矮化中间砧与乔砧‘寒富’苹果幼树碳素营养特征研究

发布时间：2018-06-13 08:55

  本文选题：寒富苹果 + 矮化中间砧　； 参考：《沈阳农业大学》2016年博士论文

【摘要】：矮化密植是世界苹果栽培的趋势和标志,应用矮化中间砧在气候冷凉和多风、土壤瘠薄、高海拔地区建立高密苹果园已经成为世界上许多国家一种新的苹果栽培模式。有关矮化中间砧是否会阻滞光合同化物向地下部的运输在一定程度上还存在着争议。为此,本研究以沈阳农业大学自主选育的‘寒富’苹果为试材,选取‘寒富/山荆子’和‘寒富/GM256/山荆子’两种嫁接方式的苹果幼树,利用稳定同位素13C示踪技术和光合气体交换参数及叶绿素荧光参数分析技术,并将多种生理生化测试分析方法有机结合,深入研究矮化中间砧苹果幼树的光合特性和光合同化物运转与分配规律,由此揭示并阐明矮化中间砧苹果树光合生理基础、碳素同化物运转与分配规律及可能的调控机制,以期为‘寒富’苹果幼树矮化栽培技术措施的制定提供理论依据。研究结果如下：1.GM256作为矮化中间砧加快了接穗品种‘寒富’苹果叶片的生长发育进程,与乔砧树相比,矮化中间砧树的叶面积和比叶重增加的速度更快、成熟度更高,较高的叶绿素a(Chl a)含量和叶绿素a/叶绿素b (Chl a/b)值提高了叶片中参与光合作用被能量激发的分子数和光能转换为电能的效率。在相同的光强和CO2浓度条件下,矮砧树较乔砧树有更高的净光合速率(Pn)和固定CO2能力。2.在叶片建造期间,‘寒富’苹果叶片的转色末期为转折期,自此时期开始矮化中间砧树叶片的净光合速率和光系统活性、光能转化效率等显著提升,并较乔砧树具有更强的PS Ⅱ光化学效率(Fv/Fm)、潜在活性(Fv/F0)和光化学综合性能指数(PIABS),能将所吸收的光能更有效地转化为化学能。3.从叶片形态、质量、光合色素含量、光系统活性和光能转化效率等方面探明了矮化中间砧与乔砧‘寒富’苹果幼树对弱光和水分胁迫的反应机制。与乔砧树相比,矮化中间砧树对弱光具有更好的适应性,表现出叶面积增大、Chl b含量升高,比叶重和Chla/b值降低,进而提高了PS Ⅱ系统Fv/Fm、Fv/F0和PIABS。在受到水分胁迫时,乔砧‘寒富’苹果幼树较矮化中间砧幼树叶片中Chl a/b值变化幅度小,具有更低的初始荧光(F0)和更高的Fv/Fm和PIABS,对干旱和淹水胁迫表现出更强的抗性。4.矮化中间砧与乔砧‘寒富’苹果幼树叶片的比叶重、光合色素含量和光合生理参数等特性在不同季节存在差异。比叶重在年动态变化中呈单峰曲线变化,在养分回流期之前中间砧树的比叶重大于乔砧树,至落叶前小于乔砧树；Chla、Chlb和Chl(a+b)含量呈双峰曲线变化,两种嫁接方式的苹果叶片的Chlb含量差异不显著,但在养分回流期之前矮化中间砧树叶片较乔砧树有着更高的Chla、Chl(a+b)含量和Chla/b比值,较高的Chla含量是引起Chl(a+b)含量和Chla/b值升高进而导致中间砧树有着更强光合能力的主要原因之一。5.矮化中间砧与乔砧‘寒富’苹果幼树在春季和秋季的Pn日变化呈单峰状,最高峰在11：00左右,在夏季呈现典型的中午降低型双峰曲线,最高峰在11：00左右,次高峰在15：00左右。在养分回流期之前矮化中间砧‘寒富’苹果幼树叶片对强光和弱光的利用能力以及固定CO2的能力强于乔砧树,因而有着更强的光合能力。在秋季矮化中间砧树较乔砧树提早进入养分回流期,其对光能的利用效率以及固定CO2的能力也变为低于乔砧树。6.矮化中间砧与乔砧‘寒富’苹果幼树叶片的叶绿素荧光特性同样随季节的变化而变化,F0、Fm和Fv周年变化呈“勺”型变化,PIABs呈双峰曲线变化,而Fv/Fm受环境的影响较小。在叶片建造完成时,中间砧树叶片的F0、Fm、Fv和PIABS高于乔砧树,进一步证明了矮化中间砧可以提高叶片建造过程期间光系统的光化学能力。夏季强光和高温引发光抑制保护机制,矮砧树对强光反应更为敏感,能及早启动消耗过剩光能的保护性反应来保护光合机构免遭强光的破坏,表现出对强光环境的适应性,这从光合作用过程中对光能的吸收、传递和耗散方面间接为进一步研究矮化中间砧与乔砧‘寒富’苹果幼树叶片秋季光合速率差异提供新的证据。7.13C脉冲标记显示,在新梢迅速生长期和停长期,矮化中间砧树较乔砧树可以固定更多的13C,提高了光合同化物的输出量,但并不能提高同化物的输出率。在养分回流期矮砧树叶片固定13C光合同化物能力弱于乔砧树。8.证明了矮化中间砧韧皮部运输阻力并不是主导同化物分配的主要因素,GM256作为矮化中间砧虽然积累大量13C光合同化物,但并不阻滞13C向地下部的运输,中间砧的库强和对光合同化物的竞争改变了光合同化物的分配格局,这是矮化中间砧与乔砧苹果树碳素运转与分配产生差异的主要原因。9.阐明了矮化中间砧与乔砧‘寒富’苹果幼树在不同生育阶段光合同化物的运转与分配规律。在新梢迅速生长期,乔砧树光合固定的13C同化物分配顺序依次为：新梢、主干、一年生枝、基砧、粗根、细根；矮化中间砧树依次为：新梢、中间砧、一年生枝、细根、粗根、基砧、主干。在新梢停长期,乔砧树光合固定的13C同化物分配顺序依次为：细根、主干、粗根、一年生枝、新梢、基砧；矮化中间砧树依次为：细根、中间砧、一年生枝、新梢、粗根、基砧、主干。在养分回流期,乔砧树光合固定的13C同化物分配顺序依次为：细根、粗根、主干、新梢、一年生枝、基砧；矮化中间砧树依次为：细根、粗根、一年生枝、新梢、中间砧、主干、基砧。10.与乔砧树相比,矮化中间砧树细根对碳素同化物的竞争力更强,13C同化物运输到中间砧树地下部后向细根分配增多,而向粗根分配降低。光合同化物在根系分配的这种差异是矮化中间砧与乔砧苹果树根系构型产生差异的一个重要原因。11.矮化中间砧与乔砧‘寒富’苹果幼树叶片中的总糖含量年动态变化趋势相似,整体呈现先下降再升高的趋势,矮化中间砧‘寒富’苹果幼树叶片中的总糖含量高于乔砧树叶片。在秋季之前,矮化中间砧‘寒富’苹果幼树叶片中的山梨醇、蔗糖和果糖的含量整体高于乔砧树叶片,而葡萄糖的含量整体低于乔砧树叶片。在养分回流期,矮化中间砧树叶片中的山梨醇和葡萄糖的含量高于乔砧树叶片,蔗糖和果糖的含量低于乔砧树叶片。12.矮化中间砧通过改变接穗叶片中的山梨醇脱氢酶(SDH)活性来调控山梨醇的合成量与降解速度,进而影响叶片光合作用的效率,矮化中间砧与乔砧‘寒富’苹果幼树叶片中SDH活性整体年动态变化呈降低趋势,在养分回流期之前矮化中间砧树叶片中的SDH活性高于乔砧树,在养分回流期低于乔砧树。酸性转化酶(AI)活性与总糖含量的年动态变化趋势相似,呈现先下降再升高的趋势。蔗糖合成酶(SS)和蔗糖磷酸合成酶(SPS)活性在新梢迅速生长期降低,在新梢停长期活性变化不大,在养分回流期活性再降低。
[Abstract]:Dwarfing and dense planting is the trend and symbol of apple cultivation in the world. The application of dwarf intermediate anvil in cold and windy climate, poor soil and high density apple orchard has become a new apple cultivation model in many countries in the world. There are still disputes. Therefore, this study took "Hanfu" apple, selected by Shenyang Agricultural Uinversity as a test material, to select two young apple trees of "cold rich / mountain Jingzi" and "Han Fu /GM256/ mountain Jingzi", using stable isotope 13C tracer technology and photosynthetic gas exchange parameters and chlorophyll fluorescence parameters analysis technology. A variety of physiological and biochemical testing and analysis methods were combined to study the photosynthetic characteristics and the operation and distribution of the light contract chemicals in the dwarf intermediate anvil, which revealed and clarified the physiological basis of the photosynthesis of the dwarf intermediate anvil apple tree, the regulation of the operation and distribution of carbon assimilates and the possible regulation mechanism, so as to be 'cold rich' apple. The results are as follows: 1.GM256 as a dwarf intermediate anvil accelerates the growth and development of the leaf of "Hanfu" apple. Compared with the jo anvil, the leaf area and the specific leaf weight of the dwarf anvil increase faster, the maturity is higher, and the higher chlorophyll a (Chl a). Content and chlorophyll a/ chlorophyll b (Chl a/b) value increased the number of molecules involved in photosynthesis by energy and the conversion of light energy into electrical energy. Under the same light intensity and CO2 concentration, the dwarf anvil tree had higher net photosynthetic rate (Pn) and the fixed CO2 capacity.2. in the leaf construction, 'cold rich' apple leaves during the leaf construction. In this period, the net photosynthetic rate, the activity of light system, the efficiency of light energy conversion, and the PS II photochemical efficiency (Fv/Fm), the potential activity (Fv/F0) and the photochemical comprehensive energy index (PIABS), can be more effectively converted to the light energy absorbed by the dwarf anvil. The reaction mechanism of chemical energy.3. from leaf morphology, mass, photosynthetic pigment content, light system activity and light energy conversion efficiency was explored. The response mechanism of the dwarf intermediate anvil and the young apple tree of the jo anvil to the weak light and water stress was explored. Compared with the Qiao Zhen tree, the dwarf middle anvil had better adaptability to weak light, showing the increase of leaf area, Chl The content of B increased, the specific leaf weight and the Chla/b value were reduced, and then the PS II system Fv/Fm, Fv/F0 and PIABS. were lower in the Chl a/b value in the young tree of the Apple Rootstock than that of the dwarf middle anvil, with a lower initial fluorescence (F0) and a higher Fv/Fm and PIABS, which was stronger for drought and flooding stress. The specific leaf weight, photosynthetic pigment content and photosynthetic physiological parameters of the young tree leaves of the resistant.4. dwarf middle anvil and the jo anvil 'cold rich' apple were different in different seasons. The specific leaf weight was single peak curve in the annual dynamic change. The specific leaf weight of the middle anvil tree was larger than the jo anvil before the nutrient reflux period, and before the deciduous leaf was smaller than the jo anvil tree. The content of Chla, Chlb and Chl (a+b) changes in Shuangfeng curve. The difference of Chlb content in the leaves of the two grafting methods is not significant, but the leaves of the dwarf middle anvil have higher Chla, Chl (a+b) content and Chla/b ratio before the nutrient reflux period, and the higher Chla content is caused by Chl (a+b) content and higher value. One of the main reasons for the better photosynthetic capacity of the middle anvil is one of the.5. dwarf intermediate anvil and the young tree of the apple anvil 'cold rich' apple, which has a single peak in the spring and autumn, and the peak is about 11:00 in the spring and autumn. In the summer, it presents a typical noon decreasing Shuangfeng curve. The peak is at the left right at 11:00 and the secondary peak is around. Before the period, the leaves of the dwarf middle anvil 'Hanfu' apple tree had stronger ability to use the strong light and weak light and stronger ability of fixing CO2 than the arbor tree, and thus had stronger photosynthetic capacity. In the fall, the middle anvil tree entered the nutrient reflux period earlier than the jo anvil tree, and its utilization efficiency of light energy and the ability to fix the CO2 became lower than that of Joe. The chlorophyll fluorescence characteristics of the leaves of the anvil.6. dwarf middle anvil and the jo anvil 'cold rich' Apple also changed with the seasons, and the annual changes of F0, Fm and Fv showed a "spoon" change, and the PIABs showed a change in the Shuangfeng curve, while Fv/Fm was less affected by the environment. The F0, Fm, Fv and PIABS of the middle anvil leaves were higher than the jo anvil when the leaves were completed. The tree, further proved that the dwarf intermediate anvil can improve the photochemical ability of the optical system during the process of leaf construction. The summer strong light and high temperature luminescence inhibit the protection mechanism, the dwarf anvil tree is more sensitive to the strong light response, and can start the protective reaction that consumes the excess light energy early to protect the photosynthetic mechanism from the destruction of the strong light, showing the strong light. The adaptation of the environment to the absorption, transfer and dissipation of light energy in the process of photosynthesis provides a new evidence for further study on the difference of the photosynthetic rate of the leaves of the young apple leaves of the dwarf intermediate anvil and the jo anvil of the Apple Rootstock in autumn. The.7.13C pulse indicates the rapid growth of the new shoots and the long term and the long term, and the dwarf intermediate anvil tree is more than the jo anvil tree. More 13C can be fixed to increase the output of the light contract, but it can not increase the output of the assimilates. The ability to immobilizing the 13C light contract on the dwarf anvil leaves in the nutrient reflux period is weaker than the.8. of the rootstock tree, which proves that the transport resistance of the phloem in the dwarf intermediate anvil is not the main factor for the distribution of the dominant assimilates, and GM256 is used as a dwarf intermediate. Although the anvil accumulates a large number of 13C light contracts, it does not block the transport of 13C to the lower part of the ground. The strength of the middle anvil and the competition for the light contract change the distribution pattern of the light contract, which is the main reason for the difference between the operation and distribution of carbon in the dwarf intermediate anvil and the apple anvil tree,.9. clarifies the dwarf intermediate anvil and the Joe stock. In the rapid growth period of the new shoots, the distribution sequence of 13C assimilates in the fast growing period of the new shoots is the new shoot, the trunk, the annual branch, the base anvil, the coarse root, and the fine root. The dwarf middle anvil tree is the new shoot, the middle anvil, the annual branch, the fine root, the coarse root, the base anvil, and the main root. In the long term, the distribution sequence of 13C assimilates fixed in the tree anvil tree is fine root, trunk, coarse root, annual branch, new shoot, base anvil, and the dwarf middle anvil tree is followed by fine root, middle anvil, annual branch, new shoot, coarse root, base anvil and main stem. In the nutrient reflux period, the distribution order of 13C assimilates fixed by Joe anvil tree is in turn. The fine root, the trunk, the new shoot, the annual branch, the base anvil, and the dwarf intermediate anvil are the fine roots, the coarse roots, the annual branches, the new shoots, the middle anvil, the base anvil.10., and the fine roots of the dwarf middle anvil are more competitive to the carbon assimilates, and the 13C assimilates are transported to the subsurface of the middle anvil to increase, and to be distributed to the fine roots. The difference in the root distribution of the light contracted objects in the root system is an important reason for the difference of the root system configuration between the dwarf intermediate anvil and the apple anvil. The trend of the annual dynamic change of the total sugar content in the leaves of the young apple tree of the apple anvil and the "cold rich 'apple" is similar, and the whole body presents a trend of decreasing and then rising first, and in the dwarf.11. The total sugar content in the leaves of the young apple of the anvil was higher than that of the jo anvil leaves. Before autumn, the content of sorbitol, sucrose and fructose in the leaves of the dwarf middle anvil 'Hanfu' Apple was higher than that of the rootstock leaves, but the content of glucose was lower than that of the jo anvil leaves. The content of sorbitol and glucose is higher than that of the leaves of the rootstock. The content of sucrose and fructose is lower than that of the.12. dwarf anvil of the leaves of the rootstock. By changing the activity of sorbitol dehydrogenase (SDH) in the scion leaves, the synthesis and degradation rate of sorbitol are regulated and the efficiency of photosynthesis of the leaves is affected. The dynamic changes of SDH activity in the leaves of the young apple trees decreased. The SDH activity in the leaves of the dwarf intermediate anvil was higher than that of the jo anvil before the nutrient reflux period. The activity of the acid invertase (AI) was similar to the annual dynamic change of the total sugar content in the nutrient reflux period. The activity of synthetase (SS) and sucrose phosphate synthase (SPS) decreased during the rapid growth stage of new shoots, and their activities in the new shoots stopped little.
【学位授予单位】：沈阳农业大学
【学位级别】：博士
【学位授予年份】：2016
【分类号】：S661.1
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本文编号：2013450