【作者】 李晶；

【导师】 姜远茂；

【作者基本信息】 山东农业大学 ， 果树学， 2013， 博士

【摘要】 矮砧集约高效栽培是苹果栽培的发展方向，目前我国在矮砧利用上主要是矮化中间砧。生产上由于栽培技术研究滞后，中间砧苹果旺长与早衰现象共存，如何既保证稳健生长又及时成花结果是栽培管理的核心，这就需要明确中间砧苹果C、N营养特性。为此，我们利用稳定性同位素¹⁵N和¹³C双标记技术，研究了SH₁₈、SH₂₈、SH₃₈、SHc、CG₂₄和Mark等6个中间砧对苹果生长发育及碳氮营养的影响，以生产上最常用的富士/M₂₆/平邑甜茶为试材研究不同氮素水平下年周期碳氮营养利用特性，以宫藤富士/SH₁/八棱海棠、宫藤富士/SH₆/平邑甜茶等为试材研究了摘心和摘老叶等栽培措施对苹果生长发育及碳氮营养的影响，以期为中间砧苹果优质丰产栽培技术制定提供依据。主要结果如下：1.不同中间砧苹果幼树在不同氮水平下的碳氮特性研究：根据生物量进行聚类分析可把供试中间砧分为3类：对生长抑制作用最强的有CG₂₄，居中的有SH₁₈、SH_c、SH₃₈、Mark，抑制最弱的是SH₂₈。不同类型中间砧幼树在不同施氮水平下树体生物量差异显著，在不施氮肥（N0）、适宜施氮肥（N100）和过量施氮肥（N200）三个氮素水平下，对树体生长抑制最弱的SH₂₈中间砧苹果幼树随着施氮量增加树体生物量显著增加，而其他中间砧苹果幼树表现出在适宜供氮条件下生物量最大，继续增加氮素供应反而抑制生长，表明单纯通过增加氮肥用量不能促进抑制作用较强中间砧苹果的生长，需要在适应供氮条件下才能促进生长。不同中间砧在不同供氮水平下对¹⁵N的利用及其分配差异显著，对树体生长抑制最弱的SH₂₈中间砧苹果幼树随着施氮量增加¹⁵N利用率逐渐升高，¹⁵N更多分配到地上部，根系的¹⁵N分配率逐渐减少；而CG₂₄和SH₃₈中间砧苹果幼树在适宜供氮条件下氮素利用率最高，但¹⁵N分配率存在显著差异；CG₂₄在过量施氮条件下¹⁵N更多分配到根系，地上部的¹⁵N分配率逐渐减少，不施氮肥和适宜施氮条件下更多分配到地上部，根系¹⁵N分配率减少；SH₃₈在适宜施氮条件下¹⁵N更多的分配到根系，地上部的¹⁵N分配率减少，而不施氮肥和过量施氮条件下更多的分配到地上部，根系¹⁵N分配率减少。不同中间砧在不同施氮水平下苹果幼树地上部和根系的¹³C分配率差异显著，对树体生长抑制最弱的SH₂₈中间砧苹果幼树在过量施氮肥条件下¹³C更多分配到地上部，根系¹³C分配减少，而不施氮肥和适宜施氮肥条件下¹³C更多分配到根系，地上部¹³C分配率减少；SH₃₈中间砧苹果幼树随着施氮量增加¹³C更多分配到地上部，根系¹³C分配逐渐减少；而CG₂₄在适宜施氮肥条件下¹³C更多分配到地上部(根系¹³C分配率最低17.63%)，不施氮肥条件下其次，过量施氮肥条件下最低(根系¹³C分配率最高27.14%)。2.烟富3/M₂₆/平邑甜茶幼树年周期碳氮特性研究：从秋梢旺长期不同氮素水平下烟富3/M₂₆/平邑甜茶幼树生物量来看，M₂₆中间砧为抑制作用较强的类型。年周期中其对不同氮素水平的反应，在春梢旺长期三者间差异不显著，随着施氮量增加生物量有增加趋势；到春梢停长期适宜施氮肥和过量施氮肥树体生物量显著高于不施氮肥，仍然呈随着施氮肥量增加生物量增加趋势；而在秋梢旺长期适宜施氮肥最高。适宜施氮肥处理较过量施氮肥处理在秋梢旺长期生物量发生改变与春梢停长期适宜施氮肥处理树体细根显著高于过量施氮肥处理有关。在年周期不同氮素水平对¹⁵N的利用及其分配影响方面，虽然最终表现为适宜施氮肥处理¹⁵N的利用率最高，但在春梢旺长期是以不施氮肥处理最高，到春梢停长期开始以适宜施氮肥最高。各施氮肥处理均表现为随着生长期的延长，根系¹⁵N分配率逐渐减少，地上部¹⁵N分配率逐渐增加。施氮对¹³C分配影响显著，随着生长期的延长，不施氮肥树体根系¹³C分配率逐渐增高，地上部¹³C分配率逐渐减少；适宜施氮肥和过量施氮肥初期促进根系¹³C分配率显著升高，但随着生长期的延长逐渐减少，地上部¹³C分配率逐渐增高。适宜施氮利于维持均衡的碳氮代谢，使碳氮同化产物在地上和地下部分配均衡，对翌年树体生长的促进作用较当年更为显著。淀粉累积量与果树成花显著相关，本试验整个生长期内，叶片淀粉含量均表现为不施氮肥条件下最高，同时不施氮肥条件下叶片ABA含量也表现为最高。因此不同氮水平下树体光合产物的分配与贮藏形态的差异是由ABA的合成差异所造成。3.摘心对不同中间砧苹果碳氮特性影响的研究：在4年生宫藤富士/SH1/八棱海棠大树和1年生宫藤富士/SH6/平邑甜茶幼树结果均表明，与对照相比摘心可显著抑制树体长势，氮吸收量和碳固定量显著减少；摘心后¹³C和¹⁵N向贮藏器官分配比率显著升高，而新生器官（叶片、新梢和细根）的比率显著降低；叶片IAA含量显著降低，而ABA含量显著上升。4.摘老叶对中间砧苹果幼树碳氮特性影响的研究：以1年生宫藤富士/SH6/平邑甜茶幼树为试材进行摘老叶试验，结果表明，与对照相比摘除中心干下部老叶显著促进中心干生长，碳固定量显著下降，氮吸收量差异不显著；¹³C和¹⁵N向当年新生器官（叶片和新梢）分配比率显著升高，而贮藏器官的比率显著降低；叶片ABA含量逐渐降低，而GA3含量逐渐上升。更多还原

【Abstract】 Dwarf rootstock efficient intensive cultivation is a new developing trend of appleindustry with the dwarfing interstocks using the most widely. Since the backward oncultivation technique study and the vigorous growth and premature senescence of theinterstock making the robust growth of interstocks consistent with timely flowering andbearing is the heart of management, which needs to study deep into the nutritioncharacteristics on carbon and nitrogen of interstocks. Therefore, double labelling technique of¹⁵N and¹³C stable isotopes was used to study the effect on growth and development and thecarbon and nitrogen nutrition of SH₁₈、SH₂₈、SH₃₈、SHc、CG₂₄and Mark, the yearly C、Nutilization characteristics of different nitrogen levels of commonly-used Fuji/M₂₆/M.hupehensis Rehd.in production, and effects of pinching and old leaf picking on growth andcarbon and nitrogen nutrition of Borkh cv. Fuji/SH1/Malus robusta Rehd. and Borkh cv.Fuji/SH6/M. hupehensis Rehd., in order to provide evidence for high quality and yieldcultivation technique of trees with interstocks. The main results were as follows:1.C、N characteristics of different nitrogen levels on young apple trees with differentinterstocks: the testing interstocks were classified into three different types based onclustering analysis of biomass: CG₂₄has the strongest effect on growth inhibition, while SH₂₈the weakest, and SH₁₈、SHc、SH₃₈and Mark showed medium inhibition effect. Significantdifferences can be found on biomass of different nitrogen levels on young apple trees, of theno nitrogen （N0）、proper nitrogen（N100）and over-dose nitrogen（N200）treatment, biomassof SH₂₈increased with the increasing application of nitrogen, while biomass of the other treesshowed the best results with the proper amount of nitrogen application, and continuousapplication of nitrogen existed inhibition effect, indicating that proper amount of nitrogen isessential for the promotion of young apple tree with strongly inhibition interstocks.Significant differences can be seen on¹⁵N utilization and distribution of various nitrogenapplication,¹⁵N utilization rate of trees with interstock SH₂₈increased with the increasingapplication of nitrogen,and the¹⁵N is mostly distributed into the aboveground, while the root¹⁵N distribution rate decreases; however, trees with interstock CG₂₄and SH₃₈shows thehighest¹⁵N utilization rate with the treatment of proper amount of nitrogen; and¹⁵Ndistribution differs significantly:¹⁵N is mostly distributed to the root of CG₂₄with over-doseapplication of nitrogen, while the aboveground¹⁵N distribution rate decreased, the¹⁵N distribution into aboveground of CG24was superior to root system in the treatment of nonitrogen application and proper nitrogen application;¹⁵N of trees with interstock SH₃₈wasmostly distributed to root system with proper nitrogen application, and the¹⁵N distributionrate of aboveground was superior to root system in the treatment of no nitrogen applicationand over-dose application of nitrogen.¹³C distribution rate of different nitrogen levels on treeswith various interstocks differs significantly:¹³C was mostly distributed to aboveground intrees with interstock SH₂₈with the treatment of over-dose nitrogen application, while¹³Cdistribution rate of the root system decreased;¹³C distribution rate of aboveground wassuperior to root system with the treatment of no nitrogen application and proper nitrogenapplication;¹³C was mostly distributed to aboveground in trees with interstock SH₃₈with theincreasing application amount of nitrogen;¹³C distribution of different nitrogen levels on treeswith different interstocks differs significantly,¹³C was mostly distributed to aboveground intrees with interstock SH₂₈under the treatment of over-dose nitrogen application, and¹³C of theroot system decreased, and¹³C distribution into aboveground was superior to root system inthe treatment of no nitrogen application and proper nitrogen application;¹³C was mostlydistributed to aboveground with interstock SH₃₈under the increasing application amount ofnitrogen, and the¹³C distribution rate of rootstock decreased; in trees with interstock CG24,¹³C is mostly distributed to the aboveground under proper nitrogen application (¹³Cdistribution rate of root being the lowest17.63%), and then no nitrogen treatment,while theover-dose application of nitrogen showed the lowest¹³C distribution rate(¹³C distribution rateof root being the highest27.14%).2. Study on annual cycle of carbon and nitrogen characteristics of Yantai Fuji3/M₂₆/M.hupehensis Rehd.: Biomass during autumn shoots growth period under different nitrogenlevels showed that M₂₆interstock is the type of strong inhibition. According to their annualcycle effects under different nitrogen levels, no obvious difference were found above thosethree treatments during the stage of vigorous growth of spring shoots. With the increase ofnitrogen levels, the biomass also increased; at spring shoots growth arrest stage, the biomassof medium nitrogen and excess nitrogen were higher than the treatment with no nitrogen, andthere were significantly differences among those three treatments, with the increase ofnitrogen, the biomass also increased; however at autumn shoots growth period, the biomass ofmedium nitrogen was the highest. Compared to over-dose nitrogen, the biomass of mediumnitrogen has changed at autumn shoots growth period, this related to the fine root biomass ofmedium nitrogen is higher than that of excess nitrogen. About the respect of utilization anddistribution of¹⁵N, although the utilization of medium nitrogen was the highest, no nitrogen was the highest during the stage of vigorous growth of spring shoots, until to spring shootsgrowth arrest stage, medium nitrogen was the highest. With the development of growth period,every treatment showed that the distribution of¹⁵N of root was decreased, with aerial partincreased. The treatment of no nitrogen, the distribution of¹³C of root increased, while aerialpart decreased; those were contrary to the treatment of medium nitrogen and over-dosenitrogen treatment. Medium nitrogen could keep balance of C and N metabolism, madecarbon and nitrogen assimilated broaden and deepen the optimization between abovegroundand underground portion, this could promote tree growth next year. Starch content wassignificantly related to flower formation, In this study, starch content and ABA content ofleaves were the highest in the treatment of no nitrogen. Therefore the distribution and storageform of photosynthetic products were caused by ABA synthetic differences.3. C、N characteristics of pinching on apple trees with different interstocks: Study onfour-year-old Borkh cv. Fuji/SH1/M. hupehensis Rehd. and one-year-old Borkh cv.Fuji/SH6/Malus robusta Rehd. showed that pinching significantly inhibited the tree vigorcompared with control, and decreased the uptake of nitrogen and the carbon fixationsignificantly, while increased¹³C and¹⁵N distribution rate on storage organs significantly, anddecreased the ratio of new organs（leaves, shoot and fine roots） significantly; IAA contents ofleaves significantly decreased, while the ABA contents significantly increased.4. C、N characteristics of picking old leaf on young apple trees: with old leaves pickingon one-year-old Borkh cv. Fuji/SH6/Malus robusta Rehd., results showed that, picking oldleaves could significantly improve the growth of center stem compared to control, the carbonfixation significantly reduced, with no obvious difference can be seen on the uptake ofnitrogen. The¹³C and¹⁵N distribution rate on neonatal organs（leaves and shoot） significantlyincreased, with the ratio of storage organs significantly decreased; ABA contents of leavesgradually reduced, while GA3contents gradually increased.更多还原