日光温室柑橘诱导成花及落果机理研究
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摘要
本研究以枳（Poncirus trifoliate Raf.）砧砂糖橘（Citrus reticulata“Shatangju”）和枳砧南丰蜜橘（Citrus reticulata “Nanfeng”）为试材，探索了日光温室柑橘生长发育特性、低温诱导柑橘成花效应及生理响应、柑橘幼果脱落的生理机理，为日光温室柑橘栽培生产及调控提供了理论基础。主要研究结果如下：
1.日光温室条件下，柑橘春梢叶片叶绿素含量最低，硝态氮和氨基态氮含量均显著低于夏梢和秋梢。有叶果坐果率显著高于无叶果。
2.低温诱导柑橘茎段成花处理中，10/5℃（昼12h/夜12h）处理下柑橘离体茎段成花率最高，4℃（恒温24h）处理茎段没有形成花芽。15/10℃（昼12h/夜12h）处理下，诱导成花的同时也促进了茎段萌芽，随着萌芽率的升高，成花率下降。日光温室条件下，整个冬季砂糖橘和南丰秋梢成花率呈升高趋势，表现为2月平均成花率＞1月＞12月。有叶花/无叶花比值与＜5℃低温累积小时数之间呈极显著负相关。5℃～20℃低温累积小时数达到1309h时砂糖橘对应的成花率趋于稳定为40.51%，5℃～20℃低温累积小时数达到1456h时南丰对应的成花率趋于稳定为39.22%。低温对柑橘成花的诱导效应存在品种间差异。
3.4℃处理柑橘茎段C/N值逐渐降低，主要是由全碳含量降低引起的。日光温室条件下和不同低温诱导成花过程中，柑橘枝条C/N值变化和脯氨酸含量的积累对成花表现出积极响应。低温诱导下，柑橘枝条ZRs含量与成花相关，高水平含量的ZRs有利于柑橘成花。柑橘枝条的ZRs/GA、ABA/GA、（ABA+ZRs）/（IAA+GA）与成花率间表现出一定的相关性。
4.谢花后60d内是日光温室柑橘生理落果期，其中以谢花后20d~40d落果最为严重，50d后落果量显著减少，60d后趋于结束。谢花后20d~40d的生理落果期间，柑橘幼果的蔗糖代谢以转化分解为主，且以AI作用为主，果实可溶性糖和淀粉含量降低，且相邻叶片的蔗糖合成相关酶活性降低，导致蔗糖含量减少；在氮代谢方面，果实的硝酸还原酶活性偏低限制了硝态氮的还原同化，导致果实氨基态氮含量偏低，同时相邻叶片内GS和硝酸还原酶活性降低，氮代谢活动受到限制；激素代谢方面，落果期间多胺分解代谢增强，ABA含量出现积累。说明果实和叶片内的糖代谢（尤其是蔗糖和淀粉）、氮代谢以及果实的多胺代谢和ABA含量的变化，对幼果脱落产生了重要影响。
谢花后20d~40d落果期间，无叶果落果率显著高于有叶果。其原因是无叶果相邻叶片蔗糖合成受到影响，往果实运输的蔗糖减少，淀粉含量显著低于有叶果。而有叶果相邻叶片蔗糖合成能力逐渐增强，光合同化产物（蔗糖）由源（叶片）到库（幼果）运输量增多，进一步转化为淀粉贮藏起来，促进了坐果。同时无叶果幼果内硝酸还原酶活性低于有叶果，多胺分解代谢更强，ABA含量积累量更大，加剧了幼果的脱落。
5.施用有机肥和硫磺粉均能明显降低土壤pH值，混合施肥中随施硫量的增加，土壤有效硫含量升高，土壤有机质、全氮、碱解氮含量呈先下降后升高的趋势。施用有机肥、硫磺粉后南丰蜜橘叶片内可溶性蛋白质含量增加、硝酸还原酶活性增强。当施硫量不超过100mg/kg时，柑橘的枝粗和叶绿素含量显著增加，可有效缓解叶片失绿黄化问题，同时可促进柑橘幼果淀粉的积累，有利于坐果。
Open-field cultivation has been the main mode of citrus cultivation in China, withmajor blooming peak in spring and citrus fruits only shipped to markets in October throughDecember. Under protected cultivation, it is possible to force the flowering stage of citrusto an advanced date via temperature control for earlier shipments of fruits to markets. Innorth China, early autumn low temperatures in solar greenhouses have been used to induceearly citrus flower bud differentiation. Induction of flowers by the warming effect withplastic film covers or heating forced sprouting, in order that fruits ripening stage can beadvanced. Flowering time and fruit setting rate of citrus would directly affect the yield andbenefit of citrus cultivation. The examination used “Shatangju” tangerine (Citrus reticulata“Shatangju”) and “Nanfeng” tangerine （Citrus reticulata “Nanfeng”）seedlings withstock of trifoliate orange (Poncirus trifoliate Raf.) to study the biological characteristic andphysiological mechanism of flower induction by low temperature and fruitlet abscission ofcitrus under solar greenhouse, in order to offer the theoretical basis for citrus cultivation insolar greenhouse. The results were as follows:
1. Under the condition of solar greenhouse, the chlorophyll, nitrate nitrogen andamino nitrogen contents of spring leaves were the lowest. In addition, fruit setting rate ofleafy fruits was higher than leafless fruits.
2. Floral ratio of axillary buds was the highest treated with low temperature10/5℃(15℃at day for12h,10℃at night for12h), while no flower buds formed under lowtemperature4℃(at constant temperature for24h). Axillary buds was not only inducted toflower, but also was promoted germination treated with low temperature15/10℃(15℃atday for12h,10℃at night for12h). Floral ratio apparently decreased while germinationratio sharply increased. Under solar greenhouse conditions, floral ratio of “Shatangju”tangerine and “Nanfeng” tangerine apparently increased during the whole of winter, whichshowed order of average floral ratio: February> January> December. A significantlynegative correlation was noted between leafy to leafless flower ratio and accumulatedhours of <5℃. When accumulated time of low temperature5~20℃reached1309hours,floral ratio of “Shatangju” tangerine stabilized at40.51%. Floral ratio of “Nanfeng”tangerine stabilized at39.22%with low temperature5~20℃reaching1456hours. Theresults indicated there were differences in the response of different citrus variety flower to low temperature.
3. Under low temperature4℃, C/N ratio was gradually decreased, mainly owing tothe increased total C content. The response of the change in C/N ratio and proline contentof citrus was positive to flower induction. ZRs content was related with flower formationand beneficial to flower formation for citrus with high content. The results revealed thatfloral ratio of citrus was related to ZRs/GA, ABA/GA,(ABA+ZRs)/(IAA+GA) of shoots.
4. Within60days after withering of flowers was physiological fruit-drop period ofcitrus under solar greenhouse, Fruitlets abscission was the most serious during20~40daysafter withering of flowers. The number of dropping fruits significantly reduced on the50day and fruit drop tended to the end. During fruitlets largely dropped, sucrose metabolismin citrus fruitlets was mainly in catabolism, that acid invertase played a major role. Solublesugar and starch content decreaced in fruitlets. While sucrose synthase activity reduced inadjacent leaves, resulting in sucrose decreasing. At the same time, amino nitrogen contentcould be reduced by the lower activity nitrate reductase of leafy and leafless fruits, whichlimited reduction and assimilation of nitrate nitrogen. Glutamine synthetase and nitratereductase activity reduced in adjacent leaves, which affected nitrogen metabolism of leaves.Meanwhile, polyamines catabolism enhanced and ABA content accumulated in fruitlets.The results suggest that sugar metabolism (especially in sucrose and starch) and nitrogenmetabolism in fruits and leaves, polyamine metabolism and the change of abscisic acidcontent in fruits had important influence on the fruitlets abscission.
0~20days after withering of flowers, fruitlets abscission ratio of leafless fruits washigher than leafy fruits. The reason was that sucrose anabolism of the adjacent leaves ofleafless fruits was effected, which led sucrose content transferred from leaves to fruitsdecreasing. The starch content of leafless fruitlets was significantly lower than leafyfruitlets. While the capacity of sucrose anabolism of leafy bearing shoots was graduallyincreased. The supply capacity of leafy bearing shoots that photosynthate with sucrosetransported from leaf to fruit enhanced. Sucrose was converted to starch and stored in fruit,which was favorable to leafy fruitlets set. At the same time, compared with leafy fruits,nitrate reductase activity was lower, polyamine catabolism was stronger and ABA contentwas more in leafless fruits.
5. Results showed that soil pH value decreased obviously after organic fertilizer andsulfur application. Under mixed application, soil organic matter, total nitrogen andavailable nitrogen content decreased at first but later increased with the increase of amountof sulfur application，while soil available sulfur content increased gradually. Under organic fertilizer and sulfur application treatment, soluble protein content of “Nanfeng” tangerineincreased and nitrate reductase activity was promoted. When sulfur application was lessthan100mg/kg, the branch diameter and the chlorophyll content both increased undermixed application. This could effectively alleviate the problem of leaf chorosis. Undermixed application(sulfur application was100mg/kg), starch accumulation in leafy fruitswas promoted, which was beneficial to fruit set.
1.日光温室条件下，柑橘春梢叶片叶绿素含量最低，硝态氮和氨基态氮含量均显著低于夏梢和秋梢。有叶果坐果率显著高于无叶果。
2.低温诱导柑橘茎段成花处理中，10/5℃（昼12h/夜12h）处理下柑橘离体茎段成花率最高，4℃（恒温24h）处理茎段没有形成花芽。15/10℃（昼12h/夜12h）处理下，诱导成花的同时也促进了茎段萌芽，随着萌芽率的升高，成花率下降。日光温室条件下，整个冬季砂糖橘和南丰秋梢成花率呈升高趋势，表现为2月平均成花率＞1月＞12月。有叶花/无叶花比值与＜5℃低温累积小时数之间呈极显著负相关。5℃～20℃低温累积小时数达到1309h时砂糖橘对应的成花率趋于稳定为40.51%，5℃～20℃低温累积小时数达到1456h时南丰对应的成花率趋于稳定为39.22%。低温对柑橘成花的诱导效应存在品种间差异。
3.4℃处理柑橘茎段C/N值逐渐降低，主要是由全碳含量降低引起的。日光温室条件下和不同低温诱导成花过程中，柑橘枝条C/N值变化和脯氨酸含量的积累对成花表现出积极响应。低温诱导下，柑橘枝条ZRs含量与成花相关，高水平含量的ZRs有利于柑橘成花。柑橘枝条的ZRs/GA、ABA/GA、（ABA+ZRs）/（IAA+GA）与成花率间表现出一定的相关性。
4.谢花后60d内是日光温室柑橘生理落果期，其中以谢花后20d~40d落果最为严重，50d后落果量显著减少，60d后趋于结束。谢花后20d~40d的生理落果期间，柑橘幼果的蔗糖代谢以转化分解为主，且以AI作用为主，果实可溶性糖和淀粉含量降低，且相邻叶片的蔗糖合成相关酶活性降低，导致蔗糖含量减少；在氮代谢方面，果实的硝酸还原酶活性偏低限制了硝态氮的还原同化，导致果实氨基态氮含量偏低，同时相邻叶片内GS和硝酸还原酶活性降低，氮代谢活动受到限制；激素代谢方面，落果期间多胺分解代谢增强，ABA含量出现积累。说明果实和叶片内的糖代谢（尤其是蔗糖和淀粉）、氮代谢以及果实的多胺代谢和ABA含量的变化，对幼果脱落产生了重要影响。
谢花后20d~40d落果期间，无叶果落果率显著高于有叶果。其原因是无叶果相邻叶片蔗糖合成受到影响，往果实运输的蔗糖减少，淀粉含量显著低于有叶果。而有叶果相邻叶片蔗糖合成能力逐渐增强，光合同化产物（蔗糖）由源（叶片）到库（幼果）运输量增多，进一步转化为淀粉贮藏起来，促进了坐果。同时无叶果幼果内硝酸还原酶活性低于有叶果，多胺分解代谢更强，ABA含量积累量更大，加剧了幼果的脱落。
5.施用有机肥和硫磺粉均能明显降低土壤pH值，混合施肥中随施硫量的增加，土壤有效硫含量升高，土壤有机质、全氮、碱解氮含量呈先下降后升高的趋势。施用有机肥、硫磺粉后南丰蜜橘叶片内可溶性蛋白质含量增加、硝酸还原酶活性增强。当施硫量不超过100mg/kg时，柑橘的枝粗和叶绿素含量显著增加，可有效缓解叶片失绿黄化问题，同时可促进柑橘幼果淀粉的积累，有利于坐果。
Open-field cultivation has been the main mode of citrus cultivation in China, withmajor blooming peak in spring and citrus fruits only shipped to markets in October throughDecember. Under protected cultivation, it is possible to force the flowering stage of citrusto an advanced date via temperature control for earlier shipments of fruits to markets. Innorth China, early autumn low temperatures in solar greenhouses have been used to induceearly citrus flower bud differentiation. Induction of flowers by the warming effect withplastic film covers or heating forced sprouting, in order that fruits ripening stage can beadvanced. Flowering time and fruit setting rate of citrus would directly affect the yield andbenefit of citrus cultivation. The examination used “Shatangju” tangerine (Citrus reticulata“Shatangju”) and “Nanfeng” tangerine （Citrus reticulata “Nanfeng”）seedlings withstock of trifoliate orange (Poncirus trifoliate Raf.) to study the biological characteristic andphysiological mechanism of flower induction by low temperature and fruitlet abscission ofcitrus under solar greenhouse, in order to offer the theoretical basis for citrus cultivation insolar greenhouse. The results were as follows:
1. Under the condition of solar greenhouse, the chlorophyll, nitrate nitrogen andamino nitrogen contents of spring leaves were the lowest. In addition, fruit setting rate ofleafy fruits was higher than leafless fruits.
2. Floral ratio of axillary buds was the highest treated with low temperature10/5℃(15℃at day for12h,10℃at night for12h), while no flower buds formed under lowtemperature4℃(at constant temperature for24h). Axillary buds was not only inducted toflower, but also was promoted germination treated with low temperature15/10℃(15℃atday for12h,10℃at night for12h). Floral ratio apparently decreased while germinationratio sharply increased. Under solar greenhouse conditions, floral ratio of “Shatangju”tangerine and “Nanfeng” tangerine apparently increased during the whole of winter, whichshowed order of average floral ratio: February> January> December. A significantlynegative correlation was noted between leafy to leafless flower ratio and accumulatedhours of <5℃. When accumulated time of low temperature5~20℃reached1309hours,floral ratio of “Shatangju” tangerine stabilized at40.51%. Floral ratio of “Nanfeng”tangerine stabilized at39.22%with low temperature5~20℃reaching1456hours. Theresults indicated there were differences in the response of different citrus variety flower to low temperature.
3. Under low temperature4℃, C/N ratio was gradually decreased, mainly owing tothe increased total C content. The response of the change in C/N ratio and proline contentof citrus was positive to flower induction. ZRs content was related with flower formationand beneficial to flower formation for citrus with high content. The results revealed thatfloral ratio of citrus was related to ZRs/GA, ABA/GA,(ABA+ZRs)/(IAA+GA) of shoots.
4. Within60days after withering of flowers was physiological fruit-drop period ofcitrus under solar greenhouse, Fruitlets abscission was the most serious during20~40daysafter withering of flowers. The number of dropping fruits significantly reduced on the50day and fruit drop tended to the end. During fruitlets largely dropped, sucrose metabolismin citrus fruitlets was mainly in catabolism, that acid invertase played a major role. Solublesugar and starch content decreaced in fruitlets. While sucrose synthase activity reduced inadjacent leaves, resulting in sucrose decreasing. At the same time, amino nitrogen contentcould be reduced by the lower activity nitrate reductase of leafy and leafless fruits, whichlimited reduction and assimilation of nitrate nitrogen. Glutamine synthetase and nitratereductase activity reduced in adjacent leaves, which affected nitrogen metabolism of leaves.Meanwhile, polyamines catabolism enhanced and ABA content accumulated in fruitlets.The results suggest that sugar metabolism (especially in sucrose and starch) and nitrogenmetabolism in fruits and leaves, polyamine metabolism and the change of abscisic acidcontent in fruits had important influence on the fruitlets abscission.
0~20days after withering of flowers, fruitlets abscission ratio of leafless fruits washigher than leafy fruits. The reason was that sucrose anabolism of the adjacent leaves ofleafless fruits was effected, which led sucrose content transferred from leaves to fruitsdecreasing. The starch content of leafless fruitlets was significantly lower than leafyfruitlets. While the capacity of sucrose anabolism of leafy bearing shoots was graduallyincreased. The supply capacity of leafy bearing shoots that photosynthate with sucrosetransported from leaf to fruit enhanced. Sucrose was converted to starch and stored in fruit,which was favorable to leafy fruitlets set. At the same time, compared with leafy fruits,nitrate reductase activity was lower, polyamine catabolism was stronger and ABA contentwas more in leafless fruits.
5. Results showed that soil pH value decreased obviously after organic fertilizer andsulfur application. Under mixed application, soil organic matter, total nitrogen andavailable nitrogen content decreased at first but later increased with the increase of amountof sulfur application，while soil available sulfur content increased gradually. Under organic fertilizer and sulfur application treatment, soluble protein content of “Nanfeng” tangerineincreased and nitrate reductase activity was promoted. When sulfur application was lessthan100mg/kg, the branch diameter and the chlorophyll content both increased undermixed application. This could effectively alleviate the problem of leaf chorosis. Undermixed application(sulfur application was100mg/kg), starch accumulation in leafy fruitswas promoted, which was beneficial to fruit set.
引文
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