【作者】 于洋；

【导师】 周宝利；

【作者基本信息】 沈阳农业大学 ， 蔬菜学， 2016， 博士

【摘要】 果实着色优劣是衡量果实商品价值的重要指标之一,番茄果实色素种类和含量变化导致番茄果实色泽由绿色转为红色,主要是叶绿素含量的下降以及类胡萝卜素含量的升高。类胡萝卜素的组成及含量还关系到番茄果实的营养保健价值,果实中的糖是影响果实甜度的物质,也是某些类胡萝卜素、内源激素合成的基础原料,是影响果实品质的重要因子。内源激素可以调控类胡萝卜素和叶绿素的生物合成以及糖分的积累。外源植物激素能够调节类胡萝卜素、叶绿素、糖和内源激素的代谢。本试验以粉红(金冠五号)、橙红(石英大红)、红棕(Hongz-11)、桔黄(金冠)和浅绿(Lvfanq-05)5个不同颜色果实的番茄品种为试材,研究番茄果实不同部位类萝卜素积累规律,探讨外源GA3和ABA对北方普遍栽培的粉红色番茄果实不同部位中主要色素代谢、糖积累和内源激素含量的影响,并阐明主要色素与糖、内源激素与主要色素和糖之间的关系,初步明确外源GA3和ABA调控番茄主要色素代谢的生理机制,以期为科学调控番茄果实品质提供理论依据。本文研究的主要结果如下：1.番茄果实中类胡萝卜素积累和分布特点本试验供试的粉红、橙红、红棕、桔黄和浅绿5种颜色番茄果实中,番茄红素含量都是随着成熟进程逐渐增加,p-胡萝卜素和叶黄素含量呈现先下降后上升的趋势,红熟期番茄红素在果皮组织中含量最高,p-胡萝卜素和叶黄素在胶质组织中含量最高。不同颜色类型番茄果实红熟期时果皮组织番茄红素含量依次为：红棕>粉红>橙红>桔黄>浅绿；β-胡萝卜素含量依次为：红棕>橙红>粉红>浅绿>桔黄；叶黄素含量依次为：桔黄>橙红>红棕>粉红>浅绿。2.外源GA3和ABA对番茄果实不同部位中主要色素的影响本试验以粉红色番茄“金冠五号”为试材,外源涂抹GA3处理绿熟期番茄果实,结果表明,外源GA3处理抑制成熟过程中果实各部位类胡萝卜和番茄红素含量的增合成；对p-胡萝卜素含量的影响表现为先抑制后促进的作用；降低各部位叶黄素的含量；延缓叶绿素的降解。外源涂抹ABA处理绿熟期番茄果实,结果表明,外源ABA处理增加了番茄果实中类胡萝卜素和β-胡萝卜素的含量；对番茄红素和叶黄素表现为先抑制后促进的作用；促进叶绿素的降解。3.外源GA3和ABA对番茄果实不同部位主要色素合成相关物质糖的影响番茄果实成熟过程中各糖分的变化规律不一致,果实不同部位糖积累也有差异。番茄果实中主要是葡萄糖和果糖,其中葡萄糖含量高于果糖,各部位蔗糖含量很低。三种糖在成熟果实果皮组织中含量最高。在绿熟期用100 mg·L-1GA3涂抹番茄果实,结果表明,外源GA3抑制番茄果实各部位葡萄糖和果糖的积累,抑制了蔗糖的降解。绿熟期用100 mg·L-1外源ABA处理番茄果实,结果表明,外源ABA促进果实各部位葡萄糖、果糖和蔗糖的积累。4.外源GA3和ABA对番茄果实不同部位主要色素合成相关物质内源激素的影响本试验明确了绿熟期后番茄果实成熟过程各部位内源激素的变化趋势。内源GA3含量呈现逐渐下降趋势,成熟时果皮组织中含量最低；内源ABA含量呈现“V”型曲线变化,绿熟期后8d降至最低；内源ZT含量呈现先上升再下降的趋势,绿熟期后20d最低；内源ETH含量变化趋势呈单峰曲线,不同部位峰值出现时间不同。ABA/GA3逐渐上升；GA3/Z1、ABA/ZT和ABA/ETH先下降后上升；ETH/GA3先上升后下降；ETH/ZT没有明显规律。外源GA3处理绿熟期番茄果实,结果表明,外源GA3促进内源GA3和ABA含量增加；抑制内源ZT含量下降；降低果实中内源ETH释放量,延迟出现峰值；提高GA3/ZT、ABA7ETH和ABA/ZT比值；对ABA/GA3比值是先抑制后促进的作用；ETH/ZT和ETH/GA3与对照相比先升高后降低。外源ABA处理绿熟期番茄果实,结果表明,外源ABA使内源GA3含量降低；对内源ABA含量变化的影响表现为抑制-促进-抑制作用；促进番茄果实内源ZT含量下降；提前出现内源ETH含量峰值,对内源ETH释放量影响不大；促进ABA/GA3变大；对ETH/GA3、ABA/Z、GA3/ZT、 ETH/ZT和ABA/ETH无规律性影响。5.番茄果实主要色素与糖代谢之间的关系研究结果表明,番茄果实成熟过程中葡萄糖和果糖积累有利于各部位中类胡萝卜素、番茄红素和β-胡萝卜素的合成,促进叶绿素的降解；蔗糖的积累抑制类胡萝卜素、番茄红素和β-胡萝卜素的合成,延缓叶绿素的降解；果皮组织中叶黄素积累受糖含量的影响高于其它部位。6.番茄果实内源激素与主要色素之间的关系本试验结果表明,番茄果实中内源GA3含量高促进叶黄素和叶绿素的积累,抑制β-胡萝卜素的合成；内源ABA含量高有利于总类胡萝卜素、番茄红素和β-胡萝卜素的合成,抑制叶绿素的积累；内源ZT含量高有利于叶黄素和叶绿素的积累,不利于类胡萝卜素、番茄红素和p-胡萝卜素的合成；番茄果实中主要色素合成与内源ETH含量相关性不显著。通过激素平衡与主要色素积累相关关系分析,结果表明,较高的ABA/GA3和ABA/ZT有利于番茄果实各部位主要色素中类胡萝卜素、番茄红素和β-胡萝卜素的积累；较低的ABA/GA3和ABA/ZT抑制叶绿素降解。7.番茄果实中内源激素与糖积累之间的关系本试验结果表明,番茄果实成熟过程中内源GA3和ZT含量高有利于蔗糖积累,不利于葡萄糖和果糖合成；内源ABA含量高有利于果糖形成；较高的ABA/GA3和ABA/ZT有利于番茄果实各部位葡萄糖和果糖的积累,不利于蔗糖的形成。8.外源GA3对番茄果实主要色素调控生理机制外源GA3对番茄果实主要色素作用的生理机制初步认为是：外源GA3抑制类胡萝卜素和番茄红素积累,是通过抑制葡萄糖和果糖积累,抑制蔗糖降解实现的；促进p-胡萝卜素合成主要是由于外源GA3提高内源ABA含量；抑制叶黄素积累可能是提高ABA/GA3;延缓叶绿素降解是由于外源GA3处理增加了内源GA3含量,抑制葡萄糖和果糖积累,抑制蔗糖降解所致。9.外源ABA对番茄果实主要色素调控生理机制外源ABA对主要色素调控的生理机制初步认为是：外源ABA促进番茄果实中类胡萝卜素和番茄红素的积累是由于外源ABA处理降低内源ZT含量,提高ABA/GA3比值,促进葡萄糖和果糖的积累所致；对叶黄素积累的促进作用,是由于外源ABA处理增加蔗糖含量；促进p-胡萝卜素的积累是由于外源ABA处理降低内源GA3和ZT含量,提高ABA/GA3,促进葡萄糖和果糖的积累；加速叶绿素的降解是由于外源ABA处理降低内源GA3含量,增加葡萄糖和果糖的含量,提高ABA/GA3比值所致。更多还原

【Abstract】 Fruit coloration is one of the important indicators to measure the commodity value of the fruit. And the changes of pigment in tomato fruits may affect the color from green to red, mainly due to the decrease of chlorophyll content and the increase of carotenoid content. The composition and content of carotenoid also affect the nutrient value of tomato fruit. The sugar in fruit is an important factor affecting fruit quality, which affects fruit sweetness and also is the based raw materials for the synthesis of carotenoid and endogenous hormone. Endogenous hormone has the capability to regulate carotenoid biosynthesis, chlorophyll biosynthesis and the accumulation of sugar. Plant exogenous hormone has the capability to regulate the metabolism of carotenoid, chlorophyll, sugar and endogenous hormones.In this research, the accumulation patterns of carotenoid were studied in the different tissues of 5 different tomato phenotypes, such as pink (JinGuan 5), orange (ShiYingDaHong), red (Hongz-11), yellow (JinGuan) and green (Lvfanq-05). The effects of exogenous GA3 and ABA were explored on the metabolism of main pigment, accumulation of sugar and the content of endogenous hormone in the different tissues of pink tomato fruit which commonly cultivated in north China. And the relationship between the main pigment and sugar was elucidated, meanwhile, the relationship among the endogenous hormones, the main pigment and sugar also was clarified. Physiological mechanism about the metabolism of main pigment was definited by regulation of exogenous GA3 and ABA. This study provided the theory basis for scientific regulation of tomato fruit quality. The main results were as follows.1. Accumulation and distribution of carotenoids in tomato fruitThe experiment tested many tomato fruits with 5 different tomato phenotypes-pink, orange, red, yellow and green. The content of lycopene in tomato fruit was increased gradually with the fruit becoming rippen, while the highest content of lycopene was in pericarp when it was in red ripening period. The P-carotene and xanthophyll were showed a trend of increasing after decreasing. And the highest contents of β-carotene and xanthophyll were in pectin.The highest content of lycopene was in the peel tissue of red fruit, then was in the pink fruit, the following was orange fruit, yellow fruit and green fruit. The contents of p-carotene from high to low were in red fruit, orange fruit, pink fruit, green fruit and yellow fruit. The highest content of xanthophyll was in the yellow fruit, and second was in the orange fruit, following was in the red fruit, pink fruit and green fruit.2. Effects of exogenous GA3 and ABA on the main pigment in different parts of tomato fruitThis study chose pink tomato strain Golden Crown No.5, and treated tomato fruit of green maturity with 100 mg ·L-1 GA3 and ABA. The results showed that the contents of total carotenoids and lycopene in the fruits were inhibited by exogenous GA3 treatment. The effect on the content of β-carotene was inhibited firstly and then facilitated. And the content of xanthophyll in various parts was significantly reduced. And it delayed the degradation of chlorophyll and inhibited the formation of tomato fruit-specific color and luster. After dealing with exogenous ABA, the contents of total carotenoids and P-carotene in tomato fruit were increased, while it inhibited firstly and then facilitated, the growth of lycopene and xanthophyll. The treatment of ABA promoted the degradation of chlorophyll and the formation of tomato fruit color.3. Effects of exogenous GA3 and ABA to sugar related substances of main pigment synthesis in different parts of tomato fruitDuring the ripening of tomato fruit, the variation of sugar content was different, and the sugar accumulation in different parts of fruit was also different. Tomato fruit mainly contained glucose and fructose, in which the glucose content was higher than fructose, and sucrose content of each part was very low. The three sugars had the highest content in mature fruit pericarp tissue. The results showed that exogenous GA3 inhibited the accumulation of glucose and fructose in various parts of tomato fruit, and inhibited the degradation of sucrose by exogenous GA3 to treat tomato fruit with green maturity. Then, the fruits were treated with exogenous ABA, and the results showed that exogenous ABA promoted the accumulation of glucose, fructose and sucrose in all parts of fruits.4. Effects of exogenous GA3 and ABA on endogenous hormones related to main pigment synthesis in different parts of tomato fruits.In this study, the change trend of endogenous hormones was determined in different parts of tomato fruit after green ripening during the fruit ripening process. The content of endogenous GA3 showed a decreasing trend, and the content of endogenous GA3 was the lowest at maturity. The content of endogenous ABA demonstrated a "V"-shaped curve, and reached its minimum at 8 d after green ripening. At 20 d after green ripening, the content of endogenous ABA in peel tissue was the highest, and the content of endogenous ZT showed a trend of increasing first and then descending, and the lowest was at 20 d after green ripening. The trend of endogenous ETH displayed a single peak curve in different parts at different peak time. ABA/GA3 increased gradually. Endogenous GA3/ZT, ABA/ZT and ABA/ETH decreased first and then increased. ETH/GA3 first increased and then decreased, but ETH/ZT no obvious law.Smearing of GA3 to treat tomato fruit with green maturity, the consequence showed that exogenous GA3 could promote the increase of endogenous GA3 and ABA contents. Inhibition of endogenous ZT content decreased, decreased the release of endogenous ETH and delayed the peak value, increased GA3/ZT, ABA/ETH, ABA/ZT. And the ratio of ABA/GA3 was first inhibited and then promoted. ETH/ZT and ETH/GA3 increased and then decreased compared with the control. The results showed that exogenous ABA reduced the content of endogenous GA3 after smearing of exogenous ABA on green ripe tomato fruit. The effect of endogenous ABA content was inhibited-promoted-inhibited, promoted the decrease of endogenous ZT content in tomato fruit. The peak of endogenous ETH appeared earlier, which had little effect on the release of endogenous ETH, promoted the ABA/GA3 to become bigger. And no influence of ETH/GA, ABA/ZT, GA3/ZT, ETH/ZT and ABA/ETH.5. The relationship between main pigment and sugar metabolism of tomato fruitThe accumulation of glucose and fructose was beneficial to the synthesis of carotenoids, lycopene and P-carotene, the main pigment in various parts of tomato fruit, and promoted the degradation of chlorophyll. The accumulation of sucrose was disadvantageous to the carotenoids, lycopene and (3-carotene, and was beneficial to delay the degradation of chlorophyll. The accumulation of xanthophyll in the peel was affected by sugar content higher than other parts.6. The relationship between endogenous hormones and main pigments in tomato fruitThe high content of endogenous GA3 in tomato fruit was beneficial to the accumulation of xanthophyll and chlorophyll in the main pigment of tomato fruit, which was not conducive to the synthesis of β-carotene. The high content of endogenous ABA was favorable to the synthesis of carotenoid, lycopene and P-carotene, and inhibited the accumulation of chlorophyll. The high content of endogenous ZT was beneficial to the accumulation of xanthophyll and chlorophyll, which was not conducive to the synthesis of carotenoid, lycopene and β-carotene.There was no significant correlation between the main pigment synthesis and the content of endogenous ETH in tomato fruit.Analysis the relationship of hormone balance and main pigment accumulation, this experiment confirmed that the higher ABA/GA3 and ABA/ZT favored the accumulation of carotenoid, lycopene and P-carotene in the main pigments of tomato fruit, the lower of that inhibited chlorophyll degradation.7. The relationship between endogenous hormone and sugar accumulation in tomato fruitThe high contents of endogenous GA3 and ZT in the ripening process of tomato fruit were beneficial to the accumulation of sucrose and was not beneficial to the accumulation of glucose and fructose.The high content of endogenous ABA was beneficial to the formation of fructose. Based on the analysis of the relationship between hormone balance and sugar accumulation, the results showed that higher ABA/GA3 and ABA/ZT were beneficial to the accumulation of glucose and fructose in various parts of tomato fruit, which was not beneficial to the formation of sucrose.8. Physiological mechanism of exogenous GA3 on main pigment regulation in tomato fruitIn this study, tomato fruit was treated with exogenous GA3. The results showed that the physiological mechanism of exogenous GA3 on main pigment regulation was believed to inhibit the accumulation of carotenoids and lycopene by inhibiting of glucose, fructose accumulation and sucrose degradation. To promote the synthesis of β-carotene by increasing exogenous ABA, to inhibit the xanthophyll accumulation by increasing the ratio ABA to GA3, to delay chlorophyll degradation by increasing endogenous GA3 content, inhibiting of glucose, fructose accumulation and sucrose degradation.9. Physiological mechanism of exogenous ABA on main pigment regulation in tomato fruitIn this study, the tomato fruit was treated with exogenous ABA. The results showed that the physiological mechanism of exogenous ABA to primary pigments was considered as follows:exogenous ABA treatment reduced the content of endogenous ZT, thus increased ABA/GA3 ratio, and the accumulation of glucose and fructose were enhanced, thereby promoted the accumulation of carotenoid and lycopene. Exogenous ABA increased xanthophyll content by increaing the content of sucrose. exogenous ABA treatment reduced the contents of endogenous GA3 and ZT, thus increased ABA/GA3 ratio, and the accumulation of glucose and fructose were enhanced, thereby promoted the accumulation of β-carotene, exogenous ABA treatment reduced the contents of endogenous GA3, thus increased ABA/GA3 ratio, and the accumulation of glucose and fructose were enhanced, thereby decreased the chlorophyll.更多还原