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恩施州不同海拔下烤烟光合作用与产量、质量的差异性研究

Study on Difference of Photosynthesis, Yield and Quality of Flue-cured Tobacco at Different Altitudes in Enshi

【作者】 王瑞

【导师】 刘国顺;

【作者基本信息】 河南农业大学 , 烟草学, 2010, 博士

【摘要】 以3个遗传背景不同的烤烟品种K326、云烟87、中烟103为材料,在湖北省恩施地区,选择同一山脉的3个不同海拔点(500 m、900 m、1300 m)进行盆栽和大田试验,研究了不同海拔下烤烟光合作用与产、质量的差异。研究的主要内容为:烤烟叶片全展后气体交换参数、光合色素含量、RuBPcase初始活性以及叶绿体超微结构的变化;烤烟叶片全展时光合日变化及其与环境因子的关系;烤烟叶片全展后碳、氮物质以及部分次生代谢物质的变化;烤后烟叶产、质量及其香型风格的分析;光合生产能力与烤烟产、质量的关系。主要研究结果如下:(1)烤烟叶片全展后,光合功能不断衰退。随着海拔的升高,光合功能衰退的越慢。主要表现为Pn、光合色素含量以及RuBPCase的初始活性的下降率降低,叶绿体结构和类囊体片层结构解体时间推迟,净光合速率高值持续期(APD)延长,叶源量(LSC)和单位面积叶源量(PALSC)增加。通径分析表明,APD对LSC和PALSC直接贡献最大,因此延长烤烟光合功能期是提高烤烟叶片光合生产能力的主要措施。(2)随着海拔的升高,日光合总量(∑Pn)、单位面积日光合总量(PA∑Pn)以及水分利用率(WUE)逐渐增加,而日蒸腾总量(∑Tr)和单位面积日蒸腾总量(PA∑Tr)以海拔900 m最高,海拔500 m次之。在晴天中午时段,烤烟叶片表现出“光合抑制”的现象,随着海拔的升高,受抑制的程度减轻。高温加剧了强光对烤烟的光合抑制程度,是影响光合作用的主要限制因子。影响光合日变化的主要生理因子是气孔导度(Gs),生态因子因海拔不同而有所差异。在海拔500 m,环境温度(Ta)和空气相对湿度(RH)是Pn提高的制约因子。在海拔900 m,烤烟叶片光合作用主要受光通量密度(PFD)的影响。在海拔1300 m,环境温度(Ta)和空气湿度(RH)是影响Pn的主要因子。(3)生长在不同海拔下的烤烟光合生理特性存在一定的差异。随着海拔的升高,烤烟对强光的适应性和利用强光的能力增强,对弱光的适应性和利用弱光的能力降低。相对低海拔而言,生长在高海拔地区烤烟有着较高的碳吸收能力。Pn随光强的变化规律可用方程Pn = Pn max (1-C0exp(-aPFD/ Pn max)进行模拟,随CO2浓度(Ci)的变化规律可用方程Pn= Pn max (1-exp(-a Ci / Pn max)-Rd进行模拟。(4)烤烟叶片全展后,光合产物主要以淀粉的形式积累。随着生育进程的推进,淀粉含量、碳水化合物总量、比叶重以及碳氮比值呈先上升后下降的趋势。随着海拔的升高,光合产物以及干物质积累的时间延长。淀粉含量、碳水化合物总量、比叶重以及碳氮比值随着叶片全展后天数的变化规律均可用二次多项式进行模拟。叶片全展后,叶中N含量呈下降趋势。随着海拔的升高,叶中N含量的下降率降低。叶中N含量随着叶片全展后天数的变化规律均可用渐进回归模型进行模拟。叶中N含量与叶绿素含量、Pn均呈极显著正相关。(5)海拔对烤烟叶片叶源量(LSC)、单位面积叶源量(PALSC)、日光合总量(∑Pn)以及单位面积日光合总量(PA∑Pn)的影响均达到极显著水平。品种仅对PALSC的影响达到显著水平,而对其他3个光合生产能力指标的影响不显著。海拔与品种的互作效应对烤烟各光合生产能力指标的影响均达到极显著水平。以PALSC、LSC,PA∑Pn以及∑Pn为评价光合生产能力的指标,将不同品种海拔组合进行聚类分析,可以分成3类:较强型,包括云烟87-1300 m,中烟103-1300 m;中等型,包括K326-900 m,K326-1300 m,云烟87-900 m,中烟103-900 m;较低型,包括K326-500 m,云烟87-500 m,中烟103-500 m。(6)海拔和品种对多数产、质量指标有着显著的影响。海拔和品种间的互作对烤烟各产、质量指标有着显著的影响。以单叶重和群体产量作为评定烤烟产量的评价指标,将不同品种海拔组合进行聚类分析,可以分成3类:即高产型,包括中烟103-1300 m,中烟103-900 m以及云烟87-1300 m;中产型,包括中烟103-500 m,云烟87-900 m,云烟87-500 m,K326-900 m以及K326-1300 m,低产型,为K326-500 m。以产值和均价作为烟农收益的评价指标,将不同品种海拔组合进行聚类分析,可以分成3类:即高收益型,包括K326-900 m,云烟87-500 m,云烟87-900 m,云烟87-1300 m,中烟103-1300 m;中收益型,包括K326-1300 m,中烟103-900 m;低收益型,包括K326-500 m,中烟103-500 m。以评吸质量和外观质量作为烟叶质量的评价指标,将不同品种海拔组合进行聚类分析,也可以分成3类:质量较好型,包括K326-900 m和云烟87-900 m,质量中等型,包括K326-1300 m,云烟87-500 m,云烟87-1300 m,中烟103-1300 m,中烟103-900 m;质量较差型,包括K326-500 m和中烟103-500 m。(7)以河南平顶山、云南曲靖、贵州毕节、吉林敦化以及重庆地区这5个具有不同香型风格烤烟C3F等级为参比对象,将恩施州不同品种海拔组合的烤烟进行聚类和判别分析。聚类分析结果表明生态环境是决定烤烟香气成分的主要因素,不仅在大尺度生态条件下(地理位置跨度较大的不同省份或地区)烤烟香型风格存在较大差异,而且在同一区域因海拔梯度形成的生态环境差异会导致烟叶风格的改变。判别分析表明,有3个样本归为重庆中间偏浓香型一类,包括K326-1300m、中烟103-900m、中烟103-1300m;有3个样本归为河南浓香型一类,包括K326-900m、云烟87-500m、中烟103-500m;有2个样本归为云南清香型一类,包括云烟87-900m和云烟87-1300m;1个样本归为吉林中间香型一类,为K326-500m。(8)总体看来,在恩施州烟区,海拔900 m是烤烟种植的最适宜区,海拔1300 m次之。云烟87的品种适应性较强,在各个海拔下均表现出适宜的光合生产能力和较好的产、质量水平。K326最适宜于海拔900 m区域,中烟103最适宜于海拔1300 m区域。

【Abstract】 Pot and field experiment were conducted at the altitude of 500 m, 900 m and 1300 m in Enshi Autonomous Prefecture of Hubei to study difference of photosynthesis, yield and quality of flue-cured tobacco K326, Yunyan87, and Zhongyan103, which genetic background is different. Changes of gas exchange parameters, photosynthetic pigment content, RuBPcase initial activity, and chloroplast ultrastructure after full expansion of middle leaf, diurnal changes of photosynthesis and relationship with environmental factors, changes of carbon and nitrogen as well as some secondary metabolites, analysis of yield, quality and aroma-style, and relations between photosynthetic production capacity and yield and quality were studied in this paper. The main conclusions were as follows:(1) Photosynthetic function declined after full expansion of leaf in flue-cured tobacco. With the increase of altitude, photosynthetic function decline was delayed. The decline rate of Pn, photosynthetic pigment content, and RuBPcase initial activity were reduced, disintegration time of chloroplast ultrastructure and thylakoid lamellar structure were postponed, active photosynthesis duration (APD)was prolonged, leaf Source Capacity (LSC) and per area leaf source capacity (PALSC) were increased. Pathway analysis showed that the direct action of APD affecting LSC and PALSC was largest, therefore, prolonging APD was the main measure of increasing photosynthetic production capacity of leaf in flue-cured tobacco.(2)With the increase of altitude, daily photosynthesis(∑Pn), per area daily photosynthesis(PA∑Pn), and water use efficiency (WUE) were increased. At altiude of 900 m, flue-cured tobacco’s leaves had the highest Daily transpiration(∑Tr) and per area(PA∑Tr), followed by altiute of 500 m. In the sunny noon, flue-cured tobacco’s leaves appeared photosynthetic inhibition, and the degree was lightened with the increase of altitude. High temperature was the main limitied factor affecting photosynthesis and it could exacerbate photosynthetic inhibition. The main physiological factor affecting diurnal change of photosynthes was stomatal conductance(Gs), and ecological factors were different with altitude variation. At altitude of 500 m, ambient temperature (Ta) and air relative humidity (RH) were the main limitied factors to Pn. At altitude of 900 m, Pn mainly was impacted by photons flux density (PFD) . At altitude of 1300 m, Ta and RH were the main factors affecting Pn.(3) There were some differences on photosynthetic physiological characteristics among flue-cured tobacco growing at different altitudes. With the increase of altitude, adaptability and utilization ability for strong light increased, while it reduced for weak light. Flue-cured tobacco growing at high altitude had higher efficiency of carbon uptake than at low altitude. Response curves of Pn to PFD of flue-cured tobaccos’leaves at different altitudes were simulated with equation of Pn =Pnmax (1-C0exp(-aPFD/ Pn max), and equation of response curves of Pn to Ci was Pn=Pn max (1-exp(-a Ci / Pn max)-Rd.(4) Photosynthetic products mainly accumulated in the form of starch after full expansion of leaf in flue-cured tobacco. Content of starch and total carbohydrate, specific leaf weight, and ratio of carbon to nitrogen increased firstly and decreased afterwards in the process of leaf development. With the increase of altitude, the accumulating time of assimilate and dry matter were prolonged. The variation of content of starch and total carbohydrate, specific leaf weight, and ratio of carbon to nitrogen to days after full expansion were simulated with quadratic polynomial. Leaf N concentration was declined after full expansion, its variation was simulated with progressive regression model. The decline rate of leaf N concentration reduced with the increase of altitude. There were significant positive correlation between Leaf N concertration, Pn and chlorophyll content .(5) Effects of altitude on LSC, PALSC,∑Pn and PA∑Pn reached very significant level, however, only PALSC impacted by variety was significant level. The interactive effects of altitude and variety on every indexs for photosynthetic capacity reached very significant level. With PALSC, LSC,PA∑Pn and∑Pn as evaluation index for photosynthetic production capacity, different combinations of altitude and varieties were cluster analyzed, it had been divided into three categories, that were strong type including Yunyan87-1300 m and Zhongyan103-1300 m, medium type including K326-900 m, K326-1300 m and Yunyan87-900 m, lower type, including K326-500 m, Yunyan87-500 m and Zhongyan103- 500 m.(6) The effects of altitude and variety on most indexs for yield and quality production were significant. The interactive effects of altitude and variety on every indexs for yield and quality reached significant level. With weight per leaf and population yield as evaluation indexs for yield, different combinations of altitude and varieties were cluster analyzed, it had been divided into three categories, that were high-yield type including Zhongyan103-1300m, Zhongyan103-900m and Yunyan87-1300m, middle-yield type including Zhongyan103-500m, Yunyan87-900 m, Yunyan87-500m, K326-900m and K326-1300m, low-yield type, including K326-500m. With output value and average price as evaluation indexs for tobacco growers’income, different combinations of altitude and varieties were cluster analysised, it had been divided into three categories, that were high-income type including K326-900 m, Yunyan87-500m, Yunyan87-900m, Yunyan87-1300m and Zhongyan103-1300m, middle-income type including K326-1300m, Zhongyan103-900m,low-income type including K326-500m and Zhongyan103-500m. With smoking quality and appearance quality as evaluation indexs for tobacco quality, different combinations of altitude and varieties were cluster analysised, it had been divided into three categories, that were better quality type including K326-900m and Yunyan87-900m, middle quality type including K326-1300m, Yunyan87-500m, Yunyan87 -1300m, Zhongyan103-900m, Zhongyan103-1300m, lower quality type including K326-500m and Zhongyan103 -500m.(7) With grade C3F of flue-cured tobacco leaves as reference materials produced at different regions including Pingdingshan area in Henan, Qujing area in Yunnan, Bijie area in Guizhou, Dunhua area in Jilin, and Chongqing area which have different aroma type style, different combinations of altitude and varieties were cluster and discrimination analysised. Cluster analysis indicated that aroma type style showed great difference not only in large scale regions but also at different altitudes in the same region. Discrimination analysis indicated that three sample were classified as neutral tend to full aroma type in Chongqing including K326-1300 m, Zhongyan103-900 m, Zhongyan103-1300 m, three sample were classified as full aroma type in Henan including K326-900m, Yunyan87-500m, Zhongyan103 -500 m, two sample were classified as faint aroma type in Yunnan including Yunyan87-900m and Yunyan87 -1300m, one sample were classified as neutral aroma type in Jilin which was K326-500m.(8) Overall, altitude of 900 m in Enshi was the most suitable area for tobacco cultivation, followed by altitude of 1300m. Flue-cured Yunyan87 had stronger adaptability, exhibited more suitable photosynthetic production capacity and better yield and quality at every altitude. K326 was most appropriate at altitude of 900 m, while Zhongyan103 was most appropriate at altitude of 1300 m.

【关键词】 烤烟海拔光合作用产量质量恩施州
【Key words】 Flue-cured tobaccoAltitudePhotosynthesisYieldQualityEnshi prefecture
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