發(fā)布時間：2018-06-13 08:55

  本文選題：寒富蘋果 + 矮化中間砧��； 參考：《沈陽農(nóng)業(yè)大學》2016年博士論文

【摘要】：矮化密植是世界蘋果栽培的趨勢和標志,應用矮化中間砧在氣候冷涼和多風、土壤瘠薄、高海拔地區(qū)建立高密蘋果園已經(jīng)成為世界上許多國家一種新的蘋果栽培模式。有關(guān)矮化中間砧是否會阻滯光合同化物向地下部的運輸在一定程度上還存在著爭議。為此,本研究以沈陽農(nóng)業(yè)大學自主選育的‘寒富’蘋果為試材,選取‘寒富/山荊子’和‘寒富/GM256/山荊子’兩種嫁接方式的蘋果幼樹,利用穩(wěn)定同位素13C示蹤技術(shù)和光合氣體交換參數(shù)及葉綠素熒光參數(shù)分析技術(shù),并將多種生理生化測試分析方法有機結(jié)合,深入研究矮化中間砧蘋果幼樹的光合特性和光合同化物運轉(zhuǎn)與分配規(guī)律,由此揭示并闡明矮化中間砧蘋果樹光合生理基礎、碳素同化物運轉(zhuǎn)與分配規(guī)律及可能的調(diào)控機制,以期為‘寒富’蘋果幼樹矮化栽培技術(shù)措施的制定提供理論依據(jù)。研究結(jié)果如下：1.GM256作為矮化中間砧加快了接穗品種‘寒富’蘋果葉片的生長發(fā)育進程,與喬砧樹相比,矮化中間砧樹的葉面積和比葉重增加的速度更快、成熟度更高,較高的葉綠素a(Chl a)含量和葉綠素a/葉綠素b (Chl a/b)值提高了葉片中參與光合作用被能量激發(fā)的分子數(shù)和光能轉(zhuǎn)換為電能的效率。在相同的光強和CO2濃度條件下,矮砧樹較喬砧樹有更高的凈光合速率(Pn)和固定CO2能力。2.在葉片建造期間,‘寒富’蘋果葉片的轉(zhuǎn)色末期為轉(zhuǎn)折期,自此時期開始矮化中間砧樹葉片的凈光合速率和光系統(tǒng)活性、光能轉(zhuǎn)化效率等顯著提升,并較喬砧樹具有更強的PS Ⅱ光化學效率(Fv/Fm)、潛在活性(Fv/F0)和光化學綜合性能指數(shù)(PIABS),能將所吸收的光能更有效地轉(zhuǎn)化為化學能。3.從葉片形態(tài)、質(zhì)量、光合色素含量、光系統(tǒng)活性和光能轉(zhuǎn)化效率等方面探明了矮化中間砧與喬砧‘寒富’蘋果幼樹對弱光和水分脅迫的反應機制。與喬砧樹相比,矮化中間砧樹對弱光具有更好的適應性,表現(xiàn)出葉面積增大、Chl b含量升高,比葉重和Chla/b值降低,進而提高了PS Ⅱ系統(tǒng)Fv/Fm、Fv/F0和PIABS。在受到水分脅迫時,喬砧‘寒富’蘋果幼樹較矮化中間砧幼樹葉片中Chl a/b值變化幅度小,具有更低的初始熒光(F0)和更高的Fv/Fm和PIABS,對干旱和淹水脅迫表現(xiàn)出更強的抗性。4.矮化中間砧與喬砧‘寒富’蘋果幼樹葉片的比葉重、光合色素含量和光合生理參數(shù)等特性在不同季節(jié)存在差異。比葉重在年動態(tài)變化中呈單峰曲線變化,在養(yǎng)分回流期之前中間砧樹的比葉重大于喬砧樹,至落葉前小于喬砧樹；Chla、Chlb和Chl(a+b)含量呈雙峰曲線變化,兩種嫁接方式的蘋果葉片的Chlb含量差異不顯著,但在養(yǎng)分回流期之前矮化中間砧樹葉片較喬砧樹有著更高的Chla、Chl(a+b)含量和Chla/b比值,較高的Chla含量是引起Chl(a+b)含量和Chla/b值升高進而導致中間砧樹有著更強光合能力的主要原因之一。5.矮化中間砧與喬砧‘寒富’蘋果幼樹在春季和秋季的Pn日變化呈單峰狀,最高峰在11：00左右,在夏季呈現(xiàn)典型的中午降低型雙峰曲線,最高峰在11：00左右,次高峰在15：00左右。在養(yǎng)分回流期之前矮化中間砧‘寒富’蘋果幼樹葉片對強光和弱光的利用能力以及固定CO2的能力強于喬砧樹,因而有著更強的光合能力。在秋季矮化中間砧樹較喬砧樹提早進入養(yǎng)分回流期,其對光能的利用效率以及固定CO2的能力也變?yōu)榈陀趩陶铇洹?.矮化中間砧與喬砧‘寒富’蘋果幼樹葉片的葉綠素熒光特性同樣隨季節(jié)的變化而變化,F0、Fm和Fv周年變化呈“勺”型變化,PIABs呈雙峰曲線變化,而Fv/Fm受環(huán)境的影響較小。在葉片建造完成時,中間砧樹葉片的F0、Fm、Fv和PIABS高于喬砧樹,進一步證明了矮化中間砧可以提高葉片建造過程期間光系統(tǒng)的光化學能力。夏季強光和高溫引發(fā)光抑制保護機制,矮砧樹對強光反應更為敏感,能及早啟動消耗過剩光能的保護性反應來保護光合機構(gòu)免遭強光的破壞,表現(xiàn)出對強光環(huán)境的適應性,這從光合作用過程中對光能的吸收、傳遞和耗散方面間接為進一步研究矮化中間砧與喬砧‘寒富’蘋果幼樹葉片秋季光合速率差異提供新的證據(jù)。7.13C脈沖標記顯示,在新梢迅速生長期和停長期,矮化中間砧樹較喬砧樹可以固定更多的13C,提高了光合同化物的輸出量,但并不能提高同化物的輸出率。在養(yǎng)分回流期矮砧樹葉片固定13C光合同化物能力弱于喬砧樹。8.證明了矮化中間砧韌皮部運輸阻力并不是主導同化物分配的主要因素,GM256作為矮化中間砧雖然積累大量13C光合同化物,但并不阻滯13C向地下部的運輸,中間砧的庫強和對光合同化物的競爭改變了光合同化物的分配格局,這是矮化中間砧與喬砧蘋果樹碳素運轉(zhuǎn)與分配產(chǎn)生差異的主要原因。9.闡明了矮化中間砧與喬砧‘寒富’蘋果幼樹在不同生育階段光合同化物的運轉(zhuǎn)與分配規(guī)律。在新梢迅速生長期,喬砧樹光合固定的13C同化物分配順序依次為：新梢、主干、一年生枝、基砧、粗根、細根；矮化中間砧樹依次為：新梢、中間砧、一年生枝、細根、粗根、基砧、主干。在新梢停長期,喬砧樹光合固定的13C同化物分配順序依次為：細根、主干、粗根、一年生枝、新梢、基砧；矮化中間砧樹依次為：細根、中間砧、一年生枝、新梢、粗根、基砧、主干。在養(yǎng)分回流期,喬砧樹光合固定的13C同化物分配順序依次為：細根、粗根、主干、新梢、一年生枝、基砧；矮化中間砧樹依次為：細根、粗根、一年生枝、新梢、中間砧、主干、基砧。10.與喬砧樹相比,矮化中間砧樹細根對碳素同化物的競爭力更強,13C同化物運輸?shù)街虚g砧樹地下部后向細根分配增多,而向粗根分配降低。光合同化物在根系分配的這種差異是矮化中間砧與喬砧蘋果樹根系構(gòu)型產(chǎn)生差異的一個重要原因。11.矮化中間砧與喬砧‘寒富’蘋果幼樹葉片中的總糖含量年動態(tài)變化趨勢相似,整體呈現(xiàn)先下降再升高的趨勢,矮化中間砧‘寒富’蘋果幼樹葉片中的總糖含量高于喬砧樹葉片。在秋季之前,矮化中間砧‘寒富’蘋果幼樹葉片中的山梨醇、蔗糖和果糖的含量整體高于喬砧樹葉片,而葡萄糖的含量整體低于喬砧樹葉片。在養(yǎng)分回流期,矮化中間砧樹葉片中的山梨醇和葡萄糖的含量高于喬砧樹葉片,蔗糖和果糖的含量低于喬砧樹葉片。12.矮化中間砧通過改變接穗葉片中的山梨醇脫氫酶(SDH)活性來調(diào)控山梨醇的合成量與降解速度,進而影響葉片光合作用的效率,矮化中間砧與喬砧‘寒富’蘋果幼樹葉片中SDH活性整體年動態(tài)變化呈降低趨勢,在養(yǎng)分回流期之前矮化中間砧樹葉片中的SDH活性高于喬砧樹,在養(yǎng)分回流期低于喬砧樹。酸性轉(zhuǎn)化酶(AI)活性與總糖含量的年動態(tài)變化趨勢相似,呈現(xiàn)先下降再升高的趨勢。蔗糖合成酶(SS)和蔗糖磷酸合成酶(SPS)活性在新梢迅速生長期降低,在新梢停長期活性變化不大,在養(yǎng)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.
【學位授予單位】：沈陽農(nóng)業(yè)大學
【學位級別】：博士
【學位授予年份】：2016
【分類號】：S661.1
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本文編號：2013450