增强UV-B辐射和模拟酸雨对C_4植物玉米和苋菜生长与代谢的影响
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摘要
人类活动打破了大气平流层臭氧原有的平衡,使臭氧层变薄,直接导致辐射到地表的UV-B增强。而各国工业除尘和绿化工作的开展降低了大气颗粒物特别是碱性大颗粒物对酸雨的缓冲作用,同时紫外辐射增加有助于细粒子的形成和转化,导致大气酸性增强,使酸雨区以外的地区将来也可能面临酸雨污染问题。
为科学评估全球气候变化背景下环境因子对植物产生的种种生物学和生态学效应,本研究选择NADP-苹果酸酶光合亚型单子叶C4植物玉米[糯玉米(Waxy corn) (Zea mays L. certain Kulesh),品种“渝糯7号”]和NAD-苹果酸酶光合亚型双子叶C4植物苋菜[edible amaranth (Amaranthus mangostanus L.),品种“红圆叶苋菜”]为试验对象,利用生态模拟的研究方法,采用自控的设施模拟酸雨处理[pH6.5(Ao),4.5(A1)和3.5(A2)]和增强UV-B辐射处理[0(Bo),2.88(B1)和5.76(B2,玉米)或4.32(B2',苋菜)kJ-m-2·d-1]处理,研究近似大田环境下,增强UV-B辐射和模拟酸雨及其复合作用对不同类型C4植物生长和种群生理生态的影响,以了解UV-B辐射增强和酸雨区面积扩大的现状下不同类型C4植物特别是重要农作物抵御环境胁迫的机理,为寻找有效手段维持和增加粮食与蔬菜产量,乃至为有关全球气候变化应对措施的研究提供参考和理论支持。本试验主要结果如下：
1.增强UV-B辐射和模拟酸雨单独或复合作用影响玉米和苋菜生物量分配,两种作物生长发育均受到抑制。UV-B辐射下,玉米生育期的滞后主要体现在发育后期,苋菜则从抽苔期即开始发育迟缓。UV-B辐射下玉米的株高和叶面积降低,玉米发育后期株高和叶面积受UV-B辐射影响最严重,而苋菜抽苔期株高和收获期叶面积对UV-B辐射响应最大。UV-B辐射对玉米和苋菜根、茎、叶和果实生物量均有显著影响。除高强度UV-B辐射使玉米根冠比降低外,其余UV-B辐射单独处理时对两种植物根冠比均无显著影响。玉米平均UV-B胁迫反应指数小于苋菜。酸雨对玉米生育期影响最大的时期是拔节期和收获期,而在开花期和收获期对苋菜影响较大。两种植物的株高和叶面积在不同发育时期对酸雨的敏感性也不同,玉米在收获期最为敏感,而酸雨对苋菜叶面积影响最大的时期是抽苔期。玉米的根和果实生物量在酸雨胁迫下降低,而苋菜的茎、叶和果实生物量在酸雨影响下均显著下降。玉米根冠比随酸雨pH降低显著降低,轻度酸雨使苋菜低根冠降低的程度较重度酸雨的大。玉米平均酸雨胁迫反应指数亦较苋菜低。UV-B辐射和模拟酸雨复合处理时,玉米更多地表现为因子间对于生长发育延迟和生物量降低的拮抗作用,而苋菜多表现为协同效应。复合处理下玉米平均胁迫反应指数为0.85,较苋菜平均胁迫反应指数1.73小。相对于整个生长季而言,玉米对逆境因子的耐受性更好,苋菜受到的影响更大。
2.增强UV-B辐射和模拟酸雨单独或复合作用对玉米和苋菜光合生理有较大影响。单独UV-B辐射下,玉米净光合速率随UV-B辐射强度增加显著降低。玉米在发育前期叶片内光合色素(包括叶绿素a、叶绿素b和类胡萝卜素)含量增加,发育后期有所降低；叶绿素a与叶绿素b的比率在拔节期和收获期显著下降,苗期和开花期与对照相比基本无变化；类胡萝卜素与总叶绿素的比率相比对照组无显著性差异。UV-B辐照还造成玉米PEPCase活性、RuBPCase活性,以及PEPCase舌性与RuBPCase活性比率的降低。苋菜净光合速率、光合色素含量、PEPCase舌性、RuBPCase舌性和PEPCase活性与RuBPCase活性比率均随UV-B辐射增强而显著下降,但其叶绿素a与叶绿素b的比率和类胡萝卜素与总叶绿素的比率对照相比则无显著差异。模拟酸雨单独处理时,玉米净光合速率降低,苋菜在发育前期净光合速率几乎不受酸雨影响,在开花期和抽苔期净光合速率下降。酸雨也造成玉米和苋菜叶片内光合色素含量的变化,在玉米发育前期和苋菜所有发育阶段,重度酸雨条下的光合色素含量均较轻度酸雨胁迫下的高；叶绿素a与叶绿素b的比率和类胡萝卜素与总叶绿素的比率大多不受酸雨影响。酸雨造成玉米和苋菜PEPCase活性降低；玉米除收获期外RuBPCase舌性下降,苋菜RuBPCase活性对轻度酸雨不敏感,重度酸雨下除开花期苋菜RuBPCase舌性均下降。苋菜PEPCase活性与RuBPCase舌性比率随酸雨pH值降低而减小,而玉米在发育前期两种酶活性比率上升,除收获期A1处理外其余阶段与对照相比无显著差异。UV-B辐射和模拟酸雨复合处理下更多的表现为两因子的拮抗作用,但植物自我修复能力有限,玉米和苋菜净光合速率降低,光合色素含量发生变化,PEPCase活性和RuBPCase活性下降,PEPCase活性与RuBPCase活性比率大多下降。玉米叶绿素a与叶绿素b的比率在拔节期和收获期显著下降,胡萝卜素与总叶绿素的比率与对照组相比差异不显著；而苋菜叶绿素a与叶绿素b的比率除抽苔期外与对照基本一致,除收获期A282处理外类胡萝卜素与总叶绿素的比率均显著上升。苋菜无论UV-B辐射和模拟酸雨单独或复合处理下平均光合指标的可塑性较均玉米大。UV-B辐射对玉米和苋菜光合生理指标的可塑性大于酸雨。
3.增强UV-B辐射和模拟酸雨单独或复合作用对玉米和苋菜物质代谢有显著影响。UV-B辐射单因子作用时可使玉米和苋菜可溶性蛋白质含量下降,影响最大的时期是收获期和开花期。低强度UV-B辐照下两种作物总游离氨基酸含量除收获期玉米外均较对照有所增大,高强度UV-B辐射除收获期外对两种作物总游离氨基酸含量影响不大。两种作物的可溶性糖含量和ATP含量均随UV-B辐射增强而显著降低,而类黄酮含量和多酚含量则随UV-B辐射增强而增加。酸雨胁迫也会造成玉米和苋菜可溶性蛋白质含量下降,但玉米在拔节期和开花期对A1强度酸雨不敏感,苋菜在抽苔期对两种强度的酸雨均不敏感。酸雨胁迫下玉米和苋菜游离氨基酸含量上升,在整个发育阶段均大于对照且呈现先升后降的趋势；A1条件下两种作物游离氨基酸含量大于A2组。玉米和苋菜可溶性糖含量和ATP含量在酸雨处理时与对照组相比有所降低。酸雨对玉米叶片类黄酮含量基本无影响,对苗期到开花期的苋菜影响不大,在收获期使苋菜黄酮含量显著增加。两种作物多酚含量则随酸雨pH值降低而升高。模拟酸雨和增强UV-B辐射复合处理时在各发育阶段多表现为两因子的拮抗效应,但两种植物可溶性蛋白质含量、可溶性糖含量和ATP含量等代谢物指标仍较对照有所下降,类黄酮含量和多酚含量均增加；各指标都未能恢复到对照水平。玉米和苋菜平均代谢物指标的可塑性指数相等,但就具体的代谢物指标而言,酸雨对两种植物游离氨基酸含量的可塑性较大,而UV-B辐射对其余代谢物指标的可塑性较大。方差分析结果显示UV-B辐射对两种作物代谢的影响更显著。
4.增强UV-B辐射和模拟酸雨单独或复合作用对玉米和苋菜活性氧代谢有明显影响,造成植物膜系统损伤。单独UV-B辐射胁迫时,玉米和苋菜叶片内细胞质膜透性随UV-B辐射的增强显著增大,在这种伤害在发育后期更为显著。两种强度的UV-B辐射均使玉米MDA含量显著增加,但对苋菜而言,高强度UV-B辐射造成MDA含量上升的幅度较低强度UV-B辐射的大。UV-B辐照下玉米SOD活性显著增加,随处理时间延长在收获期SOD活性较发育前期大幅下降,已接近对照组水平。苋菜在发育前期SOD活性显著提高,后期则显著下降。两种植物的POD活性和CAT活性均在UV-B辐射胁迫下降低。模拟酸雨下两种植物细胞质膜透性增大,叶片内MDA含量亦有不同程度的增加。酸雨条件下玉米苋菜SOD活性变化趋势与UV-B辐射胁迫下的类似,POD活性除抽苔期苋菜和开花期A1玉米外均显著降低。轻度酸雨对玉米和苋菜叶片内CAT活性的影响不大,但随着酸雨酸度进一步增加,CAT活性急剧下降。复合处理下较多地表现出因子间的拮抗作用,玉米和苋菜细胞质膜透性在各发育时期均表现为复合处理小于单独UV-B辐照处理,但细胞质膜透性仍大于对照组,MDA含量也较对照显著增加,说明植物利用多种逆境因子交互作用抵御伤害的能力有限。两种物种的保护酶SOD、POD和CAT活性可在逆境因子条件下相互影响和制约以减轻外界伤害。从两种植物平均活性氧代谢指标的可塑性指数来看,苋菜受到的伤害更大,玉米膜系统对UV-B或/和酸雨伤害的抗性较强。从平均活性氧代谢各指标的可塑性指数可以看出UV-B辐射对两种植物膜系统的伤害比酸雨更大。
以上结果说明,增强UV-B辐射对玉米和苋菜的伤害大于酸雨,二者复合处理可在一定程度上降低UV-B辐射的伤害,但其能力有限。在复合处理下,两个因子对不同指标在不同处理时间和复合水平下,表现出一定的协同或拮抗效应可能与作物对环境变化产生一定的抗逆性和适应性有关。在不同时期两种C4植物对紫外辐射与酸雨敏感性不同。本试验条件下与苋菜相比,玉米对外界环境变化的抗性更强。
The balance of ozone layer in stratosphere was broken due to human activities, which reduced the ozone layer and increased the irradiation of ultraviolet-B (UV-B) to earth surface. However, the buffer action of atmospheric particulates on acid rain (AR), especially alkaline large particles, was lowered by industry dust removal and afforested work. Additionally, increased UV irradiation benefits the formation and transformation of fine particles and thus enhancing the acidity of atmosphere. As a result, acid rain issues might occur outside the acid ram regions.
To scientifically estimate the biological and ecological effects of environmental factors on plants considering the global climate changing, NADP-ME C4 photosynthetic pathway monocotyledon plant waxy corn(Zea mays L. certain Kulesh) and NAD-ME C4 photosynthetic pathway dicotyledon plant edible amaranth (Amaranthus mangostanus L.) were studied through ecological simulation. The samples were treated with simulated acid rain [pH6.5 (Ao),4.5 (A1) and 3.5 (A2)] and UV-B irradiation [0 (Bo),2.88 (B,) and 5.76 (B2, maize) or 4.32 (B2, amaranth) kJ·m-2·d-1] used self-control facility. The present research aimed at explored the influences of enhanced UV-B irradiation and simulated acid rain alone or combined each other on individual growth and population physiological ecology of different C4 plants in the similar field conditions. Moreover, the mechanism on different C4 plants, especially important crops, to resist the environmental stress when UV-B irradiation increasing and acid rain area augmenting were also discussed in this dissertation. This research would provide references and theoretical supports to countermeasures of global climate changing and explore the effective means in maintaining and enhancing grains and vegetables yield. Results in this study are as follows:
1. The biomass distributions of maize and edible amaranth were affected by enhanced UV-B irradiation, simulated acid rain or their combination, and the growth and development of both plants were inhibited. Under UV-B irradiation, maize growth period lagged in the late growth stage while that of edible amaranth was in the beginning of bolting stage. The maize height and leaf area reduced and reached the least in the late growth stage, while edible amaranth height in bolting stage and leaf area in harvesting stage showed most significant response to UV-B irradiation. UV-B irradiation significantly affected biomass of root, stem and leaf of both maize and amaranth. The UV-B irradiation had no significant effect on the root/shoot ratio of these two plants except for root/shoot ratio of maize reduced by high level UV-B irradiation. Average stress response index under UV-B stress of maize was lower than that of edible amaranth. On the other hand, acid rain had the most effects on maize in jointing stage and harvesting stage, while flowering stage and harvesting stage were the most vulnerable stages for edible amaranth. The sensitivity of two plants height and leaf area to acid rain varied in different developmental stages:maize was most sensitive in harvesting stage, while leaf area of amaranth was most significant influenced in bolting stage. Root and fruit biomass of maize decreased, and stem, leaf and fruit biomass of amaranth significantly decreased under acid rain conditions. Root/shoot ratio of maize decreased significantly with the decline of acid rain pH, while that of amaranth decreased more under mild acid rain than severe acid rain. Average stress response index in maize under acid rain stress was also less than edible amaranth. What's more, under combination of UV-B and acid rain treatments, maize displayed antagonistic effects on growth delay and biomass decline while edible amaranth showed synergistic effects. The average stress response index of maize was 0.85, lower than that of edible amaranth,1.73. In terms of the whole growth season, maize showed a better tolerance to adversity and edible amaranth was more easily to be influenced.
2. Enhanced UV-B irradiation and simulated acid rain singly and in combination played a great role on plant photosynthesis physiological traits of maize and edible amaranth. Under single UV-B irradiation treatment, net photosynthetic rate of maize decreased with UV-B irradiation increased. In the early development stage of maize, the content of photosynthetic pigments increased, including chlorophyll a, chlorophyll b and carotenoid, but decreased at late development stage. The chlorophyll a/b ratio decreased significantly in jointing stage and harvesting stage, while there showed a stable tendency in seedling and flowering stage in contrast with the control treatment, and no significant difference was found on the ratio of carotenoid and total chlorophyll between treatment and the control treatment. In addition, PEPCase and RuBPCase activity, and the ratio of PEPCase activity/RuBPCase activity of maize decreased under UV-B irradiation. In terms of edible amaranth, net photosynthetic rate, photosynthetic pigment content, PEPCase activity, RuBPCase activity, and PEPCase activity/RuBPCase activity ratio decreased significantly with UV-B irradiation increased. However, its chlorophyll a/b ratio and carotenoid/total chlorophyll ratio had no significant difference to the control treatment. The net photosynthetic rate of maize decreased under simulative acid rain treatment. The net photosynthetic rate was hardly affected in the early development stage of edible amaranth, but decreased in flowering and bolting stage. The content of photosynthetic pigment of maize and edible amaranth influenced by acid rain, at the early development of maize and the whole growth stage of edible amaranth, the content of photosynthetic pigment was higher under severe acid rain than mild acid rain stress. The chlorophyll a/b ratio and the ratio of carotenoid and total chlorophyll were mostly uninfluenced. Under acid rain treatment, PEPCase activity of maize and edible amaranth decreased and RuBPCase activity of maize decreased except in harvesting stage. RuBPCase activity of edible amaranth was not sensitivity to mild acid rain, but it decreased under severe acid rain except in flowering stage. The ratio of PEPCase activity/RuBPCase activity of edible amaranth decreased with decreased pH of acid rain. But the ratio of PEPCase activity/ RuBPCase activity increased in early development stage of maize, except for A1 treatment in harvesting stage, the ratio of the others treatments in various stages had no significant difference. Under the combined stresses of UV-B radiation and simulated acid rain, antagonistic effects between two factors were shown. But the plant self-restoration capability was limited, net photosynthetic rate of maize and edible amaranth decreased, the content of photosynthetic pigment had changed, PEPCase activity and RuBPCase activity decreased, and the ratio of PEPCase activity/RuBPCase activity mostly decreased. The chlorophyll a/b ratio of maize decreased significantly in jointing stage and harvesting stage, the ratio of carotenoid and total chlorophyll had no significant difference compared with the control treatment. In terms of edible amaranth, the chlorophyll a/b ratio was similar to the control treatment except for bolting stage; ratio of carotenoid and total chlorophyll almost increased significantly except for A2B2 treatment in harvesting stage. The edible amaranth had larger mean plasticity index of photosynthetic traits than maize no matter under UV-B irradiation, acid rain or their combination, and the plasticity index of photosynthetic traits of maize and amaranth under UV-B irradiation was better than acid rain.
3. Enhanced UV-B irradiation and simulated acid rain singly and in combination had significant influence on material metabolism of maize and edible amaranth. Under single UV-B irradiation treatment, soluble protein content of maize and edible amaranth decreased, and the harvesting and flowering stage were the most influenced stage. Content of total free amino acid of two crops was higher than the control treatment except for it in harvesting of maize at lower level enhanced UV-B irradiation, while those were little influenced at high level enhanced UV-B irradiation except for in harvesting both of the two plants. Content of soluble sugar and ATP decreased significantly with UV-B irradiation intensity increasing, but flavonoids and polyphenol content increased. The acid rain stress could also cause the decline of soluble protein content of maize and edible amaranth. However, maize was insensitive to A1 treatment of acid rain in jointing stage and flowering stage, while insensitivity was found in edible amaranth in two degrees of acid rain in bolting stage. Content of free amino acid increased along acid rain stress, which was larger than the control treatment in the whole growth stage, with the tendency of increased first but decreased afterwards, and it was higher under A1 condition than A2 in both two plants. It showed a less content of soluble sugar and ATP of maize and edible amaranth under acid rain than the control treatment. There was almost no influence of acid rain to flavonoids of maize leaf under single acid rain treatment, and the same with amaranth from seeding to flowering stage. However, that of edible amaranth increased significantly in harvesting stage. In addition, content of polyphenol in both of the crops increased with decreased pH of simulated acid rain. On the other hand, with the combination of enhanced UV-B irradiation and simulated acid rain, antagonistic effect between two factors was showed. Soluble protein, soluble sugar, and ATP content still had small decline compared with the control treatment, but flavonoids and polyphenol content increased. Every index failed to restore to the former level. Equal to mean plasticity index of various metabolite indices of maize and edible amaranth was found. However, in terms of specific metabolite index, acid rain had greater effect on plasticity of their free amino acid content, while UV-B had larger plasticity to other metabolite index. Variance analysis showed that UV-B irradiation has more significant influence on metabolism of the two crops.
4. Enhanced UV-B irradiation, simulated acid rain or their combination had significant influence on active oxygen metabolism of maize and edible amaranth, damaged plant membrane system. Under single UV-B irradiation treatment, cell membrane permeability of maize and edible amaranth leaf increased significantly with the intensity of enhanced UV-B irradiation, and showed a more significant damage at later developmental stage. MDA content of maize increased significantly under the both levels of enhanced UV-B irradiation, while in terms of edible amaranth, it increased more under high level UV-B irradiation than low level irradiation. SOD activity of maize showed a significant increase under UV-B irradiation, and along with extension of the experimental treatment, SOD activity at harvesting stage, which was close to the control treatment, decreased sharply in contrast with early development stage. SOD activity of edible amaranth in early development stage increased significantly but decreased later. POD and CAT activity of two crops decreased as well. Meanwhile, under simulated acid rain stress, plasmalemma permeability of these two plants improved and leaf MAD content increased to different extents. The change of SOD activity in maize and edible amaranth experienced similar changes under single AR and UV-B treatment. POD activity decreased significantly except for that of edible amaranth in bolting stage and A1 treatment of maize in flowering stage. Light acid rain had little influence on CAT activity of maize and edible amaranth leaves. However, CAT activity decreased sharply in further decreased pH of acid rain. More antagonistic effect of two factors was indicated than additive, synergistic effects under the dual stresses (AR+UV-B). Plasmalemma permeability of maize and edible amaranth in each development stage under combined stress was lower than single UV-B treatment, whiles those still higher than the control treatment, with a same trend as MDA content, and showed a limited ability of plants to utilize interaction of multiple adversity factors to resist impairment. The activity of protective enzymes, such as SOD, POD, and CAT, could interact and alleviate the damage from outside under adversity factors. In terms of the average plasticity index of active oxygen metabolism indices between two crops, edible amaranth hurt even more, and membrane system of maize has better resistance to UV-B irradiation and/or acid rain. Moreover, each plasticity index of active oxygen metabolism indices showed more serious damages under enhanced UV-B irradiation than under on two crops'membrane system.
In sum, the present study suggested that damages caused by enhanced UV-B irradiation on maize and edible amaranth was more serious than acid rain. Impairment from UV-B irradiation could be reduced through combined treatment to some extent, but this capability was limited. The dynamic changes of physiological and biochemical indices under the dual stresses (AR+UV-B) were not identical, and none of them showed a simple linear variation, but displayed additive, synergistic or antagonistic effects depending on the different treatment periods and/or degree of stresses, indicating that plants had different mechanisms of resistance and adaptability under different stresses. The two different C4 plants had different sensitivity to AR and UV-B treatment at different development stages. Maize was in general more tolerant to stresses than edible amaranth under the experimental conditions.
为科学评估全球气候变化背景下环境因子对植物产生的种种生物学和生态学效应,本研究选择NADP-苹果酸酶光合亚型单子叶C4植物玉米[糯玉米(Waxy corn) (Zea mays L. certain Kulesh),品种“渝糯7号”]和NAD-苹果酸酶光合亚型双子叶C4植物苋菜[edible amaranth (Amaranthus mangostanus L.),品种“红圆叶苋菜”]为试验对象,利用生态模拟的研究方法,采用自控的设施模拟酸雨处理[pH6.5(Ao),4.5(A1)和3.5(A2)]和增强UV-B辐射处理[0(Bo),2.88(B1)和5.76(B2,玉米)或4.32(B2',苋菜)kJ-m-2·d-1]处理,研究近似大田环境下,增强UV-B辐射和模拟酸雨及其复合作用对不同类型C4植物生长和种群生理生态的影响,以了解UV-B辐射增强和酸雨区面积扩大的现状下不同类型C4植物特别是重要农作物抵御环境胁迫的机理,为寻找有效手段维持和增加粮食与蔬菜产量,乃至为有关全球气候变化应对措施的研究提供参考和理论支持。本试验主要结果如下：
1.增强UV-B辐射和模拟酸雨单独或复合作用影响玉米和苋菜生物量分配,两种作物生长发育均受到抑制。UV-B辐射下,玉米生育期的滞后主要体现在发育后期,苋菜则从抽苔期即开始发育迟缓。UV-B辐射下玉米的株高和叶面积降低,玉米发育后期株高和叶面积受UV-B辐射影响最严重,而苋菜抽苔期株高和收获期叶面积对UV-B辐射响应最大。UV-B辐射对玉米和苋菜根、茎、叶和果实生物量均有显著影响。除高强度UV-B辐射使玉米根冠比降低外,其余UV-B辐射单独处理时对两种植物根冠比均无显著影响。玉米平均UV-B胁迫反应指数小于苋菜。酸雨对玉米生育期影响最大的时期是拔节期和收获期,而在开花期和收获期对苋菜影响较大。两种植物的株高和叶面积在不同发育时期对酸雨的敏感性也不同,玉米在收获期最为敏感,而酸雨对苋菜叶面积影响最大的时期是抽苔期。玉米的根和果实生物量在酸雨胁迫下降低,而苋菜的茎、叶和果实生物量在酸雨影响下均显著下降。玉米根冠比随酸雨pH降低显著降低,轻度酸雨使苋菜低根冠降低的程度较重度酸雨的大。玉米平均酸雨胁迫反应指数亦较苋菜低。UV-B辐射和模拟酸雨复合处理时,玉米更多地表现为因子间对于生长发育延迟和生物量降低的拮抗作用,而苋菜多表现为协同效应。复合处理下玉米平均胁迫反应指数为0.85,较苋菜平均胁迫反应指数1.73小。相对于整个生长季而言,玉米对逆境因子的耐受性更好,苋菜受到的影响更大。
2.增强UV-B辐射和模拟酸雨单独或复合作用对玉米和苋菜光合生理有较大影响。单独UV-B辐射下,玉米净光合速率随UV-B辐射强度增加显著降低。玉米在发育前期叶片内光合色素(包括叶绿素a、叶绿素b和类胡萝卜素)含量增加,发育后期有所降低；叶绿素a与叶绿素b的比率在拔节期和收获期显著下降,苗期和开花期与对照相比基本无变化；类胡萝卜素与总叶绿素的比率相比对照组无显著性差异。UV-B辐照还造成玉米PEPCase活性、RuBPCase活性,以及PEPCase舌性与RuBPCase活性比率的降低。苋菜净光合速率、光合色素含量、PEPCase舌性、RuBPCase舌性和PEPCase活性与RuBPCase活性比率均随UV-B辐射增强而显著下降,但其叶绿素a与叶绿素b的比率和类胡萝卜素与总叶绿素的比率对照相比则无显著差异。模拟酸雨单独处理时,玉米净光合速率降低,苋菜在发育前期净光合速率几乎不受酸雨影响,在开花期和抽苔期净光合速率下降。酸雨也造成玉米和苋菜叶片内光合色素含量的变化,在玉米发育前期和苋菜所有发育阶段,重度酸雨条下的光合色素含量均较轻度酸雨胁迫下的高；叶绿素a与叶绿素b的比率和类胡萝卜素与总叶绿素的比率大多不受酸雨影响。酸雨造成玉米和苋菜PEPCase活性降低；玉米除收获期外RuBPCase舌性下降,苋菜RuBPCase活性对轻度酸雨不敏感,重度酸雨下除开花期苋菜RuBPCase舌性均下降。苋菜PEPCase活性与RuBPCase舌性比率随酸雨pH值降低而减小,而玉米在发育前期两种酶活性比率上升,除收获期A1处理外其余阶段与对照相比无显著差异。UV-B辐射和模拟酸雨复合处理下更多的表现为两因子的拮抗作用,但植物自我修复能力有限,玉米和苋菜净光合速率降低,光合色素含量发生变化,PEPCase活性和RuBPCase活性下降,PEPCase活性与RuBPCase活性比率大多下降。玉米叶绿素a与叶绿素b的比率在拔节期和收获期显著下降,胡萝卜素与总叶绿素的比率与对照组相比差异不显著；而苋菜叶绿素a与叶绿素b的比率除抽苔期外与对照基本一致,除收获期A282处理外类胡萝卜素与总叶绿素的比率均显著上升。苋菜无论UV-B辐射和模拟酸雨单独或复合处理下平均光合指标的可塑性较均玉米大。UV-B辐射对玉米和苋菜光合生理指标的可塑性大于酸雨。
3.增强UV-B辐射和模拟酸雨单独或复合作用对玉米和苋菜物质代谢有显著影响。UV-B辐射单因子作用时可使玉米和苋菜可溶性蛋白质含量下降,影响最大的时期是收获期和开花期。低强度UV-B辐照下两种作物总游离氨基酸含量除收获期玉米外均较对照有所增大,高强度UV-B辐射除收获期外对两种作物总游离氨基酸含量影响不大。两种作物的可溶性糖含量和ATP含量均随UV-B辐射增强而显著降低,而类黄酮含量和多酚含量则随UV-B辐射增强而增加。酸雨胁迫也会造成玉米和苋菜可溶性蛋白质含量下降,但玉米在拔节期和开花期对A1强度酸雨不敏感,苋菜在抽苔期对两种强度的酸雨均不敏感。酸雨胁迫下玉米和苋菜游离氨基酸含量上升,在整个发育阶段均大于对照且呈现先升后降的趋势；A1条件下两种作物游离氨基酸含量大于A2组。玉米和苋菜可溶性糖含量和ATP含量在酸雨处理时与对照组相比有所降低。酸雨对玉米叶片类黄酮含量基本无影响,对苗期到开花期的苋菜影响不大,在收获期使苋菜黄酮含量显著增加。两种作物多酚含量则随酸雨pH值降低而升高。模拟酸雨和增强UV-B辐射复合处理时在各发育阶段多表现为两因子的拮抗效应,但两种植物可溶性蛋白质含量、可溶性糖含量和ATP含量等代谢物指标仍较对照有所下降,类黄酮含量和多酚含量均增加；各指标都未能恢复到对照水平。玉米和苋菜平均代谢物指标的可塑性指数相等,但就具体的代谢物指标而言,酸雨对两种植物游离氨基酸含量的可塑性较大,而UV-B辐射对其余代谢物指标的可塑性较大。方差分析结果显示UV-B辐射对两种作物代谢的影响更显著。
4.增强UV-B辐射和模拟酸雨单独或复合作用对玉米和苋菜活性氧代谢有明显影响,造成植物膜系统损伤。单独UV-B辐射胁迫时,玉米和苋菜叶片内细胞质膜透性随UV-B辐射的增强显著增大,在这种伤害在发育后期更为显著。两种强度的UV-B辐射均使玉米MDA含量显著增加,但对苋菜而言,高强度UV-B辐射造成MDA含量上升的幅度较低强度UV-B辐射的大。UV-B辐照下玉米SOD活性显著增加,随处理时间延长在收获期SOD活性较发育前期大幅下降,已接近对照组水平。苋菜在发育前期SOD活性显著提高,后期则显著下降。两种植物的POD活性和CAT活性均在UV-B辐射胁迫下降低。模拟酸雨下两种植物细胞质膜透性增大,叶片内MDA含量亦有不同程度的增加。酸雨条件下玉米苋菜SOD活性变化趋势与UV-B辐射胁迫下的类似,POD活性除抽苔期苋菜和开花期A1玉米外均显著降低。轻度酸雨对玉米和苋菜叶片内CAT活性的影响不大,但随着酸雨酸度进一步增加,CAT活性急剧下降。复合处理下较多地表现出因子间的拮抗作用,玉米和苋菜细胞质膜透性在各发育时期均表现为复合处理小于单独UV-B辐照处理,但细胞质膜透性仍大于对照组,MDA含量也较对照显著增加,说明植物利用多种逆境因子交互作用抵御伤害的能力有限。两种物种的保护酶SOD、POD和CAT活性可在逆境因子条件下相互影响和制约以减轻外界伤害。从两种植物平均活性氧代谢指标的可塑性指数来看,苋菜受到的伤害更大,玉米膜系统对UV-B或/和酸雨伤害的抗性较强。从平均活性氧代谢各指标的可塑性指数可以看出UV-B辐射对两种植物膜系统的伤害比酸雨更大。
以上结果说明,增强UV-B辐射对玉米和苋菜的伤害大于酸雨,二者复合处理可在一定程度上降低UV-B辐射的伤害,但其能力有限。在复合处理下,两个因子对不同指标在不同处理时间和复合水平下,表现出一定的协同或拮抗效应可能与作物对环境变化产生一定的抗逆性和适应性有关。在不同时期两种C4植物对紫外辐射与酸雨敏感性不同。本试验条件下与苋菜相比,玉米对外界环境变化的抗性更强。
The balance of ozone layer in stratosphere was broken due to human activities, which reduced the ozone layer and increased the irradiation of ultraviolet-B (UV-B) to earth surface. However, the buffer action of atmospheric particulates on acid rain (AR), especially alkaline large particles, was lowered by industry dust removal and afforested work. Additionally, increased UV irradiation benefits the formation and transformation of fine particles and thus enhancing the acidity of atmosphere. As a result, acid rain issues might occur outside the acid ram regions.
To scientifically estimate the biological and ecological effects of environmental factors on plants considering the global climate changing, NADP-ME C4 photosynthetic pathway monocotyledon plant waxy corn(Zea mays L. certain Kulesh) and NAD-ME C4 photosynthetic pathway dicotyledon plant edible amaranth (Amaranthus mangostanus L.) were studied through ecological simulation. The samples were treated with simulated acid rain [pH6.5 (Ao),4.5 (A1) and 3.5 (A2)] and UV-B irradiation [0 (Bo),2.88 (B,) and 5.76 (B2, maize) or 4.32 (B2, amaranth) kJ·m-2·d-1] used self-control facility. The present research aimed at explored the influences of enhanced UV-B irradiation and simulated acid rain alone or combined each other on individual growth and population physiological ecology of different C4 plants in the similar field conditions. Moreover, the mechanism on different C4 plants, especially important crops, to resist the environmental stress when UV-B irradiation increasing and acid rain area augmenting were also discussed in this dissertation. This research would provide references and theoretical supports to countermeasures of global climate changing and explore the effective means in maintaining and enhancing grains and vegetables yield. Results in this study are as follows:
1. The biomass distributions of maize and edible amaranth were affected by enhanced UV-B irradiation, simulated acid rain or their combination, and the growth and development of both plants were inhibited. Under UV-B irradiation, maize growth period lagged in the late growth stage while that of edible amaranth was in the beginning of bolting stage. The maize height and leaf area reduced and reached the least in the late growth stage, while edible amaranth height in bolting stage and leaf area in harvesting stage showed most significant response to UV-B irradiation. UV-B irradiation significantly affected biomass of root, stem and leaf of both maize and amaranth. The UV-B irradiation had no significant effect on the root/shoot ratio of these two plants except for root/shoot ratio of maize reduced by high level UV-B irradiation. Average stress response index under UV-B stress of maize was lower than that of edible amaranth. On the other hand, acid rain had the most effects on maize in jointing stage and harvesting stage, while flowering stage and harvesting stage were the most vulnerable stages for edible amaranth. The sensitivity of two plants height and leaf area to acid rain varied in different developmental stages:maize was most sensitive in harvesting stage, while leaf area of amaranth was most significant influenced in bolting stage. Root and fruit biomass of maize decreased, and stem, leaf and fruit biomass of amaranth significantly decreased under acid rain conditions. Root/shoot ratio of maize decreased significantly with the decline of acid rain pH, while that of amaranth decreased more under mild acid rain than severe acid rain. Average stress response index in maize under acid rain stress was also less than edible amaranth. What's more, under combination of UV-B and acid rain treatments, maize displayed antagonistic effects on growth delay and biomass decline while edible amaranth showed synergistic effects. The average stress response index of maize was 0.85, lower than that of edible amaranth,1.73. In terms of the whole growth season, maize showed a better tolerance to adversity and edible amaranth was more easily to be influenced.
2. Enhanced UV-B irradiation and simulated acid rain singly and in combination played a great role on plant photosynthesis physiological traits of maize and edible amaranth. Under single UV-B irradiation treatment, net photosynthetic rate of maize decreased with UV-B irradiation increased. In the early development stage of maize, the content of photosynthetic pigments increased, including chlorophyll a, chlorophyll b and carotenoid, but decreased at late development stage. The chlorophyll a/b ratio decreased significantly in jointing stage and harvesting stage, while there showed a stable tendency in seedling and flowering stage in contrast with the control treatment, and no significant difference was found on the ratio of carotenoid and total chlorophyll between treatment and the control treatment. In addition, PEPCase and RuBPCase activity, and the ratio of PEPCase activity/RuBPCase activity of maize decreased under UV-B irradiation. In terms of edible amaranth, net photosynthetic rate, photosynthetic pigment content, PEPCase activity, RuBPCase activity, and PEPCase activity/RuBPCase activity ratio decreased significantly with UV-B irradiation increased. However, its chlorophyll a/b ratio and carotenoid/total chlorophyll ratio had no significant difference to the control treatment. The net photosynthetic rate of maize decreased under simulative acid rain treatment. The net photosynthetic rate was hardly affected in the early development stage of edible amaranth, but decreased in flowering and bolting stage. The content of photosynthetic pigment of maize and edible amaranth influenced by acid rain, at the early development of maize and the whole growth stage of edible amaranth, the content of photosynthetic pigment was higher under severe acid rain than mild acid rain stress. The chlorophyll a/b ratio and the ratio of carotenoid and total chlorophyll were mostly uninfluenced. Under acid rain treatment, PEPCase activity of maize and edible amaranth decreased and RuBPCase activity of maize decreased except in harvesting stage. RuBPCase activity of edible amaranth was not sensitivity to mild acid rain, but it decreased under severe acid rain except in flowering stage. The ratio of PEPCase activity/RuBPCase activity of edible amaranth decreased with decreased pH of acid rain. But the ratio of PEPCase activity/ RuBPCase activity increased in early development stage of maize, except for A1 treatment in harvesting stage, the ratio of the others treatments in various stages had no significant difference. Under the combined stresses of UV-B radiation and simulated acid rain, antagonistic effects between two factors were shown. But the plant self-restoration capability was limited, net photosynthetic rate of maize and edible amaranth decreased, the content of photosynthetic pigment had changed, PEPCase activity and RuBPCase activity decreased, and the ratio of PEPCase activity/RuBPCase activity mostly decreased. The chlorophyll a/b ratio of maize decreased significantly in jointing stage and harvesting stage, the ratio of carotenoid and total chlorophyll had no significant difference compared with the control treatment. In terms of edible amaranth, the chlorophyll a/b ratio was similar to the control treatment except for bolting stage; ratio of carotenoid and total chlorophyll almost increased significantly except for A2B2 treatment in harvesting stage. The edible amaranth had larger mean plasticity index of photosynthetic traits than maize no matter under UV-B irradiation, acid rain or their combination, and the plasticity index of photosynthetic traits of maize and amaranth under UV-B irradiation was better than acid rain.
3. Enhanced UV-B irradiation and simulated acid rain singly and in combination had significant influence on material metabolism of maize and edible amaranth. Under single UV-B irradiation treatment, soluble protein content of maize and edible amaranth decreased, and the harvesting and flowering stage were the most influenced stage. Content of total free amino acid of two crops was higher than the control treatment except for it in harvesting of maize at lower level enhanced UV-B irradiation, while those were little influenced at high level enhanced UV-B irradiation except for in harvesting both of the two plants. Content of soluble sugar and ATP decreased significantly with UV-B irradiation intensity increasing, but flavonoids and polyphenol content increased. The acid rain stress could also cause the decline of soluble protein content of maize and edible amaranth. However, maize was insensitive to A1 treatment of acid rain in jointing stage and flowering stage, while insensitivity was found in edible amaranth in two degrees of acid rain in bolting stage. Content of free amino acid increased along acid rain stress, which was larger than the control treatment in the whole growth stage, with the tendency of increased first but decreased afterwards, and it was higher under A1 condition than A2 in both two plants. It showed a less content of soluble sugar and ATP of maize and edible amaranth under acid rain than the control treatment. There was almost no influence of acid rain to flavonoids of maize leaf under single acid rain treatment, and the same with amaranth from seeding to flowering stage. However, that of edible amaranth increased significantly in harvesting stage. In addition, content of polyphenol in both of the crops increased with decreased pH of simulated acid rain. On the other hand, with the combination of enhanced UV-B irradiation and simulated acid rain, antagonistic effect between two factors was showed. Soluble protein, soluble sugar, and ATP content still had small decline compared with the control treatment, but flavonoids and polyphenol content increased. Every index failed to restore to the former level. Equal to mean plasticity index of various metabolite indices of maize and edible amaranth was found. However, in terms of specific metabolite index, acid rain had greater effect on plasticity of their free amino acid content, while UV-B had larger plasticity to other metabolite index. Variance analysis showed that UV-B irradiation has more significant influence on metabolism of the two crops.
4. Enhanced UV-B irradiation, simulated acid rain or their combination had significant influence on active oxygen metabolism of maize and edible amaranth, damaged plant membrane system. Under single UV-B irradiation treatment, cell membrane permeability of maize and edible amaranth leaf increased significantly with the intensity of enhanced UV-B irradiation, and showed a more significant damage at later developmental stage. MDA content of maize increased significantly under the both levels of enhanced UV-B irradiation, while in terms of edible amaranth, it increased more under high level UV-B irradiation than low level irradiation. SOD activity of maize showed a significant increase under UV-B irradiation, and along with extension of the experimental treatment, SOD activity at harvesting stage, which was close to the control treatment, decreased sharply in contrast with early development stage. SOD activity of edible amaranth in early development stage increased significantly but decreased later. POD and CAT activity of two crops decreased as well. Meanwhile, under simulated acid rain stress, plasmalemma permeability of these two plants improved and leaf MAD content increased to different extents. The change of SOD activity in maize and edible amaranth experienced similar changes under single AR and UV-B treatment. POD activity decreased significantly except for that of edible amaranth in bolting stage and A1 treatment of maize in flowering stage. Light acid rain had little influence on CAT activity of maize and edible amaranth leaves. However, CAT activity decreased sharply in further decreased pH of acid rain. More antagonistic effect of two factors was indicated than additive, synergistic effects under the dual stresses (AR+UV-B). Plasmalemma permeability of maize and edible amaranth in each development stage under combined stress was lower than single UV-B treatment, whiles those still higher than the control treatment, with a same trend as MDA content, and showed a limited ability of plants to utilize interaction of multiple adversity factors to resist impairment. The activity of protective enzymes, such as SOD, POD, and CAT, could interact and alleviate the damage from outside under adversity factors. In terms of the average plasticity index of active oxygen metabolism indices between two crops, edible amaranth hurt even more, and membrane system of maize has better resistance to UV-B irradiation and/or acid rain. Moreover, each plasticity index of active oxygen metabolism indices showed more serious damages under enhanced UV-B irradiation than under on two crops'membrane system.
In sum, the present study suggested that damages caused by enhanced UV-B irradiation on maize and edible amaranth was more serious than acid rain. Impairment from UV-B irradiation could be reduced through combined treatment to some extent, but this capability was limited. The dynamic changes of physiological and biochemical indices under the dual stresses (AR+UV-B) were not identical, and none of them showed a simple linear variation, but displayed additive, synergistic or antagonistic effects depending on the different treatment periods and/or degree of stresses, indicating that plants had different mechanisms of resistance and adaptability under different stresses. The two different C4 plants had different sensitivity to AR and UV-B treatment at different development stages. Maize was in general more tolerant to stresses than edible amaranth under the experimental conditions.
引文
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