绿素b、总叶绿素、光合速率、气孔导度、蒸腾速率和最低叶绿素荧光具有抑制作用,对叶绿素a和叶绿素b的比值、类胡萝卜素、类胡萝卜素和总叶绿索的比值、胞间CO2浓度、最大叶绿素荧光值和最大量子效率具有促进作用。其中总叶绿素/光合速率、光合速率/气孔导度、光合速率/蒸腾速率、总叶绿素/最低叶绿素荧光、叶绿素a/叶绿素b的比值与最大量子效率、最大叶绿素荧光和最大量子效率等密切相关。通过UV-B响应指数、总叶绿素含量的变化和Fv/Fm的变化值评价烤烟品种对UV-B的敏感性,所得结果不太一致。综合来看,Fv/Fm变化值的评价结果和生产实际更为接近,但与烤烟品种的来源地关系不大
The depletion of the ozone led to the increase of ultraviolet-B(UV-B) with biological effects in the earth's surface. In the first experiment of our study, the effects of enhanced UV-B radiation on growth, development, antioxidant capacity, photosynthesis and chemical components contents were studied in the leaves of two flue-cured tobacco cultivars-"Yunyan87"(Y87) and "Honghuadajinyuan"(HD). Selected cultivars were exposed to UV-B radiation at ambient(control, CK), ambient plus5.30kJ·m-2·d-1(T1) and ambient plus8.50kJ·m-2·d-1(T2) levels that respectively simulated24.65%and39.53%ozone depletions under summer solstice field conditions in Kunming City (located at25°04' N and102°73'E). The study was meant to provide theoretical basis for the formation mechanism of quality of flue-cured tobacco leaves in different ambient UV-B regions. The results were as follows:
Low UV-B radiation significantly promote the growth of two flue-cured tobacco varieties, but also to promote reproductive developmental process of flue-cured tobacco, make squaring period, initial time of flowering and full-bloom stage shorten. The plant height and stem diameter of low UV-B radiation were the largest, whereas high UV-B radiation were the least. Influential tendencies of UV-B radiation on internode and plant height of HD are just the same. And the internode of Y87was decrease with enhanced UV-B radiation. The length, width and leaf area of HD were increase with enhanced UV-B radiation, whereas those of Y87were the largest under the low UV-B radiation condition. The low UV-B radiation promoted the increase of leaf area index, weight per unit leaf area and weight per leaf of both cultivars. Compared to control, the high UV-B radiation decreased weight per unit leaf area and weight per leaf of both cultivars, and increased leaf area index. The thickness of palisade tissues, leaf thickness, the ratio of palisade tissues thickness to spongy and to leaf thickness of low UV-B radiation were the largest, whereas high UV-B radiation were the least. Change in plant height. internode and leaf area in response to UV-B radiation indicated that Y87might be more sensitive to UV-B stress. The low UV-B radiation can improve the growth and anatomical structure of tobacco leaves. UV-B radiation remarkably enhances MDA content level of two flue-cured tobacco varieties leaves. With the enhancement of UV-B radiation, two flue-cured tobacco varieties leaves of SOD and POD. CAT activity increased significantly, and soluble protein content also increased significantly, and proline content decreased significantly. Low UV-B radiation promoted polyphenol oxidase activity and UV-B absorbing substances of two flue-cured tobacco varieties increased, but high UV-B radiation made polyphenol oxidase activity and UV-B absorption material by certain inhibition. Low UV-B radiation also promoted chlorophyll a, chlorophyll b and total chlorophyll content of two flue-cured tobacco varieties to increase, while high UV-B radiation made chlorophyll a, chlorophyll b and total chlorophyll content decreased due to the broken chlorophyll structure by high UV-B radiation. Effect of enhanced UV-B radiation on the carotenoid content of two varieties was inconsistent, the carotenoid content of HD under low UV-B radiation condition was the highest, with the further enhancement of the UV-B radiation and its content decreased, while the carotenoid of Y87is no difference between the enhanced UV-B radiation and control. Responses of two flue-cured tobacco varieties of photosynthetic rate, transpiration rate and stomatal conductance to enhance UV-B radiation were different. Photosynthetic rate, stomatal conductance and transpiration rate of HD increases with the enhancement of UV-B radiation, while photosynthetic rate, stomatal conductance and transpiration rate of Y87are highest under low UV-B radiation condition, those are lower under high UV-B radiation condition. Enhance UV-B radiation on intercellular CO2concentration of two flue-cured tobacco varieties has almost the same response, as low UV-B radiation to reduce, high UV-B radiation to increase. Water soluble total sugar and catechol levels declined markedly in leaves of both cultivars with two doses of UV-B radiation compared with CK. However, total nitrogen, potassium, free amino acids, chromoplast pigment and flavonoid levels increased in leaves of both cultivars. While nicotine level increased in the leaves of Y87, it decreased in the leaves of HD under two doses of UV-B radiation compared with CK. Changes in water soluble total sugar and total nitrogen levels in response to UV-B radiation suggested that Y87was more sensitive to UV-B stress than HD.
Through short-term UV-B radiation experiment of22flue-cured tobacco varieties seedling showed that enhanced UV-B radiation on most of flue-cured tobacco varieties the plant height, stem circumference, maximum leaf length, maximum leaf width, shoot dry weight, underground dry weightand fresh weight has a significant inhibitory effect, and higher than the degree of inhibition of the field period of the same varieties. UV-B radiation made leaf membrane lipid peroxidation of most of flue-cured tobacco varieties increased significantly, mainly to improve the activity of superoxide dismutase and polyphenol oxidase, as well as increasing proline and UV absorbing compounds to protect the flue-cured tobacco leaves from reactive oxygen and membrane lipid peroxidation harm. UV-B radiation significantly inhibited the chlorophyll a and carotenoids of most of the flue-cured tobacco varieties. This is also why the growth of most of the flue-cured tobacco varieties were inhibited. Through the comprehensive evaluation to UV-B sensitivity of growth response index and total chlorophyll content change value of the flue-cured tobacco varieties seedling, the results showed that Y87and Y85in seedling stage to UV-B patience were relatively strong, and the KRK26and HD patience were relatively weak, this result is different from the field period.
Through field experimengt of the effect of long-term enhanced UV-B radiation on growth and photosynthesis of the different flue-cured tobacco varieties showed that enhanced UV-B radiation on growth and photosynthesis of flue-cured tobacco varieties have a remarkable effect, UV-B radiation have a stimulating effect on some varieties, but for most varieties inhibited. Enhanced UV-B radiation has little effect to most of varieties of leaf length, maximum leaf width and stem circumference, has a stimulating effect on the number of varieties than inhibit the number of varieties. The response of plant height and pitch to UV-B radiation is consistent, the inhibition of the number of varieties is higher than the number of varieties of the stimulation. Enhanced UV-B radiation on most varieties of chlorophyll a, chlorophyll b, total chlorophyll, photosynthetic rate, transpiration rate, stomatal conductance and chlorophyll fluorescence has minimum inhibition. The ratio of chlorophyll a and chlorophyll b, carotenoids, carotene and total chlorophyll ratio, the intercellular CO2concentration, the maximum chlorophyll fluorescence and the maximum quantum efficiency had stimulative effect. The total chlorophyll and photosynthetic rate, photosynthetic rate and stomatal conductance, photosynthetic rate and transpiration rate, total chlorophyll and the lowest chlorophyll fluorescence, chlorophyll a and chlorophyll b ratio and the maximum quantum efficiency, the maximum chlorophyll fluorescence and the maximum quantum efficiency are closely related. Three kinds of results, among UV-B response index, total chlorophyll content changes with Fv/Fm change value to evaluate flue-cured tobacco varieties to UV-B sensitivity are less consistent. Taken together, Fv/Fm change value and the results of the evaluation of the actual production are close, but the results had little relation with the origin of flue-cured tobacco varieties.
引文
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