成熟期不同的梨品种果实生长发育机理探讨
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摘要
梨属于蔷薇科(Resaceae)梨亚科(Pomoideae)梨属(Pyrus)植物。我国是世界梨的主要原产地和栽培地之一,已有3000多年的栽培历史。目前,对梨果实采前生理即生长发育过程的系统研究较少,特别是针对我国南方梨在生长发育期间处于高温、高湿和光、热、水充裕的气候条件下其果实生长发育及其内部机理研究甚少,多集中在与果实品质相关的指标,或局限于某一类激素和某一种酶的变化,没有全面、系统的综合研究其各种内部因素的平衡关系和具体的调控机理。而针对不同梨品种果实生长发育历程不同的研究主要集中在品种遗传特性上,对影响果实生长发育的其它内部因素研究很少,或者只是简单根据一个或几个指标说明其含量变化,没有将定性和定量结合起来研究影响果实生长发育期的各种内部因子与生长发育期差异产生的相关性。另外,在梨果实生长发育期间关于果实内果糖、葡萄糖、蔗糖、IAA、ABA、GAs、和CTK等物质的变化规律的报道不尽一致,有关不同品种梨果实内多胺、细胞壁酶和LOX等物质的消长规律及其差异尚未见研究报道。而在其它果树如猕猴桃、草莓、柿、葡萄、苹果等上也多利用同一成熟期品种从遗传特性或某一个影响因子来进行果实发育及调控机理研究。因此,对成熟期不同的梨果实生长发育过程中内含物质、细胞壁酶类、内源激素及内源多胺进行研究,不仅可以进一步了解梨果实生长发育中生理生化的变化规律,还可以进一步揭示梨果实生长发育及成熟机理,从而为进一步研究果树生理学和果树生产提供一定的理论依据。
本研究以成都平原引种的5年生早蜜梨和黄金梨为试材,在一般生产管理条件下,采用田间试验和室内分析相结合的方法,比较了两个成熟期不同的梨品种的基本性状、叶片光合特性及果实品质,并系统地研究了两品种果实生长发育过程中果实的生长动态、各种内含物质、PE、PG、Cx和LOX的活性以及内源IAA、GA_3、ZT、ABA、内源Put、Spd和Spm含量的变化规律和特点,探讨了两个品种果实生长发育的异同点。其主要结果如下:
1、早蜜梨和黄金梨在成都平原的引种表现都较好,基本性状相似。两品种植株生长旺盛,发育良好,物候期较其引种地早,果实品质优良,但早蜜梨口味较黄金梨更甜。在相同季节早蜜梨叶片的光合速率值略高于黄金梨,叶绿素a及总量也高于黄金梨。两品种叶片的光合速率主要取决于叶绿素,特别是叶绿素a含量。早蜜梨较黄金梨更为耐阴。
2、两品种梨果实发育的各个阶段长短差异明显。早蜜梨和黄金梨的整个果实生长发育过程都可划分为3个阶段。早蜜梨果实生育期(或果实发育天数)为100d,坐果至花后30d,为果实发育前期;花后30d～70d,为果实发育中期;花后70d～100d,为果实发育后期。黄金梨果实生育期(或果实发育天数)为130d,坐果至花后50d,为果实发育前期;花后50d～100d,为果实发育中期,花后100d～130d,为果实发育后期。就3个时期所持续的时间而言,早蜜梨为30d、40d和30d,黄金梨为50d、50d和30d,后者在果实发育前期和中期分别长20d和10d。另外,两个品种果实纵横径生长都符合logistic曲线变化。
3、两品种果实发育期间各种内含物含量变化各有特点。
(1)两品种果实中可溶性糖、果糖、葡萄糖和蔗糖含量随着果实发育天数的增加而呈逐渐上升趋势。糖分以果糖为主,含量大小为果糖＞葡萄糖＞蔗糖,且早蜜梨果实的3种糖分含量都明显高于黄金梨,两品种果实成熟时可溶性总糖含量差异极显著(p＜0.01)。两品种在子房膨大初期和果实发育后期都出现两个含量迅速增长期,而果实发育后期是糖积累的关键时期。从增长速度来看,果糖转化和积累最快,葡萄糖次之,蔗糖最慢。另外,早蜜梨果实中果糖、葡萄糖,黄金梨果实中果糖、葡萄糖、蔗糖和可溶性总糖积累都符合logistic曲线变化。
(2)两品种果肉中蛋白质含量在果实发育初期较高,随着果实的发育而逐渐降低,但二者在果实发育的大部分时期含量差异不大。整个果实发育期间,早蜜梨出现两次含量高峰,黄金梨变化较为平缓。
(3)早蜜梨果实在果实发育初期和后期游离氨基酸含量高于黄金梨,在果实成熟时极显著高于黄金梨(p＜0.01),而在果实发育中期则低于黄金梨。
(4)两品种果肉中核酸含量在果实发育初期都较高,随着果实的发育逐渐降低。早蜜梨果肉中核酸含量分别在花后10d、50d和90d出现高峰,果实成熟时极显著高于黄金梨。黄金梨果肉在花后20d时含量达到最大。
4、两品种果实发育期间相关酶类变化趋势相似。
(1)两品种果肉中PG活性总体上随果实的发育逐渐升高。早蜜梨在花后50d出现活性高峰,黄金梨果肉呈现升降交替变化的趋势,分别在花后30d、90d和130d出现3次活性高峰。早蜜梨在果实发育中期PG活性高于黄金梨,而果实发育后期二者差异不大。果肉中PG活性与果实发育中期和后期密切相关。
(2)两品种果肉中PE活性在果实发育前期较高,随着果实的发育逐渐降低,果实成熟前有所回升。早蜜梨果肉在花后10d、70d和100d出现3次PE活性高峰,黄金梨果肉在花后10d、80d和130d也出现3次活性高峰。整个果实发育过程中,前期黄金梨PE活性高于早蜜梨,而中后期则低于后者。
(3)两品种果肉中Cx活性在果实发育前期较低,随着果实发育成熟而逐渐升高,果实成熟前有所降低。早蜜梨在花后30d～70d和花后80d～90d有2次活性迅速升高,而黄金梨在花后20d～30d、花后50d～70d和花后80d～120d共有3次活性迅速升高。
(4)两个品种果肉中LOX活性在果实发育初期较低,随着果实的发育成熟而逐渐升高。早蜜梨在花后20d～40d和50d～90d共出现2次活性快速上升,而黄金梨在果实发育的各个时期增加幅度比较均匀。
5、果肉和种子中内源激素含量变化差异是导致两品种梨果实发育历程不同的重要因素。
(1)同一生境下成熟期相对较晚的黄金梨果实发育初期果肉中IAA和ZT含量高于成熟期较早的早蜜梨,且持续时间更长,而GA_3含量除花后20d时低于早蜜梨外都较早蜜梨高。黄金梨果肉细胞分裂的强度可能大于早蜜梨,分裂时间也可能长于早蜜梨。果实发育的大部分时期黄金梨果实内3种生长促进型激素(IAA、GA_3、ZT)含量高于早蜜梨,但含量高峰出现的时间比早蜜梨滞后10d～20d。
(2)两品种果肉中ABA含量在花后30d有一次升高,可能与此期果实的脱落密切相关。两品种果实成熟时果肉中ABA含量差异不显著,但早蜜梨在成熟前出现含量高峰的时间比黄金梨早10d,其果实可能启动成熟较早,成熟期较短。
(3)早蜜梨种子中IAA和ZT含量明显高于同期黄金梨的含量,这可能导致早蜜梨比黄金梨种子更早地进入下一个发育阶段,也利于转运至果肉中,促进果肉细胞的发育。黄金梨果实发育的大部分时期种子中GA_3含量高于同期早蜜梨的含量,高含量的GA_3持续时间更长,较早蜜梨花后第1次含量高峰出现的时间提前10d,这可能与黄金梨种子自身合成GA_3能力较强有关,也利于转运至果肉中促进果肉细胞的膨大。另外,早蜜梨种子中ABA含量高峰出现的时间比黄金梨提前约30d,因此,早蜜梨果实更早地启动了成熟。
(4)早蜜梨果肉和种子中IAA比值在果实发育的大部分时期都小于1,说明其种子是IAA合成的主要场所,特别在果实发育初期合成能力更强。两品种果肉和种子中ZT含量的比值变化相似,但早蜜梨比值小于黄金梨,可能与其种子中合成ZT的能力较强有关。两品种果肉在果实发育后期合成ZT的能力极低,而种子在成熟前还具有一定的合成能力。早蜜梨果实发育期间果肉和种子中GA_3含量的比值都小于1,说明了胚乳是其合成的主要场所。
6、内源多胺与两品种果实幼果发育的差异关系密切。
(1)早蜜梨和黄金梨果肉和种子中Put、Spd和Spm及游离多胺总量在果实发育初期都较高,这与两个品种果实发育初期旺盛的细胞分裂有关。两个梨品种果肉中3种游离多胺含量在果实发育初期差异较大,早蜜梨在花后10d时3种游离多胺含量都极显著高于黄金梨(p＜0.01);种子中,除Put外其它2种游离多胺在花后20d时含量都极显著高于黄金梨(p＜0.01)。综合果肉和种子的情况,可以说明早蜜梨幼果的早期发育更为旺盛。但黄金梨果肉中Spm含量在果实发育的大部分时期高于早蜜梨,特别是在花后20d有一个明显升高的过程,与同期早蜜梨含量差异极显著(p＜0.01),推测Spm与黄金梨幼果第一次迅速生长关系密切,
(2)两品种果肉和种子中Put含量在果实发育后期出现了第2次高峰,峰值接近,但黄金梨出现的时间滞后约30d～40d,成熟期也比早蜜梨晚30d。这可能与梨果实发育后期乙烯的释放和果实成熟启动有关。
The pear is a plant which belongs to Rosaceae, Pomoideae, Pyrus. Pears are by far one of the most important deciduous tree fruits, and are widely grown in China. Fruit growth and development is one of the foremost life activities for fruit trees and is regulated by complex interactions. The early researches on it focused mainly on fruit quality, or on one endogenous hormone or enzyme only. The systematic studies are not enough. Especially, pear fruits grow and develop more differently on the condition of high temperature and humidity, rich light, caloric and water in south of China than in north. On the other hand, different procedures of growth and development of pear fruits exist among the different cultivars. The characteristics of heredity are looked as one of the important factors to elucidate this physiological phenomenon. But in fact there are so many other factors influencing fruit growth and development. Qualitative and quantitative researches are extremely necessarily combined to clarify the relevance of these factors and ripening-season of pear fruits. The past research indicated that fructose, glucose, sucrose and endogenous IAA, ABA, GAs and CTK contents in pear fruits didn't change in same regularity during fruit growth and development for different experiments. And there have no reports about the changing regularity of endogenous polyamines contents, activiety of cell wall enzymes and LOX during fruit growth and development among the different pear cultivars. Only one cultivar or factor was selected to study the mechanism of fruit growth and development in kiwifruit, strawberry, persimmon, grape, apple and some other frutis. Therefore, researches on chemical substances, such as carbohydrates, proteins, endogenous hormones and polyamines are very crucial to find the difference of growth regularity and disclose mechanism of fruit growth and development of pear fruits among the pear cultivars in different ripening-seasons. To some extent it could provide advice for fruit physiologist and field production.
Both Zaomi and WhangKeumbae pear in 5-year-old trees with uniform growing vigor were selected for this experiment. Trees were grafted on Pyrus pashia Buch-Ham rootstocks and planted in Chengdu plain. Cultural management such as fertilization and pest control were the same as those used in a commercial orchard, without fruit thinning and bagging. Characteristics of trees, photosynthetic characteristics of leaves and fruit quality were analyzed for these two pear cultivars. At the same time carbohydrates, proteins and nucleic acid contents, activities of pectinesterase (PE), polygalacturonase (PG), cellulase (Cx) and lipoxygenase (LOX) were determined along with endogenous indol-3-acetic acid (IAA), gibberellins (GA_s), zeatin (ZT), abscisin acid (ABA) and putrescine(Put), spermidine(Spd), spermine(Spm) contents during the period of fruit growth and development. The followings were the results:
1、Zaomi and Whangkeumbae pear had similar characteristics of trees in Chengdu plain. The trees grew strongly and had earlier phenophase than that in origin region. The fruits of these two cultivars were in similar shape and good in fruit quality. However, Zaomi pear fruits were sweeter than Whangkeumbae. Photosynthesis rate, chlorophyll a and the total chlorophyll contents of leaves of Zaomi pear were little higher than that of Whangkeumbae pear in the same season. Chlorophyll especially chlorophyll a was the key factor influencing photosynthesis rate for this two pear cultivars. And Zaomi pear trees had stronger tolerance of shading.
2、There were obvious differences of developmental stages of fruits between these two pear cultivars. The growth curve of fruit showed as single sigmoid and was divided into three phases. The fruit growth and development lasted 100days and 130days for Zaomi and Whangkeumbae pear, respectively. For Zaomi pear fruits, period from fruit setting to 30 days after full bloom (DAFB) was looked as the early stage, period from 30 to 70 DAFB as middle stage, and period from 70 to 100 DAFB as late stage. For Whangkeumbae pear fruits, period from fruit setting to 50DAFB was divided as the early stage, period from 50 to 100DAFB as the middle stage, period from 100 to 130DAFB as the late stage. The early stage of Whangkeumbae pear held longer for 20days than that of Zaomi pear, and the middle stage kept longer for 10days. The growth of longitudinal length and transverse diameter during fruit growth and development accorded with logistic curve for these two cultivars.
3、The contents of main chemical substances changed in different regularities during fruit growth and development for the experimental pear cultivars.
(1) Content of soluble sugar, fructose, glucose and sucrose rose gradually along with fruit growth and development for both Zaomi and Whangkeumbae pear. The fructose content was the highest at any given stage of fruit development. Glucose content was at second and higher than sucrose. The contents of three sugar components of Zaomi pear fruits were obviously higher than that of Whangkeumbae pear. Significant difference of soluble sugar contents was produced between these two cultivars at the ripening stage of fruit (p<0.01). Sugar contents increased very fast in the early stage of ovary developing and the late stage of fruit development for the two cultivars. And the latter stage was significant for accumulating sugar. Among the three sugar components fructose transformed and accumulated fastest followed with glucose and sucrose. The accumulation of fructose and glucose of Zaomi pear, and that of fructose, glucose, sucrose and soluble sugar of Whangkeumbae pear accorded with logistic curve during fruit growth and development.
(2) Protein contents kept at high level at the initial stage and decreased along with fruit development for Zaomi and Whangkeumbae pear. There were no obvious differences during the most stages between them. Zaomi pear fruits showed two peaks of content, but Whangkeumbae pear fruits changed in slow rate.
(3) For free amino acid content, it was higher in Zaomi pear fruits than that in Whangkeumbae pear fruits both at initial and late stages, and was significantly higher at fruit maturation (p<0.01). However, it was lower at the middle stage.
(4) Nucleic acid contents kept at high level at initial stage and decreased gradually along with fruit development for the two cultivars. It peaked at 10, 50 and 90 DAFB for Zaomi pear fruits and was significantly higher than that of Whangkuembae pear at fruit maturation (p<0.01). It peaked at 20DAFB for Whangkeumbae pear fruits.
4、The activity of enzymes changed in similar trend during fruit growth and development for these two pear cultivars.
(1) For the two cultivars, polygalacturonase (PG) activity increased gradually along with fruit development. It peaked at 50DAFB for Zaomi pear fruits, and peaked at 30, 90and 130DAFB for Whangkeumbae pear fruits. Zaomi pear fruits had higher activity of PG at the middle stage but had no obvious difference with Whangkeumbae pear fruits.
(2) The activity of pectinesterase (PE) kept at high level at the early stage and decreased gradually along with fruit development, but rose in slightly range before maturation for the two cultivars. It peaked at 10, 70 and 100DAFB for Zaomi pear fruits, and at 10, 80 and 130DAFB for Whangkeumbae pear fruits which had higher activity at early stage than Zaomi pear fruits, and had lower activity at the middle and late stage.
(3) The activity of cellulase (Cx) was a little low at the early stage and increased along with fruit development up to maturation followed with slightly decrease for the two cultivars. It rose sharply from 30 to 70DAFB and from 80 to 90DAFB for Zaomi pear fruits, from 20 to 30DAFB, from 50 to 70DAFB and from 80 to 120DAFB for Whangkeumbae pear fruits.
(4) The activity of lipoxygenase (LOX) was a little low at initial stage and increased along with fruit development for the two cultivars. It rose sharply from 20 to 40DAFB and from 50 to 90DAFB, but increased in slow rate for Whangkeumbae pear fruits during fruit development.
5、The difference of endogenous hormones contents both in pulp and seeds was the key factor resulting in different fruit growth and development of Zaomi and Whangkeumbae pear.
(1) In the same habitat IAA and ZT contents in pulp of Whangkeumbae pear which matured later were higher than that of Zaomi pear which matured earlier at initial stage and held in longer time. GA3 contents were higher than that of Zaomi pear fruits at the most stages except of 20DAFB. Cell division in pulp of Whangkeumbae pear was stronger and held for longer time. The three endogenous hormones IAA, GA_3 and ZT contents in pulp were higher than that of Zaomi pear at the most stages of fruit growth and development. However, peaks of contents for Whangkeumbae pear appeared late for 10 to 20days.
(2) An increase for ABA contents appeared in pulp of the two cultivars at 30DAFB. It might result in dropping of young fruits at this stage. ABA contents in pulp of Zaomi pear peaked earlier for 20days before fruit maturation. Thus Zaomi pear fruits might shift to maturate much earlier, and had shorter period of fruit growth and development.
(3) IAA and ZT contents in seeds of Whangkeumbae pear were obviously lower than that of Zaomi pear at the same stage. It might result in shifting into next developing stage for seeds, and was advantageous of being transferred into pulp for cell development. At the most stages GA_3 contents in seeds of Whankeumbae pear were higher than that of Zaomi pear at the same time. And it lasted for longer time with high contents and got to the first peak earlier for 10 days. This could indicate that Whangkeumbae pear fruits had vigorous production of GA_3 in seeds, which could be transferred into pulp to promote cell expansion. On the other hand, ABA contents in seeds of Zaomi pear peaked earlier for 30days. Therefore it could go into maturing earlier than Whangkeumbae pear.
(4) The ratio of IAA contents between pulp and seeds was less than 1 at the most stages for Zaomi pear. Therefore IAA might be mainly synthesized in seeds, especially at initial stage of fruit growth and development. For the ratio of ZT contents this two pear cultivars had similar changing trend. But Zaomi pear, which had vigorous synthesis of ZT in seeds, showed lower ratio value than Whangkeumbae pear. Few of ZT could be synthesized in pulp at the late stage for the two cultivars, but in seeds some ZT could be synthesized before maturation. The ratio of GA_3 contents was less than 1 for Zaomi pear during fruit development, so endosperm was the important place to produce endogenous GA_3.
6、Endogenous polyamines were very important for development of young fruits.
(1) This two pear cultivars contained high level of Put, Spd, Spm and free polyamines contents both in pulp and seeds at initial stage. It could stimulate vigorous cell division. Put, Spd and Spm contents in pulp of Zaomi pear were significantly higher than that of Whangkeumbae pear at 10DAFB (p<0.01). In seeds, Spd and Spm contents were significantly higher than that of Whangkeumbae pear at 20DAFB (p<0.01). It could speculate that young fruits of Zaomi pear developed more vigorous. However, the Spm contents in pulp of Whangkeumbae pear were higher at the most stages, especially it rose sharply from 10 to 20DAFB and was significantly higher than that of Zaomi pear at the same stage. So Spm contents might be very crucial for the first fast growth of young fruits of Whangkeumbae pear.
(2) Put contents both in pulp and seeds got the peak value for two times at the late stage for the two cultivars. And these two peaking values were approximate, but it appeared late for 30 to 40 days for Whangkeumbae pear. The fruit maturated late for 30days, too. This could be correlated with releasing of ethylene and starting to maturate at the late stage.
本研究以成都平原引种的5年生早蜜梨和黄金梨为试材,在一般生产管理条件下,采用田间试验和室内分析相结合的方法,比较了两个成熟期不同的梨品种的基本性状、叶片光合特性及果实品质,并系统地研究了两品种果实生长发育过程中果实的生长动态、各种内含物质、PE、PG、Cx和LOX的活性以及内源IAA、GA_3、ZT、ABA、内源Put、Spd和Spm含量的变化规律和特点,探讨了两个品种果实生长发育的异同点。其主要结果如下:
1、早蜜梨和黄金梨在成都平原的引种表现都较好,基本性状相似。两品种植株生长旺盛,发育良好,物候期较其引种地早,果实品质优良,但早蜜梨口味较黄金梨更甜。在相同季节早蜜梨叶片的光合速率值略高于黄金梨,叶绿素a及总量也高于黄金梨。两品种叶片的光合速率主要取决于叶绿素,特别是叶绿素a含量。早蜜梨较黄金梨更为耐阴。
2、两品种梨果实发育的各个阶段长短差异明显。早蜜梨和黄金梨的整个果实生长发育过程都可划分为3个阶段。早蜜梨果实生育期(或果实发育天数)为100d,坐果至花后30d,为果实发育前期;花后30d～70d,为果实发育中期;花后70d～100d,为果实发育后期。黄金梨果实生育期(或果实发育天数)为130d,坐果至花后50d,为果实发育前期;花后50d～100d,为果实发育中期,花后100d～130d,为果实发育后期。就3个时期所持续的时间而言,早蜜梨为30d、40d和30d,黄金梨为50d、50d和30d,后者在果实发育前期和中期分别长20d和10d。另外,两个品种果实纵横径生长都符合logistic曲线变化。
3、两品种果实发育期间各种内含物含量变化各有特点。
(1)两品种果实中可溶性糖、果糖、葡萄糖和蔗糖含量随着果实发育天数的增加而呈逐渐上升趋势。糖分以果糖为主,含量大小为果糖＞葡萄糖＞蔗糖,且早蜜梨果实的3种糖分含量都明显高于黄金梨,两品种果实成熟时可溶性总糖含量差异极显著(p＜0.01)。两品种在子房膨大初期和果实发育后期都出现两个含量迅速增长期,而果实发育后期是糖积累的关键时期。从增长速度来看,果糖转化和积累最快,葡萄糖次之,蔗糖最慢。另外,早蜜梨果实中果糖、葡萄糖,黄金梨果实中果糖、葡萄糖、蔗糖和可溶性总糖积累都符合logistic曲线变化。
(2)两品种果肉中蛋白质含量在果实发育初期较高,随着果实的发育而逐渐降低,但二者在果实发育的大部分时期含量差异不大。整个果实发育期间,早蜜梨出现两次含量高峰,黄金梨变化较为平缓。
(3)早蜜梨果实在果实发育初期和后期游离氨基酸含量高于黄金梨,在果实成熟时极显著高于黄金梨(p＜0.01),而在果实发育中期则低于黄金梨。
(4)两品种果肉中核酸含量在果实发育初期都较高,随着果实的发育逐渐降低。早蜜梨果肉中核酸含量分别在花后10d、50d和90d出现高峰,果实成熟时极显著高于黄金梨。黄金梨果肉在花后20d时含量达到最大。
4、两品种果实发育期间相关酶类变化趋势相似。
(1)两品种果肉中PG活性总体上随果实的发育逐渐升高。早蜜梨在花后50d出现活性高峰,黄金梨果肉呈现升降交替变化的趋势,分别在花后30d、90d和130d出现3次活性高峰。早蜜梨在果实发育中期PG活性高于黄金梨,而果实发育后期二者差异不大。果肉中PG活性与果实发育中期和后期密切相关。
(2)两品种果肉中PE活性在果实发育前期较高,随着果实的发育逐渐降低,果实成熟前有所回升。早蜜梨果肉在花后10d、70d和100d出现3次PE活性高峰,黄金梨果肉在花后10d、80d和130d也出现3次活性高峰。整个果实发育过程中,前期黄金梨PE活性高于早蜜梨,而中后期则低于后者。
(3)两品种果肉中Cx活性在果实发育前期较低,随着果实发育成熟而逐渐升高,果实成熟前有所降低。早蜜梨在花后30d～70d和花后80d～90d有2次活性迅速升高,而黄金梨在花后20d～30d、花后50d～70d和花后80d～120d共有3次活性迅速升高。
(4)两个品种果肉中LOX活性在果实发育初期较低,随着果实的发育成熟而逐渐升高。早蜜梨在花后20d～40d和50d～90d共出现2次活性快速上升,而黄金梨在果实发育的各个时期增加幅度比较均匀。
5、果肉和种子中内源激素含量变化差异是导致两品种梨果实发育历程不同的重要因素。
(1)同一生境下成熟期相对较晚的黄金梨果实发育初期果肉中IAA和ZT含量高于成熟期较早的早蜜梨,且持续时间更长,而GA_3含量除花后20d时低于早蜜梨外都较早蜜梨高。黄金梨果肉细胞分裂的强度可能大于早蜜梨,分裂时间也可能长于早蜜梨。果实发育的大部分时期黄金梨果实内3种生长促进型激素(IAA、GA_3、ZT)含量高于早蜜梨,但含量高峰出现的时间比早蜜梨滞后10d～20d。
(2)两品种果肉中ABA含量在花后30d有一次升高,可能与此期果实的脱落密切相关。两品种果实成熟时果肉中ABA含量差异不显著,但早蜜梨在成熟前出现含量高峰的时间比黄金梨早10d,其果实可能启动成熟较早,成熟期较短。
(3)早蜜梨种子中IAA和ZT含量明显高于同期黄金梨的含量,这可能导致早蜜梨比黄金梨种子更早地进入下一个发育阶段,也利于转运至果肉中,促进果肉细胞的发育。黄金梨果实发育的大部分时期种子中GA_3含量高于同期早蜜梨的含量,高含量的GA_3持续时间更长,较早蜜梨花后第1次含量高峰出现的时间提前10d,这可能与黄金梨种子自身合成GA_3能力较强有关,也利于转运至果肉中促进果肉细胞的膨大。另外,早蜜梨种子中ABA含量高峰出现的时间比黄金梨提前约30d,因此,早蜜梨果实更早地启动了成熟。
(4)早蜜梨果肉和种子中IAA比值在果实发育的大部分时期都小于1,说明其种子是IAA合成的主要场所,特别在果实发育初期合成能力更强。两品种果肉和种子中ZT含量的比值变化相似,但早蜜梨比值小于黄金梨,可能与其种子中合成ZT的能力较强有关。两品种果肉在果实发育后期合成ZT的能力极低,而种子在成熟前还具有一定的合成能力。早蜜梨果实发育期间果肉和种子中GA_3含量的比值都小于1,说明了胚乳是其合成的主要场所。
6、内源多胺与两品种果实幼果发育的差异关系密切。
(1)早蜜梨和黄金梨果肉和种子中Put、Spd和Spm及游离多胺总量在果实发育初期都较高,这与两个品种果实发育初期旺盛的细胞分裂有关。两个梨品种果肉中3种游离多胺含量在果实发育初期差异较大,早蜜梨在花后10d时3种游离多胺含量都极显著高于黄金梨(p＜0.01);种子中,除Put外其它2种游离多胺在花后20d时含量都极显著高于黄金梨(p＜0.01)。综合果肉和种子的情况,可以说明早蜜梨幼果的早期发育更为旺盛。但黄金梨果肉中Spm含量在果实发育的大部分时期高于早蜜梨,特别是在花后20d有一个明显升高的过程,与同期早蜜梨含量差异极显著(p＜0.01),推测Spm与黄金梨幼果第一次迅速生长关系密切,
(2)两品种果肉和种子中Put含量在果实发育后期出现了第2次高峰,峰值接近,但黄金梨出现的时间滞后约30d～40d,成熟期也比早蜜梨晚30d。这可能与梨果实发育后期乙烯的释放和果实成熟启动有关。
The pear is a plant which belongs to Rosaceae, Pomoideae, Pyrus. Pears are by far one of the most important deciduous tree fruits, and are widely grown in China. Fruit growth and development is one of the foremost life activities for fruit trees and is regulated by complex interactions. The early researches on it focused mainly on fruit quality, or on one endogenous hormone or enzyme only. The systematic studies are not enough. Especially, pear fruits grow and develop more differently on the condition of high temperature and humidity, rich light, caloric and water in south of China than in north. On the other hand, different procedures of growth and development of pear fruits exist among the different cultivars. The characteristics of heredity are looked as one of the important factors to elucidate this physiological phenomenon. But in fact there are so many other factors influencing fruit growth and development. Qualitative and quantitative researches are extremely necessarily combined to clarify the relevance of these factors and ripening-season of pear fruits. The past research indicated that fructose, glucose, sucrose and endogenous IAA, ABA, GAs and CTK contents in pear fruits didn't change in same regularity during fruit growth and development for different experiments. And there have no reports about the changing regularity of endogenous polyamines contents, activiety of cell wall enzymes and LOX during fruit growth and development among the different pear cultivars. Only one cultivar or factor was selected to study the mechanism of fruit growth and development in kiwifruit, strawberry, persimmon, grape, apple and some other frutis. Therefore, researches on chemical substances, such as carbohydrates, proteins, endogenous hormones and polyamines are very crucial to find the difference of growth regularity and disclose mechanism of fruit growth and development of pear fruits among the pear cultivars in different ripening-seasons. To some extent it could provide advice for fruit physiologist and field production.
Both Zaomi and WhangKeumbae pear in 5-year-old trees with uniform growing vigor were selected for this experiment. Trees were grafted on Pyrus pashia Buch-Ham rootstocks and planted in Chengdu plain. Cultural management such as fertilization and pest control were the same as those used in a commercial orchard, without fruit thinning and bagging. Characteristics of trees, photosynthetic characteristics of leaves and fruit quality were analyzed for these two pear cultivars. At the same time carbohydrates, proteins and nucleic acid contents, activities of pectinesterase (PE), polygalacturonase (PG), cellulase (Cx) and lipoxygenase (LOX) were determined along with endogenous indol-3-acetic acid (IAA), gibberellins (GA_s), zeatin (ZT), abscisin acid (ABA) and putrescine(Put), spermidine(Spd), spermine(Spm) contents during the period of fruit growth and development. The followings were the results:
1、Zaomi and Whangkeumbae pear had similar characteristics of trees in Chengdu plain. The trees grew strongly and had earlier phenophase than that in origin region. The fruits of these two cultivars were in similar shape and good in fruit quality. However, Zaomi pear fruits were sweeter than Whangkeumbae. Photosynthesis rate, chlorophyll a and the total chlorophyll contents of leaves of Zaomi pear were little higher than that of Whangkeumbae pear in the same season. Chlorophyll especially chlorophyll a was the key factor influencing photosynthesis rate for this two pear cultivars. And Zaomi pear trees had stronger tolerance of shading.
2、There were obvious differences of developmental stages of fruits between these two pear cultivars. The growth curve of fruit showed as single sigmoid and was divided into three phases. The fruit growth and development lasted 100days and 130days for Zaomi and Whangkeumbae pear, respectively. For Zaomi pear fruits, period from fruit setting to 30 days after full bloom (DAFB) was looked as the early stage, period from 30 to 70 DAFB as middle stage, and period from 70 to 100 DAFB as late stage. For Whangkeumbae pear fruits, period from fruit setting to 50DAFB was divided as the early stage, period from 50 to 100DAFB as the middle stage, period from 100 to 130DAFB as the late stage. The early stage of Whangkeumbae pear held longer for 20days than that of Zaomi pear, and the middle stage kept longer for 10days. The growth of longitudinal length and transverse diameter during fruit growth and development accorded with logistic curve for these two cultivars.
3、The contents of main chemical substances changed in different regularities during fruit growth and development for the experimental pear cultivars.
(1) Content of soluble sugar, fructose, glucose and sucrose rose gradually along with fruit growth and development for both Zaomi and Whangkeumbae pear. The fructose content was the highest at any given stage of fruit development. Glucose content was at second and higher than sucrose. The contents of three sugar components of Zaomi pear fruits were obviously higher than that of Whangkeumbae pear. Significant difference of soluble sugar contents was produced between these two cultivars at the ripening stage of fruit (p<0.01). Sugar contents increased very fast in the early stage of ovary developing and the late stage of fruit development for the two cultivars. And the latter stage was significant for accumulating sugar. Among the three sugar components fructose transformed and accumulated fastest followed with glucose and sucrose. The accumulation of fructose and glucose of Zaomi pear, and that of fructose, glucose, sucrose and soluble sugar of Whangkeumbae pear accorded with logistic curve during fruit growth and development.
(2) Protein contents kept at high level at the initial stage and decreased along with fruit development for Zaomi and Whangkeumbae pear. There were no obvious differences during the most stages between them. Zaomi pear fruits showed two peaks of content, but Whangkeumbae pear fruits changed in slow rate.
(3) For free amino acid content, it was higher in Zaomi pear fruits than that in Whangkeumbae pear fruits both at initial and late stages, and was significantly higher at fruit maturation (p<0.01). However, it was lower at the middle stage.
(4) Nucleic acid contents kept at high level at initial stage and decreased gradually along with fruit development for the two cultivars. It peaked at 10, 50 and 90 DAFB for Zaomi pear fruits and was significantly higher than that of Whangkuembae pear at fruit maturation (p<0.01). It peaked at 20DAFB for Whangkeumbae pear fruits.
4、The activity of enzymes changed in similar trend during fruit growth and development for these two pear cultivars.
(1) For the two cultivars, polygalacturonase (PG) activity increased gradually along with fruit development. It peaked at 50DAFB for Zaomi pear fruits, and peaked at 30, 90and 130DAFB for Whangkeumbae pear fruits. Zaomi pear fruits had higher activity of PG at the middle stage but had no obvious difference with Whangkeumbae pear fruits.
(2) The activity of pectinesterase (PE) kept at high level at the early stage and decreased gradually along with fruit development, but rose in slightly range before maturation for the two cultivars. It peaked at 10, 70 and 100DAFB for Zaomi pear fruits, and at 10, 80 and 130DAFB for Whangkeumbae pear fruits which had higher activity at early stage than Zaomi pear fruits, and had lower activity at the middle and late stage.
(3) The activity of cellulase (Cx) was a little low at the early stage and increased along with fruit development up to maturation followed with slightly decrease for the two cultivars. It rose sharply from 30 to 70DAFB and from 80 to 90DAFB for Zaomi pear fruits, from 20 to 30DAFB, from 50 to 70DAFB and from 80 to 120DAFB for Whangkeumbae pear fruits.
(4) The activity of lipoxygenase (LOX) was a little low at initial stage and increased along with fruit development for the two cultivars. It rose sharply from 20 to 40DAFB and from 50 to 90DAFB, but increased in slow rate for Whangkeumbae pear fruits during fruit development.
5、The difference of endogenous hormones contents both in pulp and seeds was the key factor resulting in different fruit growth and development of Zaomi and Whangkeumbae pear.
(1) In the same habitat IAA and ZT contents in pulp of Whangkeumbae pear which matured later were higher than that of Zaomi pear which matured earlier at initial stage and held in longer time. GA3 contents were higher than that of Zaomi pear fruits at the most stages except of 20DAFB. Cell division in pulp of Whangkeumbae pear was stronger and held for longer time. The three endogenous hormones IAA, GA_3 and ZT contents in pulp were higher than that of Zaomi pear at the most stages of fruit growth and development. However, peaks of contents for Whangkeumbae pear appeared late for 10 to 20days.
(2) An increase for ABA contents appeared in pulp of the two cultivars at 30DAFB. It might result in dropping of young fruits at this stage. ABA contents in pulp of Zaomi pear peaked earlier for 20days before fruit maturation. Thus Zaomi pear fruits might shift to maturate much earlier, and had shorter period of fruit growth and development.
(3) IAA and ZT contents in seeds of Whangkeumbae pear were obviously lower than that of Zaomi pear at the same stage. It might result in shifting into next developing stage for seeds, and was advantageous of being transferred into pulp for cell development. At the most stages GA_3 contents in seeds of Whankeumbae pear were higher than that of Zaomi pear at the same time. And it lasted for longer time with high contents and got to the first peak earlier for 10 days. This could indicate that Whangkeumbae pear fruits had vigorous production of GA_3 in seeds, which could be transferred into pulp to promote cell expansion. On the other hand, ABA contents in seeds of Zaomi pear peaked earlier for 30days. Therefore it could go into maturing earlier than Whangkeumbae pear.
(4) The ratio of IAA contents between pulp and seeds was less than 1 at the most stages for Zaomi pear. Therefore IAA might be mainly synthesized in seeds, especially at initial stage of fruit growth and development. For the ratio of ZT contents this two pear cultivars had similar changing trend. But Zaomi pear, which had vigorous synthesis of ZT in seeds, showed lower ratio value than Whangkeumbae pear. Few of ZT could be synthesized in pulp at the late stage for the two cultivars, but in seeds some ZT could be synthesized before maturation. The ratio of GA_3 contents was less than 1 for Zaomi pear during fruit development, so endosperm was the important place to produce endogenous GA_3.
6、Endogenous polyamines were very important for development of young fruits.
(1) This two pear cultivars contained high level of Put, Spd, Spm and free polyamines contents both in pulp and seeds at initial stage. It could stimulate vigorous cell division. Put, Spd and Spm contents in pulp of Zaomi pear were significantly higher than that of Whangkeumbae pear at 10DAFB (p<0.01). In seeds, Spd and Spm contents were significantly higher than that of Whangkeumbae pear at 20DAFB (p<0.01). It could speculate that young fruits of Zaomi pear developed more vigorous. However, the Spm contents in pulp of Whangkeumbae pear were higher at the most stages, especially it rose sharply from 10 to 20DAFB and was significantly higher than that of Zaomi pear at the same stage. So Spm contents might be very crucial for the first fast growth of young fruits of Whangkeumbae pear.
(2) Put contents both in pulp and seeds got the peak value for two times at the late stage for the two cultivars. And these two peaking values were approximate, but it appeared late for 30 to 40 days for Whangkeumbae pear. The fruit maturated late for 30days, too. This could be correlated with releasing of ethylene and starting to maturate at the late stage.
引文
[1]华南农业大学主编.果树栽培学各论(南方本,第二版).北京:中国农业出版社,1989
[2]苏艳丽,赵思东,袁德义,等.新高梨果实的生长发育特性.经济林研究2006,24(1):45-48
[3]张惠玲,陈立新.三个梨品种的枝、叶、果生长动态与生产管理.烟台果树,2005,(3):27-28
[4]曾骧主编.果树生理学.北京:北京农业大学出版社,1992
[5]吴邦良,夏春森,赵宗方编.果树开花结实生理和调控技术.上海:上海科学技术出版社,1994
[6]李学强,李作轩,李秀珍.南果梨果实外观品质的形成.果树学报,2003,20(1):27-30
[7]韦茜,李仕品,张正秋,等.美国杂交抽果实生长发育研究.广西园艺,2001,(4):3-4
[8]张秋萍.鸭梨的果实生长及离层的发生.河北师范大学学报,1983,(1):12-15
[9]于希志,徐秋萍,金锡凤.杏果实发育的研究.果树科学,1990,7(4):227-230
[10]谭余,董云波.黑穗醋栗果实生长发育规律研究.东北农学院学报,1987,18(2):119-124
[11]鄢德锐,韩德全.山楂果实生长发育规律的研究.山西果树,1989,(4):17-19
[12]李洁维,李瑞高,梁木源,等.猴桃优良株系果实生长发育规律研究.广西植物,1992,12(2):152-156
[13]卢克成,石鼎新,倪竞德.中华猕猴桃果实生长发育规律的初步研究.江苏林业科技,1999,26(4):14-16
[14]王荣花,李嘉瑞,陈理论.杏果实发育的形态解剖学研究.西北农业大学学报,2000,22(4):45-50
[15]金方伦.黔北地区两个梨品种果实生长发育规律的研究.广西园艺,2001,36(1):5-6
[16]吕金海,朱荣林,周书伟.金秋梨叶、梢和果实的生长动态及相关性.西南园艺,2006,34(1):20-21
[17]韩明玉,田玉命,张慧梅,等.秦光油桃果实生长曲线和落果波相的观察.西北植物学报,2001,21(6):1249-1253
[18]苍晶,王学东,桂明珠,等.狗枣猕猴桃果实生长发育的研究.果树学报,2002,18(2):87-90
[19]苍晶,王学东,张达,等.软枣猕猴桃果实生长发育的研究.东北农业大学学报,2004,35(1):77-83
[20]许方主编.梨树生物学.北京:科学出版社,1992
[21]Goffinet M C,Robinson T L,Lakso A N.A comparison of 'Empire' apple fruit size and anatomy in unthinned and handthinned trees.J.Hort Sci,1984,70(3):375-378
[22]曹亚波,周亚娣,许善华,等.早熟优质梨新品种-早蜜梨.宁波农业科技,2003,(2):30-31
[23]文仁德,常运涛.早熟梨新品系-“早蜜”.广西园艺,2000,(4):19-20
[24]刘炳刚.中晚熟梨品种黄金梨引种初报.中国果树,2001,(1):13-14
[25]刘长根,余国清.黄金梨生物学特性及栽培技术要点.中国南方果树,2004,33(5):89-90
[26]刘遵春.金花梨变异单系生物学特性调查比较及RAPD多态性分析.雅安:四川农业大学硕士学位论文,2004
[27]裴秦青,张国忠.鸭梨采收期与贮藏场所的研究.河北农业大学学报,1989,12(1):46-53
[28]林真二著(吴耕民译).梨.北京:农业出版社,1981
[29]梁东,马锋旺,管清美,等.蔷薇科植物中山梨醇代谢酶的研究进展.西北植物学报,2004,24(7):1362-1366
[30]胡庆祥,郡荣庭,张玉星,等.鸭梨果实发育中糖酸变化特点及配方施肥效应.河北果树,1996(1):9-11
[31]李春英,金英善,李永勋.苹果梨果实生长期间山梨糖醇及其它糖的变化规律.延边大学农学学报,2001,23(3):198-201
[32]李春英,薛桂新,李永勋.客发梨果实生长期间山梨糖醇及其它糖变化规律的研究.北方果树,1999(6):6-7,26
[33]王军虹,闰丽梅,刘武林.大香水梨成熟过程生理生化反应.北方园艺,1998,(6):24-25
[34]敖良德,王明鑫.苹果成熟过程中果实内单宁酸、苹果酸、维生素C及淀粉含量的变化.中国果树,1988,(2):35-36
[35]邓月娥,张传来,牛立元,等.桃果实发育过程中主要营养成分的动态变化及系统分析方法研究.果树科学,1998,15(1):48-52.
[36]甘霖,谢永红,吴正琴,等.‘嘉平大枣'果实发育过程中糖、酸及维生素C含量的变化.园艺学报,2000,27(5):317-320
[37]王军虹,闰丽梅,刘武林.苹果梨采前生理生化变化.东北农业大学学报,1998,29(4):388-392
[38]申曙光,马宝焜,陈四维.红富士苹果果实发育期间生理生化变化的研究.园艺学报,1991,18(1):1-3
[39]Vizzoto G,Pinto R,Varanini Z.Sucrose accumulation in developing peach fruit.Plant Physiol,1996,96:225-230
[40]Davis C,Robinson SP.Sugar accumulation in grape berries.Plant Physiol,1996,111:275-283
[41]吕英民,张大鹏.果实发育过程中糖的积累.植物生理学通讯,2000,36(3):258-265
[42]陈俊伟,张良诚,张上隆.果实中的糖分积累机理.植物生理学通讯,2000,36(6):497-503
[43]陈俊伟,张上隆,张良诚.果实中糖的运输、代谢与积累及其调控.植物生理与分子生物学学报,2004,30(1):1-10
[44]陈俊伟,谢鸣,秦巧平.植物糖信号与激素信号之间的联系.植物生理学通讯,2005,41(3):279-285
[45]谢国禄.苹果和日本梨果实发育期间果糖含量和果糖相关酶活性.国外作物育种,2001,70(1): 16-20
[46]Beruter J,Studer Feusi ME,Ruedi P.Sorbitol and sucrose partitioning in the growing apple fruit.J Plant Physiol,1997,151:269-276
[47]Beruter J,Studer Feusi ME.Comparison of sorbitol transport in excised tissue discs and cortex tissue of intact apple fruit.Plant Physiol,1995,146:95-102
[48]Beruter J,Studer Feusi ME.The effect of girdling on carbohydrate partitioning in the growing apple fruit.J Plant Physiol,1997,151:277-285
[49]Yamaguchi H,Kanayama Y,Soejima J,etal.Changes in the a mounts of the NAD dependent sorbitol dehydrogenase and its involvement in the development of apple fruit.J Amer Soc Hort Sci,1996,121(5):848-852
[50]单守明,王永章,董晓颖,等.IAA、GA和ABA对苹果果实山梨醇代谢相关酶活性的影响.园艺学报,2005,32(6):990-993
[51]Davis C,Robinson SP.Sugar accumulation in grape berries.Plant Physiol,1996,111:275-283
[52]Farrar JF.Sink strength:What is it and how do we measure it?(Forum) Plant Cell and Enviorn,1993,16:1014-1046
[53]Hirano K,Yone mori K,Sugiura A.Involvement of sugar metabolism in persimmon growth inhibition by calyx lobe removal.J Amer Soc Hort Sci,1996,121(3):456-472
[54]Moriguchi T,Yamaki S.Purification and characterization of sucrose synthase from peach(Prunus persica) fruit.Plant Cell Physiol,1988,29(8):1361-1366
[55]Sun J.Sucrose synathase in wild tomato,Lycopersicon chmielewskii,and tomato fruit sink strength.Plant Physiol,1992,98:1163-1169
[56]Sung SS,Sheih WJ,Geiger DR,etal.Growth,sucrose synthase,and invertase activities of developing Phaseolus vulgaris L.fruit.Plant Cell and Enviorn,1994,17:419-426
[57]Yamaguchi H,Kanayama Y,Yamaki S.Purification and properties of NAD dependent sorbitol dehydrogenase from apple fruit.Plant Cell Physiol,1994,35(6):887-892
[58]Yamaki S,Ishikawa K.Roles of four sorbitol related enzymes and invertase in the seasonal alteration of sugar metabolism in apple tissue.J Amer Soc Hort Sci,1986,111(1):134-137
[59]辛贺明,张喜焕.套袋对鸭梨果实内含物变化及内源激素水平的影响.果树学报,2003,20(3):233-235
[60]程和禾.黄金梨果实生长发育及贮藏生理特性研究.保定:河北农业大学硕士学位论文,2004
[61]李金昶,石晶,苏忠民,等.苹果梨成熟和贮藏过程中有机酸含量的变化.植物生理学通讯,1995,31(2):118
[62]林敏娟.黄金梨叶片、果实内营养物质含量变化及其相关关系研究.保定:河北农业大学硕士学位论文,2005
[63]潘瑞炽主编.植物生理学.北京:高等教育出版社(第三版),1995
[64]林敏娟,徐继忠,陈海江,等.黄金梨叶片、果实中矿质元素含量的周年变化动态.河北农业大学学报,2005,28(6):23-27
[65]刘国荣.不同矮化中间砧红富士苹果果实生长及其内含物含量变化的研究.保定:河北农业大学硕士学位论文,2003
[66]李树玲,黄礼森,丛佩华。等.不同种内梨品种果实维生素C含量.园艺学报,1994,21(1):17-20
[67]吕金海,伍贤进,周书伟,等.金秋梨果实发育中生长速率及主要营养成分的变化.中国农学通报,2000,20(5):35-36,44
[68]刘剑锋,张红艳,彭抒昂.梨果实发育中果肉及种子钙和果胶含量的变化.园艺学报2003,30(6):709-711
[69]刘利德,姚敦义.植物激素的概念及其新成员.生物学通报,2002,37(8):18-20
[70]李玉珍.植物激素的研究进展.河南科技学院学报(自然科学版),2007,35(4):50-53
[71]刘剑锋,程云清,彭抒昂.梨果肉与种子中钙与内源激素含量变化关系研究.植物营养与肥料学报,2005,11(2):269-272
[72]阮晓,王强,周疆明,等.香梨果实成熟衰老过程中4种内源激素的变化.植物生理学报,2000,26(5):402-406
[73]饶景萍,平田尚美.猕猴桃果实生长发育过程中内源植物激素的变化动态.西北农业大学学报,1992,20(4):41-44
[74]陶汉之,高丽萍,陈佩璁,等.猕猴桃果实发育中内源激素水平变化的研究.园艺学报,1994,21(1):35-40
[75]肖家欣,彭抒昂,何华平.单性结实和自花结实柑橘果实发育过程中IAA、ZR和GA3含量的变化.武汉植物学研究,2005,23(3):272-275
[76]李秀菊,刘用生,束怀瑞.不同成熟型苹果果实生长发育过程中几种内源植物激素含量变化.植物生理学通讯,2000,36(1):7-10
[77]彭福田,姜远茂,顾曼如,等.氮素对苹果果实内源激素变化动态与发育进程的影响.植物营养与肥料学报,2003,9(2):208-213
[78]钟晓红,马定渭,黄远飞.草莓果实发育过程中内源激素水平的变化.江西农业大学学报,2004,26(1):107-111
[79]周碧燕,季作梁,叶永昌,等.荔枝果实发育期间内源激素含量的变化.园艺学报,1998,25(3):236-240
[80]周碧燕,李建国,黄旭明,等.龙眼果实发育过程中ZRs、GA和IAA含量的变化.华南农业大学学报,1999,20(3):50-53
[81]王碧青,邱燕萍,向旭,等.荔枝结果过程中内源激素变化及单性结果的诱导.园艺学报,1997,24(1):19-24
[82]季作梁,梁立峰,柳建良,等.荔枝果实发育期间内源激素含量动态.华南农业大学学报,1992,13(3):93-98
[83]樊卫国,安华明,刘国琴,等.刺梨果实与种子内源激素含量变化及其与果实发育的关系.中国农业科学,2004,37(5):728-733
[84]陈发河,蔡慧农,冯作山,等.葡萄浆果发育过程中激素水平的变化.植物生理与分子生物学学报,2002,28(5):391-395
[85]Luckwill L C.Studies of fruit development in relation to plant hormone.Ⅳ.Alidic auxins and growth inhibitors in leaves and fruit of the apple.The Journal of Horticultural Science,1957(32):18-33
[86]丁长奎,章恢志.植物激素对枇杷果实生长发育的影响.园艺学报,1988,15(3):148-154
[87]Hopping M.E.,Bukoval M.J..Endogenous plant growth substances in developing fruit of Prunus cerasus L.Ⅵ.Extractable auxin in the seed and pericarp.The Journal of Horticultural Science.1975,100(40):399-401
[88]王忠主编.植物生理学.北京:中国农业出版社,2000
[89]吕忠恕著.果树生理.上海:上海科学技术出版社,1982
[90]郗荣庭主编.果树栽培学总论(第三版).北京:中国农业出版社,2004
[91]杨立峰,刘弦,郝风鸽.GA_3对苹果生长发育的影响研究.河北果树,2004,(5):8-10
[92]John E.Jackson.Biology of apples and pears.UK:Cambridge University Press,2003
[93]吴少华,沈德绪,林伯年,等.梨果实发育和直感同内源激素的关系.浙江农业大学学报,1986,12(1):57-61
[94]钟晓红,马定渭,黄远飞.草莓果实发育过程中内源激素水平的变化.江西农业大学学报,2004,26(1):107-111
[95]吴俊,钟家煌,徐凯,等.外源GA_3对藤稔葡萄果实生长发育及内源激素水平的影响.果树学报2001,18(4):209-212
[96]陈大成,李志勇,胡桂兵.‘塔1'椪柑果实发育、内源激素变化及品质形成特点.华南农业大学学报,1998,19(3):51-55
[97]Kiyohide Kojima,Keishi Shiozaki.Changes of endogeneous levels of ABA,IAA and GA-like substances in fruitlets of parthenocarpic persimmon.Japan Soc Hort Sci,1999,68(2):242-247
[98]张崇浩,袁玉信,李杰芬,等.桃果实后熟中内源激素的变化.北京师范大学学报(自然科学版),1989,(2):68-72
[99]Vanoli M,Marro M,Visai C.Gibberellin-like substances during the development of peach fruits.Acta Horticulturae,1993,329:159-16
[100]Cutting J G M,Lyne M C.Girdling and the reduction in shoot xylem sap concentrations Of cytokinins and gibberellins in peach.Journal of Horticultural Science,1993,68(4):619-626
[101]Dennis F G Jr.Trial of ethephon and other growth regulation for delaying bloom in tree fruits.Journal of the American Society for Horticultural Science,1976,101(3):241-245
[102]刘小阳,李玲,蔡永萍.光强对砀山酥梨石细胞形成的影响及其与内源IAA、ZR和ABA含量的关系.激光生物学报,2006,15(2):161-166
[103]邹养军,王永熙.内源激素对苹果果实生长发育的调控作用研究进展.陕西农业科学,2002(10):13-15
[104]闫树堂,徐继忠.不同矮化中间砧对红富士苹果果实内源激素、多胺与细胞分裂的影响.园艺学报,2005,32(1):81-83
[105]闫国华,甘立军,孙瑞红.赤霉素和细胞分裂素调控苹果果实早期生长发育机理的研究.园艺学报,2000,27(1):11-16
[106]季作梁,王钢涛.荔枝果实发育过程中细胞分裂素的变化.果树科学,1996,13(2):92-95
[107]李建国,周碧燕.大核和焦核“桂味”荔枝果实发育及其发育期间果皮中内源激素含量的变化比较.植物生理学通讯,2005,41(5):587-590
[108]张海娥,郭修武.激素在葡萄分子发育调控上的作用研究进展.中外葡萄与葡萄酒,2005,(2):28-29,32
[109]Salopek-Sondi B,Kovac M,Prebeg T,et al.Developing fruit direct post-floral morphogenesis in Helleborus fruit direct post-floral morphogenesis in Helleborus niger L.J Exp Bot.2002,53(376):1949-1957
[110]周蕾,魏琦超,高峰.细胞分裂素在果实及种子发育中的作用.植物生理学通讯,2006,42(3):549-553
[111]彭良志,胥耳.6-BA和GA_3对华盛顿脐橙幼果~(14)C光合产物调配的影响.园艺学报,1990,17(2):111-116
[112]倪竹如,陈俊伟,阮美颖,等.BA对椪柑果实生长发育及其同化产物分配的影响.浙江农业学报,2000,12(5):272-276
[113]Stern RA,Shargal A,Flaishman MA.Thidiazuron increases fruit size of 'Spadona'and 'Coscia' pear(Pyrus communis L.).J Hortic Sci Biotechnol,2003,78(1):51-55
[114]Sotta B,Sossountzov L,Maldiney R,etal.Abscisic acid localization by light microscopic immunohistochemistry in chenopodium polyspermum L.J Histochem Cytochem,1985,33(3):201-208
[115]Sossountzov L,Sotta B,Maldiney R,etal.Immunoelectron-microscopy Localization of abscisic acid with colloidal gold on Lowicryl-embedded tissues of chenopodium polyspermum L.Planta,1986,168:471-481
[116]Bertrand S,Benhamou N,Nadeau P,etal.Immunogold localization of free abscisic acid in tomato root cells.Canadian J Bot,1992,70(5):1001-1011
[117]贾文锁,王学臣,娄成后.蚕豆(Vicia faba L.)叶肉细胞中ABA的胶体金免疫电镜定位.植物生理学报,1994,20(4):380-384
[118]王学臣,贾文锁.水分胁迫下蚕豆(Vicia faba L.)气孔关闭与叶片细胞中ABA区隔化与再分配的关系.植物生理学报,1995,21(4):324-328
[119]Pastor A,Cortadellas N,Alegre L.Immunolocalization of abscisic acid by monoclonal antibodies in Lavandula stoechas L.leaves.Plant Growth Regulation,1995,16:287-292
[120]黄丛林,张大鹏,贾文锁.葡萄种子细胞中脱落酸的胶体金免疫电镜定位.中国农大大学学报,1999,4(5):111-117
[121]黄丛林,贾文锁,张大鹏.葡萄果肉细胞中脱落酸的区隔化研究.植物生理学报,2000,26(4):297-300
[122]Zhang D P,Huang C L,Jia W S.Immunogold electron-microscopy localization of abscisic acid in fresh cells of grape berry(Vitis vinifera L.×Vitis labrusca L.cv Kyoho).Sci Hort,1999,81:189-198
[123]刘勇,刘善军,霍光华,等.甜柿果实发育期间矿质元素与营养成分的变化.江西农业大学学报,2000,22(2):265-270
[124]郑国华,杉浦明,米森敬三.柿果(Diospyros Kaki L.)发育及成熟过程中内源GAs活性及ABA含量的变化.北京农业大学学报,1991,17(1):77-81
[125]郑国华,平野建.喷施GA_3和乙烯利对柿果实成熟及内源GAs活性,ABA含量的影响.园艺学报,1991,18(3):193-197
[126]张大鹏,张子连,陈珈.葡萄果实发育过程中脱落酸结合蛋白动力学特性的变化.植物学报,1996,38(12):930-935
[127]Kojima K,Kuraishi S,Sakurai N.Distribution of ABA in different parts of the reproductive organs of tomato.Sci Hort,1993,56:23-30
[128]I.Lara,M.Vendrell.Changes in abscisic acid levels,ethylene biosynthesis,and protein patterns during fruit maturation of 'Granny Smith' aplles.J.Amer.Soc.Hort.Sci,2000,125(2):183-189
[129]Satoru Kondo,Hiroshi Gemma.Relationship between abscisic acid(ABA) content and maturation of the sweet cherry.J.Japan.Soc.Hort.Sci,1993,62(1):63-68
[130]黄丛林,张大鹏,贾文锁.葡萄果实发育后期脱落酸来源的研究.园艺学报,2001,28(5):385-391
[131]Yuan R C,Huang H B.Litchi fruit abscission:its pattern,effects of Shading and relation to abscisic acid.Scientia Horticulturae,1988,36(3-4):281-292
[132]Coombe B G.The grape berry as a sink.Acta Hort,1989,(239):149-158
[133]Archbold D D.Abscisic acid facilitates sucrose import by strawberry fruit explants and cortex disks in vitro.HortSci,1988,23:880-881
[134]Doucette M S,Pritchard M K,Saltveit M E.ABA involvement in low temperature sweetening of potatoes.Acta Hort,1993,(343):293-294
[135]Kojima K,Yamada Y,Yamamoto M.Distribution of ABA and IAA with a developing valencia orange fruit and its parts.J Jpn Soc Hort Sci,1994,63:335-339
[136]Smith G S,Klages K V,Green T G A,etal.Changes in abscisic acid concentration,surface conductance and water content of developing kiwifruit.Sci Hort,1995,61(1-2):13-27
[137]吴有梅,顾采琴.ABA和乙烯在草莓采后成熟过程中的作用.植物生理学报,1992,18(2):167-172
[138]Leung J,Giraudat J.Abscisic acid signal transduction.Annu Rev Plant Physiol Plant Mol Biol,1998,49:199-222
[139]Dik Hagenbeek,Ralph S.Quatrano,Christopher D.Rock,et al.Trivalent ions activate abscisic acid-inducible promoters through an ABI1-dependent pathway in rice protoplasts.Plant Physiol,2000,123:1553-1560
[140]阮晓,王强,王文全,等.香梨果实发育、成熟期间乙烯、ACC、MACC变化及相互关系.园艺学报,2000,27(3):205-206
[141]韦军,田边贤二.鸭梨、茌梨果实采前乙烯生物合成特性研究.园艺学报,1994,21(3):231-234
[142]关军锋,束怀瑞,黄天栋.新红星苹果果实细胞膨大期及成熟期乙烯生成的变化.中国农业科学,1993,26(5):57-60
[143]李全梓,李兴国,郑国生,等.早熟桃果实和种子发育过程中乙烯产生及ACC含量和ACC氧化酶活性的变化.植物生理学通讯,1998,34(1):25-27
[144]王蓉花,李嘉瑞,张娟,等.杏果实生长发育生理学研究.西北农业学报,2000,9(3):76-78
[145]李全梓,李兴国,张宪省,等.新红星苹果发育过程中果实和种子乙烯生物合成的研究.山东农业大学学报,1998,29(2):151-155
[146]李正国,El-Sharkawy I,Lelievre J-M.温度、丙烯和1—MCP对西洋梨果实乙烯合成和乙烯受体ETR1同源基因表达的影响.园艺学报,2000,27(5):313-316
[147]Li Zheng-Guo,El-Sharkawy,Bouzayen.Effects of cold treatment on ethylene biosynthesis and responsiveness during ripening of Pear(Pyrus communis L.)fruits.植物生理学报,2000,26(4):283-288
[148]Lelievre J M,Tichit L,Dao P.Effects of chilling on the expression of ethylene biosynthesis genes in Passe-Crassane pear fruits.Plant Mol.Biol.,1997,101:727-739
[149]魏绍冲,陈昆松,罗云波.乙烯受体与果实成熟调控.园艺学报,2004,31(4):543-548
[150]周俊义.多胺在果树生理的作用.果树科学,1995,12(2):117-119
[151]崔凤芝,张运涛.多胺与园艺植物生长发育的关系.河北农业大学学报,1996,19(3):94-98
[152]杨洪强,接玉玲.多胺与果树生长发育的关系.山东农业大学学报,1995,27(40):514-520
[153]Bernet E,Claparolsa I,Dondini L,et al.Changes in polyamine content,arginine and ornithine decarboxylases and ransglutaminase activities during light dark phases(of initial differentiation)in maize calluses and their chloroplasts.Plant Physiol Biochem,1999,37(12):899-909
[154]Shoeb F,Yadav JS,Bajaj LS,etal.Polyamines biomarkers for plant regeneration capacity:improvement of regeneration by modulation of polyamine metabolism in different genotypes of indica rice.Plant Seience,2001,160:1229-1235
[155]Omar franco-Mora,Kenji Tanabe,Akihiro Itai,etal.Relationship between endogenous free polyamines content and ethylene evolution during fruit growth and ripening of Japanese Pear(Pyrus pyrifolia Nakai).J.Japan.Soc.Hort.Sci,2005,74(3):221-227
[156]王立英.柿(DiospyroskakiL.)果实生长发育及成熟机理研究.保定:河北农业大学硕士学位论文,2002
[157]Biasi R,Bagni N.Endogenous polyamines in apple and their relationship to fruit set and fruit growth.Physiol Plant,1988,(73):201-205
[158]王世平,宋长冰,李连朝,等.三种多胺在苹果开花及坐果初期的生理作用.园艺学报,1996,23(4):319-325
[159]陈海江,徐继忠,袁小乱,等.不同时期喷施外源多胺对新红星苹果坐果的影响.河北农业大学学报,1999,22(4):47-49
[160]Aziz A.Spermidine and related-metabolic inhibitors modulate sugar and amino acid levels in Vitis vinifera L:possible relationships with initial fruitlet abscission.J Exp Bot,2003,54:355-363
[161]徐继忠,史常青,马宝焜,等.外源多胺对苹果花粉管伸长及胚珠寿命的影响.园艺学报,1998,25:91-92
[162]钟晓红.三种多胺在沙田柚开花及坐果初期的生理作用.湖南农业大学学报(自然科学版),2000,26(6):453-456
[163]Koshgad M M,Orvos A R,Yelenosky G.Relative change in polyamines during citrus flower development.HortScience,1990,(25):946-948
[164]郑玉生,张秋明,Lovoctt C J.柑橘花期代谢及其对坐果的影响.果树科学,1997,14(4):216-219
[165]Malik AU,Singh Z.Abscission of mango fruitlets as influenced by biosynthesis of polyamines.Hort Sci Biotechnol,2003,78(5):721-727
[166]Wolukaul JN,Zhang SL,Xu GH,et al.The effect of temperature,polyamines and polyamine synthesis inhibitor on in vitro pollen germination and pollen tube growth of Prunus mume.Sci Hort,2004,99:289-299
[167]Shiozaki S,T.Ogata,S.Horiuchi.Endogenous polyamines in the pericarp and seed of the grape berry during development and ripening.Scientia Hortic,2000,83:33-41
[168]陈杰忠,骆小兵,王泽槐,等.多胺对荔枝果实坐果率及内源多胺含量的影响.园艺学进展(第五辑)2000:303-307
[169]李建国,刘顺枝,王泽槐.荔枝果实发育过程中内源多胺含量的变化.植物生理学通讯,2004,40(2):153-156
[170]李建勇,卢钢,任彦.多胺在果实生长发育中的作用研究进展.果树学报,2005,22(3):256-260
[171]Lester GE.Polyamines and their cellular anti-senescence properties in honey dew muskmelon fruit.Plant Science,2000,160:105-112
[172]陈昆松,张上隆.脂氧合酶与果实的成熟衰老.园艺学报,1998,25(4):338-344
[173]曹慧,王永章,杜俊杰,等.脂氧合酶研究进展(综述).安徽农业大学学报,2001,28(2):219-222
[174]陈晶,李正国.脂氧合酶在果实成熟衰老中的作用.广州食品工业科技,2002,18(1):50-53
[175]张荣平.脂氧合酶在植物体内的生理功能.莱阳农学院学报,1993,10(1):47-51
[176]许文平,陈昆松,徐昌杰,等.猕猴桃采后果实冷藏与货架期脂氧合酶活性和乙烯生成的变化.中国农业科学,2003,36(10):1196-1201
[177]许文平,陈昆松,李方,等.脂氧合酶、茉莉酸和水杨酸对猕猴桃果实后熟软化进程中乙烯生物合成的调控.植物生理学报,2000,26(6):507-514
[178]罗云波.脂氧合酶与番茄采后成熟的关系.园艺学报,1994,21(4):357-360
[179]Riley J C M,Willemot C,Thompson J E.Lipoxygenase and hydroperoxide lyase activities in ripening tomato fruit.Postharvest Biology and Technology,1996,7:97-107
[180]Marcelle RD.Relationship between mineral content,lipoxygenase activity,levels of 1-aminocyclopropane-1-carboxylic acid and ethylene emission in apple fruit flesh disks(cv.Jonagold)during storage.Postharvest Biology and Technology,1991,(1):101-109
[181]吴敏,陈昆松,张上隆.桃果实采后成熟过程中脂氧合酶活性变化.园艺学 报,1999,26(4):227-231
[182]谢永红,陈学年,吕斌,等.桃果实采后脂氧合酶活性与内源乙烯对果实软化衰老的影响.西南园艺,2004,32(6):1-3
[183]Todd J F,Paliyath G,Thompson J E.Characteristics of a membrane-associated lipoxygenase in tomato fruit.Plant Physiol,1990,94:1225-1232
[184]韩碧文主编.植物生长与分化.北京:中国农业大学出版社,2003
[185]寇晓虹,罗云波.植物多聚半乳糖醛酸酶研究进展.山西农业大学学报,2004,24(2):411-415
[186]彭丽桃,饶景萍,杨书珍,等.果实软化的胞壁物质和水解酶变化.热带亚热带植物学报,2002,10(3):271-280
[187]赵云峰,林河通,林娇芬,等.果实软化的细胞壁降解酶及其调控研究进展.仲恺农业技术学院学报,2006,19(1):65-70
[188]薛炳烨,束怀瑞.肥城桃两品系果实细胞壁成分和水解酶活性的比较.园艺学报,2004,31(4):499-501
[189]曾秀丽,张光伦,李春燕,等.三个脐橙品种果实主要细胞壁酶动态变化研究.亚热带植物科学,2006,35(2):12-16
[190]李春燕,张光伦,曾秀丽,等.细胞壁酶活性与甜橙果实质地的相关性研究.四川农业大学学报,2006,24(1):73-76
[191]李建国,黄旭明,黄辉白.裂果易发性不同的荔枝品种果皮中细胞壁代谢酶活性的比较.植物生理与分子生物学学报,2003,29(2):141-146
[192]薛炳烨,毛志泉,束怀瑞.草莓果实发育成熟过程中糖苷酶和纤维素酶活性及细胞壁组成变化.植物生理与分子生物学学报,2006,32(3):363-368
[193]张嵩,张光伦,曾秀丽.纤维素酶和多聚半乳糖醛酸酶与果实成熟.果树学报,2005,22(5):532-536
[194]寇晓虹,罗云波,田慧琴,等.多聚半乳糖醛酸酶(PG)反义基因转化加工番茄.食品科学,2007,28(3):187-191
[195]李雄彪.植物细胞壁酶的分子结构与生理功能.植物生理通讯,1991,27(4):246-252
[196]张昭其,段学武,陆旺金.果实多聚半乳糖醛酸酶分子生物学研究进展.生命科学,2002,14(2):92-94
[197]薛炳烨,束怀瑞.应用基因工程改良果实成熟特性研究进展.山东农业科学,2002,(4):49-52
[198]Koch K E.Carbohydrate-modulated gene expression in plants.Annu Rev Plant Physiol Plant Mol Biol,1996,47:509-540
[199]Leon P,Sheen J.Sugar and hormone connection.Trends Plant Sci,2003,8:110-116
[200]Sheen J,Zhou L,JiangLC.Sugars as signaling molecules.Curr Opin Plant Biol,1999,2:410-418
[201]Smeekens S.Sugar-induced signal transduction in plants.Annu Rev Plant Physiol Plant Mol Biol,2000,51:49-81
[202]Leon P,Sheen J.Sugar and hormone connection.Trends Plant Sci,2003,8:110-116
[203]Rolland F,Moore B,Sheen J.Sugar sensing and signaling in plants.Plant Cell,2002,14:185-205
[204]Stadler R,Truernit E,Gahrtz M,etal.The AtSUC1 sucrose carrier may represent the osmotic driving force for anther dehiscence and pollen tube growth in Arabidopsis.Plant J,1999,19:269-278
[205]Bethke P C,Schuurink P C,Jones R L.normonal signaling in cereal alcurone.J Exp Bot,1997,48:1337-1356
[206]吴有梅,刘愚.番茄果实成熟过程中钙调索含量变化及其与乙烯合成的关系.植物学报,1990,16:709-713
[207]关军锋,MaX Saure编著.果树钙素营养与生理.北京:科学出版社,2005
[208]罗充,彭抒昂,李国怀.梨不同品种果实发育中钙与钙调素的动态研究.华中农业大学学报,2001,20(3):286-288
[209]张红艳,彭抒昂.梨果实发育过程中钙调素水平的动态研究.华中农业大学学报,2004,23(3):338-340
[210]关军锋,马智宏,张会敏,等.苹果幼果发育期果柄结构和Ca、Mg、K水平的变化.植物营养与肥料学报,2005,11(1):264-268
[211]蒲富慎,黄礼森,孙秉钧,等编.梨品种.北京:农业出版社,1989
[212]华中农业大学主编.果树研究法(第二版).北京:农业出版社,1979
[213]骆建霞,孙建设主编.园艺植物科学研究导论.北京:中国农业出版社,2001
[214]束怀瑞主编.果树栽培生理学.北京:中国农业出版社,1993
[215]李锡香编著.新鲜果蔬的品质及其分析法.北京:中国农业出版社,1994
[216]中国科学院上海植物生理研究所编.现代植物生理学实验指南.北京:科学出版社,1999
[217]张友杰.以蒽酮分光光度法测定果蔬中.的葡萄糖、果糖、蔗糖和淀粉.分析化学,1980,5(3):167-171
[218]李合生主编.植物生理生化实验原理和技术.北京:高等教育出版社,2000
[219]朱广廉,钟海文,张爱琴编.植物生理学实验.北京:北京大学出版社,1990
[220]Surrey K.Spectrophotometric method for determination of lipoxidase activity.Plant physiol.1963.39:65-70
[221]徐昌杰,陈昆松,张上隆.蔗糖酶对柑桔外切纤维素酶和外切多聚半乳糖醛酸酶活性测定的干扰及排除.植物生理通讯,1997,33(1):43-46
[222]宁正祥.食品成分分析手册.北京:中国轻工业出版社,1998
[223]张飞,岳田利,费坚,等.果胶酶活力的测定方法研究.西北农业学报,2004,13(4):134-137
[224]田志喜,张玉星,于艳军,等.水杨酸对鸭梨果实PG、PME和呼吸速率的影响.果树学报,2002,19(6):381-384
[225]上海植物生理学会编.植物生理实验指导手册.上海:上海科学技术出版社,1985
[226]丁静,沈镇德,方亦雄,等.植物内源激素的提取分离和生物鉴定.植物生理学通讯,1979,2:27-29
[227]陈昆松,徐昌杰,李方,等.HPLC法检测果实组织中内源IAA、ABA方法的改进.果树学报,2003,20(1):4-7
[228]陈远平,杨文钰.卵叶韭休眠芽中GA3、IAA、ABA和ZT的高效液相色谱法测定.四川农业大学学报,2005,23(4):498-500
[229]曾庆钱,陈厚彬,鲁才浩,等.HPLC测定荔枝不同器官中内源激素流程的优化.果树学报,2006,23(1):145-148
[230]Hector E.Flores,Arthur W.Galston.Analysis of polyamines in high plants by high performance liquid chromatography.Plant Physiol.,1982,69:701-706
[231]刘俊,吉晓佳,刘友良.检测植物组织中多胺含量的高效液相色谱法.植物生理学通讯,2002,38(6):596-598
[232]Liu Ji-Hong,Kazuyoushi Nada,Pang Xiao-ming,et al.Role of polyamines in peach fruit development and storage.Tree Physiology,2006,26:791-798
[233]盛宝龙.不同梨品种光合特性及影响因素研究.南京:南京农业大学硕士学位论文,2004
[234]郭春爱,刘芳,许晓明.叶绿素b缺失与植物的光合作用.植物生理学通讯,2006,42(5):967-973
[235]汪良驹,姜卫兵,高光林,等.幼年梨树品种光合作用的研究.园艺学报,2005,32(4):571-577
[236]Caixi Zhang,Kenji Tanabe,Shiping Wang,etal.The impact of cell division and cell enlargement on the evolution of fruit size in Pyrus pyrifolia.Annals of Botany,2006,98:537-543
[237]王贵元,夏仁学,吴强盛.果实中糖分的积累与代谢研究进展.北方园艺,2007(3):56-58
[238]胡红菊,陈启亮,王友平,等.4个砂梨品种果实发育过程中主要糖酸含量的变化.华中农业大学学报,2007,26(2):251-255
[239]赵智中.柑橘果实糖积累的生理基础研究.杭州:浙江大学博士学位论文,2001
[240]龚荣高,张光伦,吕秀兰,等.脐橙在不同生境下果实蔗糖代谢相关酶的研究.园艺学报,2004,31(6):719-722
[241]刘永忠,李道高.柑橘果实糖积累与蔗糖代谢酶活性的研究.园艺学报,2003,30(4):457-459
[242]李培环,董晓颖,王永章,等.‘新红星'苹果果实蔗糖合酶的活性及亚细胞定位.园艺学报,2002,29(4):375-37;
[243]夏国海,张大鹏,贾文锁,等.IAA、GA和ABA对葡萄果实14C蔗糖输入与代谢的调控。园艺学 报,2000,27:6-10
[244]Archbold R C.Abscisic acid facilitates sucrose import by strawberry fruit explants and cortex disks in vitro.Hortic Sci,1988,23:880-883
[245]Fischer R.L.,Bennett A.B.Role of cell wall hydrolases in fruit ripening.Annu Rev Plant Physiology Plant Molecular Biology,1991,142:675-703
[246]刘剑锋,程云清,彭抒昂.梨采后细胞壁成分及果胶酶活性与果肉质地的关系.园艺学报,2004,31(5):579-583
[247]Bonghi,C.,Ferrarese,L..Endo-b-1,4-glucanases are involved in peach fruit growth and ripening,and regulated by ethylene.Physiologia Plantarum,1998,102:346-352
[248]Siedow JN.Plant lipoxygenease:structure and function.Annu.Rev.Plant MOL.Biol.1991,42:145-188
[249]Bousquest J F,Thimann K V.Lipid peroxidation forms ethylene from ACC and may operate in leaf senescence.Proc.Natn.Acad.Sci.USA,1984,81:1724-1727
[250]黄卫东,原永兵,彭宜本.温带果树结实生理.北京:北京农业大学出版社,1994
[251]Lieberman N,Bake J E,Sloge.Influence of plant hormones ethylene production in apple,tomato and avocado slices during maturation and senescence.Plant Physiol,1977,60:214-217
[252]张大鹏,许雪峰,张子连,等.果实始熟机理.园艺学报,1997,24(1):1-7
[253]李六林,张绍玲,郭艳玲.‘新高'梨花粉败育与内源多胺含量变化的关系.园艺学报,2007,34(2):301-304
[254]郭志雄.油柰坐果及果实生长发育机制的若干研究.福州:福建农业大学硕士学位论文,1999
[255]Ke D and Romani R J.1988.Effects of spermidine on ethylene production and the senescence of suspension-cultured pear fruit cells.Plant Physiol.Biochem.,26:109-116
[256]Martin Tanguy J.1997.Conjugated polyamines and reproductive development:biochemical,molecular and physiological approaches.Physiol.Plant.,100:675-688
[257]Minocha R,Lee J S,Long S,Bhatnagar P,Minocha S C.2004.Physiological responses of wild type and putrescine-overproducing transgenic cells of polar to variations in the fom and concentration of nitrogen in the medium.Tree Physiology,24:551-560
[258]Rastogi R,Davies P J.1991.Effects of light and plant growth regulators on polyamine metabolism in higher plants.In:Slocum,R.D.,Flores,H.E.(Eds.),Biochemistry and Physiology of Polyamines in Plants.CRC Press,Boca Raton,187-199
[2]苏艳丽,赵思东,袁德义,等.新高梨果实的生长发育特性.经济林研究2006,24(1):45-48
[3]张惠玲,陈立新.三个梨品种的枝、叶、果生长动态与生产管理.烟台果树,2005,(3):27-28
[4]曾骧主编.果树生理学.北京:北京农业大学出版社,1992
[5]吴邦良,夏春森,赵宗方编.果树开花结实生理和调控技术.上海:上海科学技术出版社,1994
[6]李学强,李作轩,李秀珍.南果梨果实外观品质的形成.果树学报,2003,20(1):27-30
[7]韦茜,李仕品,张正秋,等.美国杂交抽果实生长发育研究.广西园艺,2001,(4):3-4
[8]张秋萍.鸭梨的果实生长及离层的发生.河北师范大学学报,1983,(1):12-15
[9]于希志,徐秋萍,金锡凤.杏果实发育的研究.果树科学,1990,7(4):227-230
[10]谭余,董云波.黑穗醋栗果实生长发育规律研究.东北农学院学报,1987,18(2):119-124
[11]鄢德锐,韩德全.山楂果实生长发育规律的研究.山西果树,1989,(4):17-19
[12]李洁维,李瑞高,梁木源,等.猴桃优良株系果实生长发育规律研究.广西植物,1992,12(2):152-156
[13]卢克成,石鼎新,倪竞德.中华猕猴桃果实生长发育规律的初步研究.江苏林业科技,1999,26(4):14-16
[14]王荣花,李嘉瑞,陈理论.杏果实发育的形态解剖学研究.西北农业大学学报,2000,22(4):45-50
[15]金方伦.黔北地区两个梨品种果实生长发育规律的研究.广西园艺,2001,36(1):5-6
[16]吕金海,朱荣林,周书伟.金秋梨叶、梢和果实的生长动态及相关性.西南园艺,2006,34(1):20-21
[17]韩明玉,田玉命,张慧梅,等.秦光油桃果实生长曲线和落果波相的观察.西北植物学报,2001,21(6):1249-1253
[18]苍晶,王学东,桂明珠,等.狗枣猕猴桃果实生长发育的研究.果树学报,2002,18(2):87-90
[19]苍晶,王学东,张达,等.软枣猕猴桃果实生长发育的研究.东北农业大学学报,2004,35(1):77-83
[20]许方主编.梨树生物学.北京:科学出版社,1992
[21]Goffinet M C,Robinson T L,Lakso A N.A comparison of 'Empire' apple fruit size and anatomy in unthinned and handthinned trees.J.Hort Sci,1984,70(3):375-378
[22]曹亚波,周亚娣,许善华,等.早熟优质梨新品种-早蜜梨.宁波农业科技,2003,(2):30-31
[23]文仁德,常运涛.早熟梨新品系-“早蜜”.广西园艺,2000,(4):19-20
[24]刘炳刚.中晚熟梨品种黄金梨引种初报.中国果树,2001,(1):13-14
[25]刘长根,余国清.黄金梨生物学特性及栽培技术要点.中国南方果树,2004,33(5):89-90
[26]刘遵春.金花梨变异单系生物学特性调查比较及RAPD多态性分析.雅安:四川农业大学硕士学位论文,2004
[27]裴秦青,张国忠.鸭梨采收期与贮藏场所的研究.河北农业大学学报,1989,12(1):46-53
[28]林真二著(吴耕民译).梨.北京:农业出版社,1981
[29]梁东,马锋旺,管清美,等.蔷薇科植物中山梨醇代谢酶的研究进展.西北植物学报,2004,24(7):1362-1366
[30]胡庆祥,郡荣庭,张玉星,等.鸭梨果实发育中糖酸变化特点及配方施肥效应.河北果树,1996(1):9-11
[31]李春英,金英善,李永勋.苹果梨果实生长期间山梨糖醇及其它糖的变化规律.延边大学农学学报,2001,23(3):198-201
[32]李春英,薛桂新,李永勋.客发梨果实生长期间山梨糖醇及其它糖变化规律的研究.北方果树,1999(6):6-7,26
[33]王军虹,闰丽梅,刘武林.大香水梨成熟过程生理生化反应.北方园艺,1998,(6):24-25
[34]敖良德,王明鑫.苹果成熟过程中果实内单宁酸、苹果酸、维生素C及淀粉含量的变化.中国果树,1988,(2):35-36
[35]邓月娥,张传来,牛立元,等.桃果实发育过程中主要营养成分的动态变化及系统分析方法研究.果树科学,1998,15(1):48-52.
[36]甘霖,谢永红,吴正琴,等.‘嘉平大枣'果实发育过程中糖、酸及维生素C含量的变化.园艺学报,2000,27(5):317-320
[37]王军虹,闰丽梅,刘武林.苹果梨采前生理生化变化.东北农业大学学报,1998,29(4):388-392
[38]申曙光,马宝焜,陈四维.红富士苹果果实发育期间生理生化变化的研究.园艺学报,1991,18(1):1-3
[39]Vizzoto G,Pinto R,Varanini Z.Sucrose accumulation in developing peach fruit.Plant Physiol,1996,96:225-230
[40]Davis C,Robinson SP.Sugar accumulation in grape berries.Plant Physiol,1996,111:275-283
[41]吕英民,张大鹏.果实发育过程中糖的积累.植物生理学通讯,2000,36(3):258-265
[42]陈俊伟,张良诚,张上隆.果实中的糖分积累机理.植物生理学通讯,2000,36(6):497-503
[43]陈俊伟,张上隆,张良诚.果实中糖的运输、代谢与积累及其调控.植物生理与分子生物学学报,2004,30(1):1-10
[44]陈俊伟,谢鸣,秦巧平.植物糖信号与激素信号之间的联系.植物生理学通讯,2005,41(3):279-285
[45]谢国禄.苹果和日本梨果实发育期间果糖含量和果糖相关酶活性.国外作物育种,2001,70(1): 16-20
[46]Beruter J,Studer Feusi ME,Ruedi P.Sorbitol and sucrose partitioning in the growing apple fruit.J Plant Physiol,1997,151:269-276
[47]Beruter J,Studer Feusi ME.Comparison of sorbitol transport in excised tissue discs and cortex tissue of intact apple fruit.Plant Physiol,1995,146:95-102
[48]Beruter J,Studer Feusi ME.The effect of girdling on carbohydrate partitioning in the growing apple fruit.J Plant Physiol,1997,151:277-285
[49]Yamaguchi H,Kanayama Y,Soejima J,etal.Changes in the a mounts of the NAD dependent sorbitol dehydrogenase and its involvement in the development of apple fruit.J Amer Soc Hort Sci,1996,121(5):848-852
[50]单守明,王永章,董晓颖,等.IAA、GA和ABA对苹果果实山梨醇代谢相关酶活性的影响.园艺学报,2005,32(6):990-993
[51]Davis C,Robinson SP.Sugar accumulation in grape berries.Plant Physiol,1996,111:275-283
[52]Farrar JF.Sink strength:What is it and how do we measure it?(Forum) Plant Cell and Enviorn,1993,16:1014-1046
[53]Hirano K,Yone mori K,Sugiura A.Involvement of sugar metabolism in persimmon growth inhibition by calyx lobe removal.J Amer Soc Hort Sci,1996,121(3):456-472
[54]Moriguchi T,Yamaki S.Purification and characterization of sucrose synthase from peach(Prunus persica) fruit.Plant Cell Physiol,1988,29(8):1361-1366
[55]Sun J.Sucrose synathase in wild tomato,Lycopersicon chmielewskii,and tomato fruit sink strength.Plant Physiol,1992,98:1163-1169
[56]Sung SS,Sheih WJ,Geiger DR,etal.Growth,sucrose synthase,and invertase activities of developing Phaseolus vulgaris L.fruit.Plant Cell and Enviorn,1994,17:419-426
[57]Yamaguchi H,Kanayama Y,Yamaki S.Purification and properties of NAD dependent sorbitol dehydrogenase from apple fruit.Plant Cell Physiol,1994,35(6):887-892
[58]Yamaki S,Ishikawa K.Roles of four sorbitol related enzymes and invertase in the seasonal alteration of sugar metabolism in apple tissue.J Amer Soc Hort Sci,1986,111(1):134-137
[59]辛贺明,张喜焕.套袋对鸭梨果实内含物变化及内源激素水平的影响.果树学报,2003,20(3):233-235
[60]程和禾.黄金梨果实生长发育及贮藏生理特性研究.保定:河北农业大学硕士学位论文,2004
[61]李金昶,石晶,苏忠民,等.苹果梨成熟和贮藏过程中有机酸含量的变化.植物生理学通讯,1995,31(2):118
[62]林敏娟.黄金梨叶片、果实内营养物质含量变化及其相关关系研究.保定:河北农业大学硕士学位论文,2005
[63]潘瑞炽主编.植物生理学.北京:高等教育出版社(第三版),1995
[64]林敏娟,徐继忠,陈海江,等.黄金梨叶片、果实中矿质元素含量的周年变化动态.河北农业大学学报,2005,28(6):23-27
[65]刘国荣.不同矮化中间砧红富士苹果果实生长及其内含物含量变化的研究.保定:河北农业大学硕士学位论文,2003
[66]李树玲,黄礼森,丛佩华。等.不同种内梨品种果实维生素C含量.园艺学报,1994,21(1):17-20
[67]吕金海,伍贤进,周书伟,等.金秋梨果实发育中生长速率及主要营养成分的变化.中国农学通报,2000,20(5):35-36,44
[68]刘剑锋,张红艳,彭抒昂.梨果实发育中果肉及种子钙和果胶含量的变化.园艺学报2003,30(6):709-711
[69]刘利德,姚敦义.植物激素的概念及其新成员.生物学通报,2002,37(8):18-20
[70]李玉珍.植物激素的研究进展.河南科技学院学报(自然科学版),2007,35(4):50-53
[71]刘剑锋,程云清,彭抒昂.梨果肉与种子中钙与内源激素含量变化关系研究.植物营养与肥料学报,2005,11(2):269-272
[72]阮晓,王强,周疆明,等.香梨果实成熟衰老过程中4种内源激素的变化.植物生理学报,2000,26(5):402-406
[73]饶景萍,平田尚美.猕猴桃果实生长发育过程中内源植物激素的变化动态.西北农业大学学报,1992,20(4):41-44
[74]陶汉之,高丽萍,陈佩璁,等.猕猴桃果实发育中内源激素水平变化的研究.园艺学报,1994,21(1):35-40
[75]肖家欣,彭抒昂,何华平.单性结实和自花结实柑橘果实发育过程中IAA、ZR和GA3含量的变化.武汉植物学研究,2005,23(3):272-275
[76]李秀菊,刘用生,束怀瑞.不同成熟型苹果果实生长发育过程中几种内源植物激素含量变化.植物生理学通讯,2000,36(1):7-10
[77]彭福田,姜远茂,顾曼如,等.氮素对苹果果实内源激素变化动态与发育进程的影响.植物营养与肥料学报,2003,9(2):208-213
[78]钟晓红,马定渭,黄远飞.草莓果实发育过程中内源激素水平的变化.江西农业大学学报,2004,26(1):107-111
[79]周碧燕,季作梁,叶永昌,等.荔枝果实发育期间内源激素含量的变化.园艺学报,1998,25(3):236-240
[80]周碧燕,李建国,黄旭明,等.龙眼果实发育过程中ZRs、GA和IAA含量的变化.华南农业大学学报,1999,20(3):50-53
[81]王碧青,邱燕萍,向旭,等.荔枝结果过程中内源激素变化及单性结果的诱导.园艺学报,1997,24(1):19-24
[82]季作梁,梁立峰,柳建良,等.荔枝果实发育期间内源激素含量动态.华南农业大学学报,1992,13(3):93-98
[83]樊卫国,安华明,刘国琴,等.刺梨果实与种子内源激素含量变化及其与果实发育的关系.中国农业科学,2004,37(5):728-733
[84]陈发河,蔡慧农,冯作山,等.葡萄浆果发育过程中激素水平的变化.植物生理与分子生物学学报,2002,28(5):391-395
[85]Luckwill L C.Studies of fruit development in relation to plant hormone.Ⅳ.Alidic auxins and growth inhibitors in leaves and fruit of the apple.The Journal of Horticultural Science,1957(32):18-33
[86]丁长奎,章恢志.植物激素对枇杷果实生长发育的影响.园艺学报,1988,15(3):148-154
[87]Hopping M.E.,Bukoval M.J..Endogenous plant growth substances in developing fruit of Prunus cerasus L.Ⅵ.Extractable auxin in the seed and pericarp.The Journal of Horticultural Science.1975,100(40):399-401
[88]王忠主编.植物生理学.北京:中国农业出版社,2000
[89]吕忠恕著.果树生理.上海:上海科学技术出版社,1982
[90]郗荣庭主编.果树栽培学总论(第三版).北京:中国农业出版社,2004
[91]杨立峰,刘弦,郝风鸽.GA_3对苹果生长发育的影响研究.河北果树,2004,(5):8-10
[92]John E.Jackson.Biology of apples and pears.UK:Cambridge University Press,2003
[93]吴少华,沈德绪,林伯年,等.梨果实发育和直感同内源激素的关系.浙江农业大学学报,1986,12(1):57-61
[94]钟晓红,马定渭,黄远飞.草莓果实发育过程中内源激素水平的变化.江西农业大学学报,2004,26(1):107-111
[95]吴俊,钟家煌,徐凯,等.外源GA_3对藤稔葡萄果实生长发育及内源激素水平的影响.果树学报2001,18(4):209-212
[96]陈大成,李志勇,胡桂兵.‘塔1'椪柑果实发育、内源激素变化及品质形成特点.华南农业大学学报,1998,19(3):51-55
[97]Kiyohide Kojima,Keishi Shiozaki.Changes of endogeneous levels of ABA,IAA and GA-like substances in fruitlets of parthenocarpic persimmon.Japan Soc Hort Sci,1999,68(2):242-247
[98]张崇浩,袁玉信,李杰芬,等.桃果实后熟中内源激素的变化.北京师范大学学报(自然科学版),1989,(2):68-72
[99]Vanoli M,Marro M,Visai C.Gibberellin-like substances during the development of peach fruits.Acta Horticulturae,1993,329:159-16
[100]Cutting J G M,Lyne M C.Girdling and the reduction in shoot xylem sap concentrations Of cytokinins and gibberellins in peach.Journal of Horticultural Science,1993,68(4):619-626
[101]Dennis F G Jr.Trial of ethephon and other growth regulation for delaying bloom in tree fruits.Journal of the American Society for Horticultural Science,1976,101(3):241-245
[102]刘小阳,李玲,蔡永萍.光强对砀山酥梨石细胞形成的影响及其与内源IAA、ZR和ABA含量的关系.激光生物学报,2006,15(2):161-166
[103]邹养军,王永熙.内源激素对苹果果实生长发育的调控作用研究进展.陕西农业科学,2002(10):13-15
[104]闫树堂,徐继忠.不同矮化中间砧对红富士苹果果实内源激素、多胺与细胞分裂的影响.园艺学报,2005,32(1):81-83
[105]闫国华,甘立军,孙瑞红.赤霉素和细胞分裂素调控苹果果实早期生长发育机理的研究.园艺学报,2000,27(1):11-16
[106]季作梁,王钢涛.荔枝果实发育过程中细胞分裂素的变化.果树科学,1996,13(2):92-95
[107]李建国,周碧燕.大核和焦核“桂味”荔枝果实发育及其发育期间果皮中内源激素含量的变化比较.植物生理学通讯,2005,41(5):587-590
[108]张海娥,郭修武.激素在葡萄分子发育调控上的作用研究进展.中外葡萄与葡萄酒,2005,(2):28-29,32
[109]Salopek-Sondi B,Kovac M,Prebeg T,et al.Developing fruit direct post-floral morphogenesis in Helleborus fruit direct post-floral morphogenesis in Helleborus niger L.J Exp Bot.2002,53(376):1949-1957
[110]周蕾,魏琦超,高峰.细胞分裂素在果实及种子发育中的作用.植物生理学通讯,2006,42(3):549-553
[111]彭良志,胥耳.6-BA和GA_3对华盛顿脐橙幼果~(14)C光合产物调配的影响.园艺学报,1990,17(2):111-116
[112]倪竹如,陈俊伟,阮美颖,等.BA对椪柑果实生长发育及其同化产物分配的影响.浙江农业学报,2000,12(5):272-276
[113]Stern RA,Shargal A,Flaishman MA.Thidiazuron increases fruit size of 'Spadona'and 'Coscia' pear(Pyrus communis L.).J Hortic Sci Biotechnol,2003,78(1):51-55
[114]Sotta B,Sossountzov L,Maldiney R,etal.Abscisic acid localization by light microscopic immunohistochemistry in chenopodium polyspermum L.J Histochem Cytochem,1985,33(3):201-208
[115]Sossountzov L,Sotta B,Maldiney R,etal.Immunoelectron-microscopy Localization of abscisic acid with colloidal gold on Lowicryl-embedded tissues of chenopodium polyspermum L.Planta,1986,168:471-481
[116]Bertrand S,Benhamou N,Nadeau P,etal.Immunogold localization of free abscisic acid in tomato root cells.Canadian J Bot,1992,70(5):1001-1011
[117]贾文锁,王学臣,娄成后.蚕豆(Vicia faba L.)叶肉细胞中ABA的胶体金免疫电镜定位.植物生理学报,1994,20(4):380-384
[118]王学臣,贾文锁.水分胁迫下蚕豆(Vicia faba L.)气孔关闭与叶片细胞中ABA区隔化与再分配的关系.植物生理学报,1995,21(4):324-328
[119]Pastor A,Cortadellas N,Alegre L.Immunolocalization of abscisic acid by monoclonal antibodies in Lavandula stoechas L.leaves.Plant Growth Regulation,1995,16:287-292
[120]黄丛林,张大鹏,贾文锁.葡萄种子细胞中脱落酸的胶体金免疫电镜定位.中国农大大学学报,1999,4(5):111-117
[121]黄丛林,贾文锁,张大鹏.葡萄果肉细胞中脱落酸的区隔化研究.植物生理学报,2000,26(4):297-300
[122]Zhang D P,Huang C L,Jia W S.Immunogold electron-microscopy localization of abscisic acid in fresh cells of grape berry(Vitis vinifera L.×Vitis labrusca L.cv Kyoho).Sci Hort,1999,81:189-198
[123]刘勇,刘善军,霍光华,等.甜柿果实发育期间矿质元素与营养成分的变化.江西农业大学学报,2000,22(2):265-270
[124]郑国华,杉浦明,米森敬三.柿果(Diospyros Kaki L.)发育及成熟过程中内源GAs活性及ABA含量的变化.北京农业大学学报,1991,17(1):77-81
[125]郑国华,平野建.喷施GA_3和乙烯利对柿果实成熟及内源GAs活性,ABA含量的影响.园艺学报,1991,18(3):193-197
[126]张大鹏,张子连,陈珈.葡萄果实发育过程中脱落酸结合蛋白动力学特性的变化.植物学报,1996,38(12):930-935
[127]Kojima K,Kuraishi S,Sakurai N.Distribution of ABA in different parts of the reproductive organs of tomato.Sci Hort,1993,56:23-30
[128]I.Lara,M.Vendrell.Changes in abscisic acid levels,ethylene biosynthesis,and protein patterns during fruit maturation of 'Granny Smith' aplles.J.Amer.Soc.Hort.Sci,2000,125(2):183-189
[129]Satoru Kondo,Hiroshi Gemma.Relationship between abscisic acid(ABA) content and maturation of the sweet cherry.J.Japan.Soc.Hort.Sci,1993,62(1):63-68
[130]黄丛林,张大鹏,贾文锁.葡萄果实发育后期脱落酸来源的研究.园艺学报,2001,28(5):385-391
[131]Yuan R C,Huang H B.Litchi fruit abscission:its pattern,effects of Shading and relation to abscisic acid.Scientia Horticulturae,1988,36(3-4):281-292
[132]Coombe B G.The grape berry as a sink.Acta Hort,1989,(239):149-158
[133]Archbold D D.Abscisic acid facilitates sucrose import by strawberry fruit explants and cortex disks in vitro.HortSci,1988,23:880-881
[134]Doucette M S,Pritchard M K,Saltveit M E.ABA involvement in low temperature sweetening of potatoes.Acta Hort,1993,(343):293-294
[135]Kojima K,Yamada Y,Yamamoto M.Distribution of ABA and IAA with a developing valencia orange fruit and its parts.J Jpn Soc Hort Sci,1994,63:335-339
[136]Smith G S,Klages K V,Green T G A,etal.Changes in abscisic acid concentration,surface conductance and water content of developing kiwifruit.Sci Hort,1995,61(1-2):13-27
[137]吴有梅,顾采琴.ABA和乙烯在草莓采后成熟过程中的作用.植物生理学报,1992,18(2):167-172
[138]Leung J,Giraudat J.Abscisic acid signal transduction.Annu Rev Plant Physiol Plant Mol Biol,1998,49:199-222
[139]Dik Hagenbeek,Ralph S.Quatrano,Christopher D.Rock,et al.Trivalent ions activate abscisic acid-inducible promoters through an ABI1-dependent pathway in rice protoplasts.Plant Physiol,2000,123:1553-1560
[140]阮晓,王强,王文全,等.香梨果实发育、成熟期间乙烯、ACC、MACC变化及相互关系.园艺学报,2000,27(3):205-206
[141]韦军,田边贤二.鸭梨、茌梨果实采前乙烯生物合成特性研究.园艺学报,1994,21(3):231-234
[142]关军锋,束怀瑞,黄天栋.新红星苹果果实细胞膨大期及成熟期乙烯生成的变化.中国农业科学,1993,26(5):57-60
[143]李全梓,李兴国,郑国生,等.早熟桃果实和种子发育过程中乙烯产生及ACC含量和ACC氧化酶活性的变化.植物生理学通讯,1998,34(1):25-27
[144]王蓉花,李嘉瑞,张娟,等.杏果实生长发育生理学研究.西北农业学报,2000,9(3):76-78
[145]李全梓,李兴国,张宪省,等.新红星苹果发育过程中果实和种子乙烯生物合成的研究.山东农业大学学报,1998,29(2):151-155
[146]李正国,El-Sharkawy I,Lelievre J-M.温度、丙烯和1—MCP对西洋梨果实乙烯合成和乙烯受体ETR1同源基因表达的影响.园艺学报,2000,27(5):313-316
[147]Li Zheng-Guo,El-Sharkawy,Bouzayen.Effects of cold treatment on ethylene biosynthesis and responsiveness during ripening of Pear(Pyrus communis L.)fruits.植物生理学报,2000,26(4):283-288
[148]Lelievre J M,Tichit L,Dao P.Effects of chilling on the expression of ethylene biosynthesis genes in Passe-Crassane pear fruits.Plant Mol.Biol.,1997,101:727-739
[149]魏绍冲,陈昆松,罗云波.乙烯受体与果实成熟调控.园艺学报,2004,31(4):543-548
[150]周俊义.多胺在果树生理的作用.果树科学,1995,12(2):117-119
[151]崔凤芝,张运涛.多胺与园艺植物生长发育的关系.河北农业大学学报,1996,19(3):94-98
[152]杨洪强,接玉玲.多胺与果树生长发育的关系.山东农业大学学报,1995,27(40):514-520
[153]Bernet E,Claparolsa I,Dondini L,et al.Changes in polyamine content,arginine and ornithine decarboxylases and ransglutaminase activities during light dark phases(of initial differentiation)in maize calluses and their chloroplasts.Plant Physiol Biochem,1999,37(12):899-909
[154]Shoeb F,Yadav JS,Bajaj LS,etal.Polyamines biomarkers for plant regeneration capacity:improvement of regeneration by modulation of polyamine metabolism in different genotypes of indica rice.Plant Seience,2001,160:1229-1235
[155]Omar franco-Mora,Kenji Tanabe,Akihiro Itai,etal.Relationship between endogenous free polyamines content and ethylene evolution during fruit growth and ripening of Japanese Pear(Pyrus pyrifolia Nakai).J.Japan.Soc.Hort.Sci,2005,74(3):221-227
[156]王立英.柿(DiospyroskakiL.)果实生长发育及成熟机理研究.保定:河北农业大学硕士学位论文,2002
[157]Biasi R,Bagni N.Endogenous polyamines in apple and their relationship to fruit set and fruit growth.Physiol Plant,1988,(73):201-205
[158]王世平,宋长冰,李连朝,等.三种多胺在苹果开花及坐果初期的生理作用.园艺学报,1996,23(4):319-325
[159]陈海江,徐继忠,袁小乱,等.不同时期喷施外源多胺对新红星苹果坐果的影响.河北农业大学学报,1999,22(4):47-49
[160]Aziz A.Spermidine and related-metabolic inhibitors modulate sugar and amino acid levels in Vitis vinifera L:possible relationships with initial fruitlet abscission.J Exp Bot,2003,54:355-363
[161]徐继忠,史常青,马宝焜,等.外源多胺对苹果花粉管伸长及胚珠寿命的影响.园艺学报,1998,25:91-92
[162]钟晓红.三种多胺在沙田柚开花及坐果初期的生理作用.湖南农业大学学报(自然科学版),2000,26(6):453-456
[163]Koshgad M M,Orvos A R,Yelenosky G.Relative change in polyamines during citrus flower development.HortScience,1990,(25):946-948
[164]郑玉生,张秋明,Lovoctt C J.柑橘花期代谢及其对坐果的影响.果树科学,1997,14(4):216-219
[165]Malik AU,Singh Z.Abscission of mango fruitlets as influenced by biosynthesis of polyamines.Hort Sci Biotechnol,2003,78(5):721-727
[166]Wolukaul JN,Zhang SL,Xu GH,et al.The effect of temperature,polyamines and polyamine synthesis inhibitor on in vitro pollen germination and pollen tube growth of Prunus mume.Sci Hort,2004,99:289-299
[167]Shiozaki S,T.Ogata,S.Horiuchi.Endogenous polyamines in the pericarp and seed of the grape berry during development and ripening.Scientia Hortic,2000,83:33-41
[168]陈杰忠,骆小兵,王泽槐,等.多胺对荔枝果实坐果率及内源多胺含量的影响.园艺学进展(第五辑)2000:303-307
[169]李建国,刘顺枝,王泽槐.荔枝果实发育过程中内源多胺含量的变化.植物生理学通讯,2004,40(2):153-156
[170]李建勇,卢钢,任彦.多胺在果实生长发育中的作用研究进展.果树学报,2005,22(3):256-260
[171]Lester GE.Polyamines and their cellular anti-senescence properties in honey dew muskmelon fruit.Plant Science,2000,160:105-112
[172]陈昆松,张上隆.脂氧合酶与果实的成熟衰老.园艺学报,1998,25(4):338-344
[173]曹慧,王永章,杜俊杰,等.脂氧合酶研究进展(综述).安徽农业大学学报,2001,28(2):219-222
[174]陈晶,李正国.脂氧合酶在果实成熟衰老中的作用.广州食品工业科技,2002,18(1):50-53
[175]张荣平.脂氧合酶在植物体内的生理功能.莱阳农学院学报,1993,10(1):47-51
[176]许文平,陈昆松,徐昌杰,等.猕猴桃采后果实冷藏与货架期脂氧合酶活性和乙烯生成的变化.中国农业科学,2003,36(10):1196-1201
[177]许文平,陈昆松,李方,等.脂氧合酶、茉莉酸和水杨酸对猕猴桃果实后熟软化进程中乙烯生物合成的调控.植物生理学报,2000,26(6):507-514
[178]罗云波.脂氧合酶与番茄采后成熟的关系.园艺学报,1994,21(4):357-360
[179]Riley J C M,Willemot C,Thompson J E.Lipoxygenase and hydroperoxide lyase activities in ripening tomato fruit.Postharvest Biology and Technology,1996,7:97-107
[180]Marcelle RD.Relationship between mineral content,lipoxygenase activity,levels of 1-aminocyclopropane-1-carboxylic acid and ethylene emission in apple fruit flesh disks(cv.Jonagold)during storage.Postharvest Biology and Technology,1991,(1):101-109
[181]吴敏,陈昆松,张上隆.桃果实采后成熟过程中脂氧合酶活性变化.园艺学 报,1999,26(4):227-231
[182]谢永红,陈学年,吕斌,等.桃果实采后脂氧合酶活性与内源乙烯对果实软化衰老的影响.西南园艺,2004,32(6):1-3
[183]Todd J F,Paliyath G,Thompson J E.Characteristics of a membrane-associated lipoxygenase in tomato fruit.Plant Physiol,1990,94:1225-1232
[184]韩碧文主编.植物生长与分化.北京:中国农业大学出版社,2003
[185]寇晓虹,罗云波.植物多聚半乳糖醛酸酶研究进展.山西农业大学学报,2004,24(2):411-415
[186]彭丽桃,饶景萍,杨书珍,等.果实软化的胞壁物质和水解酶变化.热带亚热带植物学报,2002,10(3):271-280
[187]赵云峰,林河通,林娇芬,等.果实软化的细胞壁降解酶及其调控研究进展.仲恺农业技术学院学报,2006,19(1):65-70
[188]薛炳烨,束怀瑞.肥城桃两品系果实细胞壁成分和水解酶活性的比较.园艺学报,2004,31(4):499-501
[189]曾秀丽,张光伦,李春燕,等.三个脐橙品种果实主要细胞壁酶动态变化研究.亚热带植物科学,2006,35(2):12-16
[190]李春燕,张光伦,曾秀丽,等.细胞壁酶活性与甜橙果实质地的相关性研究.四川农业大学学报,2006,24(1):73-76
[191]李建国,黄旭明,黄辉白.裂果易发性不同的荔枝品种果皮中细胞壁代谢酶活性的比较.植物生理与分子生物学学报,2003,29(2):141-146
[192]薛炳烨,毛志泉,束怀瑞.草莓果实发育成熟过程中糖苷酶和纤维素酶活性及细胞壁组成变化.植物生理与分子生物学学报,2006,32(3):363-368
[193]张嵩,张光伦,曾秀丽.纤维素酶和多聚半乳糖醛酸酶与果实成熟.果树学报,2005,22(5):532-536
[194]寇晓虹,罗云波,田慧琴,等.多聚半乳糖醛酸酶(PG)反义基因转化加工番茄.食品科学,2007,28(3):187-191
[195]李雄彪.植物细胞壁酶的分子结构与生理功能.植物生理通讯,1991,27(4):246-252
[196]张昭其,段学武,陆旺金.果实多聚半乳糖醛酸酶分子生物学研究进展.生命科学,2002,14(2):92-94
[197]薛炳烨,束怀瑞.应用基因工程改良果实成熟特性研究进展.山东农业科学,2002,(4):49-52
[198]Koch K E.Carbohydrate-modulated gene expression in plants.Annu Rev Plant Physiol Plant Mol Biol,1996,47:509-540
[199]Leon P,Sheen J.Sugar and hormone connection.Trends Plant Sci,2003,8:110-116
[200]Sheen J,Zhou L,JiangLC.Sugars as signaling molecules.Curr Opin Plant Biol,1999,2:410-418
[201]Smeekens S.Sugar-induced signal transduction in plants.Annu Rev Plant Physiol Plant Mol Biol,2000,51:49-81
[202]Leon P,Sheen J.Sugar and hormone connection.Trends Plant Sci,2003,8:110-116
[203]Rolland F,Moore B,Sheen J.Sugar sensing and signaling in plants.Plant Cell,2002,14:185-205
[204]Stadler R,Truernit E,Gahrtz M,etal.The AtSUC1 sucrose carrier may represent the osmotic driving force for anther dehiscence and pollen tube growth in Arabidopsis.Plant J,1999,19:269-278
[205]Bethke P C,Schuurink P C,Jones R L.normonal signaling in cereal alcurone.J Exp Bot,1997,48:1337-1356
[206]吴有梅,刘愚.番茄果实成熟过程中钙调索含量变化及其与乙烯合成的关系.植物学报,1990,16:709-713
[207]关军锋,MaX Saure编著.果树钙素营养与生理.北京:科学出版社,2005
[208]罗充,彭抒昂,李国怀.梨不同品种果实发育中钙与钙调素的动态研究.华中农业大学学报,2001,20(3):286-288
[209]张红艳,彭抒昂.梨果实发育过程中钙调素水平的动态研究.华中农业大学学报,2004,23(3):338-340
[210]关军锋,马智宏,张会敏,等.苹果幼果发育期果柄结构和Ca、Mg、K水平的变化.植物营养与肥料学报,2005,11(1):264-268
[211]蒲富慎,黄礼森,孙秉钧,等编.梨品种.北京:农业出版社,1989
[212]华中农业大学主编.果树研究法(第二版).北京:农业出版社,1979
[213]骆建霞,孙建设主编.园艺植物科学研究导论.北京:中国农业出版社,2001
[214]束怀瑞主编.果树栽培生理学.北京:中国农业出版社,1993
[215]李锡香编著.新鲜果蔬的品质及其分析法.北京:中国农业出版社,1994
[216]中国科学院上海植物生理研究所编.现代植物生理学实验指南.北京:科学出版社,1999
[217]张友杰.以蒽酮分光光度法测定果蔬中.的葡萄糖、果糖、蔗糖和淀粉.分析化学,1980,5(3):167-171
[218]李合生主编.植物生理生化实验原理和技术.北京:高等教育出版社,2000
[219]朱广廉,钟海文,张爱琴编.植物生理学实验.北京:北京大学出版社,1990
[220]Surrey K.Spectrophotometric method for determination of lipoxidase activity.Plant physiol.1963.39:65-70
[221]徐昌杰,陈昆松,张上隆.蔗糖酶对柑桔外切纤维素酶和外切多聚半乳糖醛酸酶活性测定的干扰及排除.植物生理通讯,1997,33(1):43-46
[222]宁正祥.食品成分分析手册.北京:中国轻工业出版社,1998
[223]张飞,岳田利,费坚,等.果胶酶活力的测定方法研究.西北农业学报,2004,13(4):134-137
[224]田志喜,张玉星,于艳军,等.水杨酸对鸭梨果实PG、PME和呼吸速率的影响.果树学报,2002,19(6):381-384
[225]上海植物生理学会编.植物生理实验指导手册.上海:上海科学技术出版社,1985
[226]丁静,沈镇德,方亦雄,等.植物内源激素的提取分离和生物鉴定.植物生理学通讯,1979,2:27-29
[227]陈昆松,徐昌杰,李方,等.HPLC法检测果实组织中内源IAA、ABA方法的改进.果树学报,2003,20(1):4-7
[228]陈远平,杨文钰.卵叶韭休眠芽中GA3、IAA、ABA和ZT的高效液相色谱法测定.四川农业大学学报,2005,23(4):498-500
[229]曾庆钱,陈厚彬,鲁才浩,等.HPLC测定荔枝不同器官中内源激素流程的优化.果树学报,2006,23(1):145-148
[230]Hector E.Flores,Arthur W.Galston.Analysis of polyamines in high plants by high performance liquid chromatography.Plant Physiol.,1982,69:701-706
[231]刘俊,吉晓佳,刘友良.检测植物组织中多胺含量的高效液相色谱法.植物生理学通讯,2002,38(6):596-598
[232]Liu Ji-Hong,Kazuyoushi Nada,Pang Xiao-ming,et al.Role of polyamines in peach fruit development and storage.Tree Physiology,2006,26:791-798
[233]盛宝龙.不同梨品种光合特性及影响因素研究.南京:南京农业大学硕士学位论文,2004
[234]郭春爱,刘芳,许晓明.叶绿素b缺失与植物的光合作用.植物生理学通讯,2006,42(5):967-973
[235]汪良驹,姜卫兵,高光林,等.幼年梨树品种光合作用的研究.园艺学报,2005,32(4):571-577
[236]Caixi Zhang,Kenji Tanabe,Shiping Wang,etal.The impact of cell division and cell enlargement on the evolution of fruit size in Pyrus pyrifolia.Annals of Botany,2006,98:537-543
[237]王贵元,夏仁学,吴强盛.果实中糖分的积累与代谢研究进展.北方园艺,2007(3):56-58
[238]胡红菊,陈启亮,王友平,等.4个砂梨品种果实发育过程中主要糖酸含量的变化.华中农业大学学报,2007,26(2):251-255
[239]赵智中.柑橘果实糖积累的生理基础研究.杭州:浙江大学博士学位论文,2001
[240]龚荣高,张光伦,吕秀兰,等.脐橙在不同生境下果实蔗糖代谢相关酶的研究.园艺学报,2004,31(6):719-722
[241]刘永忠,李道高.柑橘果实糖积累与蔗糖代谢酶活性的研究.园艺学报,2003,30(4):457-459
[242]李培环,董晓颖,王永章,等.‘新红星'苹果果实蔗糖合酶的活性及亚细胞定位.园艺学报,2002,29(4):375-37;
[243]夏国海,张大鹏,贾文锁,等.IAA、GA和ABA对葡萄果实14C蔗糖输入与代谢的调控。园艺学 报,2000,27:6-10
[244]Archbold R C.Abscisic acid facilitates sucrose import by strawberry fruit explants and cortex disks in vitro.Hortic Sci,1988,23:880-883
[245]Fischer R.L.,Bennett A.B.Role of cell wall hydrolases in fruit ripening.Annu Rev Plant Physiology Plant Molecular Biology,1991,142:675-703
[246]刘剑锋,程云清,彭抒昂.梨采后细胞壁成分及果胶酶活性与果肉质地的关系.园艺学报,2004,31(5):579-583
[247]Bonghi,C.,Ferrarese,L..Endo-b-1,4-glucanases are involved in peach fruit growth and ripening,and regulated by ethylene.Physiologia Plantarum,1998,102:346-352
[248]Siedow JN.Plant lipoxygenease:structure and function.Annu.Rev.Plant MOL.Biol.1991,42:145-188
[249]Bousquest J F,Thimann K V.Lipid peroxidation forms ethylene from ACC and may operate in leaf senescence.Proc.Natn.Acad.Sci.USA,1984,81:1724-1727
[250]黄卫东,原永兵,彭宜本.温带果树结实生理.北京:北京农业大学出版社,1994
[251]Lieberman N,Bake J E,Sloge.Influence of plant hormones ethylene production in apple,tomato and avocado slices during maturation and senescence.Plant Physiol,1977,60:214-217
[252]张大鹏,许雪峰,张子连,等.果实始熟机理.园艺学报,1997,24(1):1-7
[253]李六林,张绍玲,郭艳玲.‘新高'梨花粉败育与内源多胺含量变化的关系.园艺学报,2007,34(2):301-304
[254]郭志雄.油柰坐果及果实生长发育机制的若干研究.福州:福建农业大学硕士学位论文,1999
[255]Ke D and Romani R J.1988.Effects of spermidine on ethylene production and the senescence of suspension-cultured pear fruit cells.Plant Physiol.Biochem.,26:109-116
[256]Martin Tanguy J.1997.Conjugated polyamines and reproductive development:biochemical,molecular and physiological approaches.Physiol.Plant.,100:675-688
[257]Minocha R,Lee J S,Long S,Bhatnagar P,Minocha S C.2004.Physiological responses of wild type and putrescine-overproducing transgenic cells of polar to variations in the fom and concentration of nitrogen in the medium.Tree Physiology,24:551-560
[258]Rastogi R,Davies P J.1991.Effects of light and plant growth regulators on polyamine metabolism in higher plants.In:Slocum,R.D.,Flores,H.E.(Eds.),Biochemistry and Physiology of Polyamines in Plants.CRC Press,Boca Raton,187-199