本文關(guān)鍵詞：山葡萄糖酸積累規(guī)律及代謝調(diào)控機(jī)理研究　出處：《西北農(nóng)林科技大學(xué)》2016年博士論文　論文類型：學(xué)位論文

  更多相關(guān)文章： 山葡萄 光合熒光特性 生理特性 糖和有機(jī)酸代謝相關(guān)酶 代謝物運(yùn)輸

【摘要】：糖和有機(jī)酸含量是決定葡萄品質(zhì)及葡萄酒質(zhì)量的關(guān)鍵因素。在實際生產(chǎn)當(dāng)中,可通過采用不同的栽培措施來調(diào)節(jié)葡萄生長的微環(huán)境、改善葡萄品質(zhì),從而獲得優(yōu)質(zhì)的葡萄酒。本研究以集安產(chǎn)區(qū)的山葡萄品種北冰紅和雙紅作為試驗試材,對其光合熒光特性以及漿果內(nèi)糖和有機(jī)酸含量及其相關(guān)代謝酶進(jìn)行了深入的研究,并采用不同架式、不同負(fù)載和不同光呼吸抑制劑處理來研究不同栽培措施對山葡萄漿果內(nèi)糖和有機(jī)酸代謝及含量的相關(guān)影響。旨在為提高當(dāng)?shù)厣狡咸褲{果品質(zhì)和改善山葡萄生產(chǎn)管理模式提供科學(xué)的理論支持。主要取得以下結(jié)果:1.對山葡萄葉片光合及熒光相關(guān)指標(biāo)的變化研究發(fā)現(xiàn):隨著山葡萄的生長發(fā)育,漿果膨大期及轉(zhuǎn)色期山葡萄葉片的凈光合速率(Pn)值均呈現(xiàn)上升態(tài)勢,而始熟期山葡萄葉片的Pn呈略微下降趨勢。漿果膨大期和轉(zhuǎn)色期的氣孔導(dǎo)度(Gs)日變化趨勢較為接近,且三個關(guān)鍵時期的蒸騰速率(Tr)變化趨勢與Gs幾乎一致。漿果膨大期的呼吸速率(Pr)值保持最高水平,而始熟期的Pr值最低。山葡萄在成熟期時的胞間CO2濃度(Ci)值明顯高于其余兩個時期,而漿果膨大期和轉(zhuǎn)色期的Ci值差異不大。在山葡萄漿果發(fā)育前期葉片的最大熒光(Fm)日變化較為波動,但成熟期的Fm日變化相對比較平穩(wěn)。漿果膨大期山葡萄葉片在接近傍晚時仍在進(jìn)行高程度的電子傳遞;成熟期的Fm值卻略有下降,雖略有下降但未下降到最低值,表明此時光系統(tǒng)Ⅱ(PSⅡ)反應(yīng)中心完全關(guān)閉的熒光產(chǎn)量仍較高,其電子傳遞水平也較高。PSⅡ的最大光化學(xué)效率(Fv/Fm)與PSⅡ的潛在活性(Fv/Fo)比值在不同時刻出現(xiàn)急劇降低表明,此刻山葡萄葉片中的PSⅡ受到了傷害,光合作用原初反應(yīng)過程受抑制和影響。2.對轉(zhuǎn)色期和成熟期山葡萄漿果韌皮部非離體糖卸載研究體系及韌皮部后糖運(yùn)輸途徑的研究發(fā)現(xiàn):無論是轉(zhuǎn)色期還是成熟期,山葡萄非離體卸載體系漿果同化物糖分的輸入在正午之前均保持較高活性,主要積累的糖分是葡萄糖和果糖;蔗糖在這兩個階段均存在,并呈緩慢下降趨勢。轉(zhuǎn)色期和成熟期對山葡萄漿果組織進(jìn)行抑制劑處理,經(jīng)“新漿果杯”法及高效液相色譜法(HPLC)法測定漿果內(nèi)的糖組分及含量以及對完整葡萄漿果同化物輸入動態(tài)和非離體漿果同化物卸載動態(tài)進(jìn)行分析,揭示出山葡萄漿果韌皮部后糖運(yùn)輸?shù)耐緩街饕再|(zhì)外體卸載途徑為主,其限速關(guān)鍵時期主要為轉(zhuǎn)色期和成熟期。3.山葡萄漿果內(nèi)糖代謝相關(guān)酶活性及其糖分含量動態(tài)變化的研究發(fā)現(xiàn):在整個生長季,兩種架式(立架和棚架)山葡萄漿果的蔗糖磷酸合成酶(SPS)和合成方向的蔗糖磷酸(SS-s)活性均遠(yuǎn)遠(yuǎn)低于酶酸性轉(zhuǎn)化酶(SAI)和SS-s的活性,且隨果實發(fā)育進(jìn)程而增強(qiáng)。從生理角度來看,較低的sps和ss-s活性和較高的sai和ss-c活性可能是成熟山葡萄漿果含糖量低的主要原因,且架式對糖代謝相關(guān)酶的影響不明顯。4.對山葡萄漿果發(fā)育過程中有機(jī)酸代謝相關(guān)酶研究及有機(jī)酸含量動態(tài)變化研究發(fā)現(xiàn):棚架和立架山葡萄漿果在成熟期cs活性均比漿果膨大期高。立架和棚架的pepc變化趨勢近乎一致,除成熟期外,其余各個時期各個時間段內(nèi)棚架的pepc酶活性均比立架模式高。兩種架式的nad-idh活性變化差異十分顯著,其成熟期活性均高于漿果膨大期。轉(zhuǎn)色期是nad-mdh活性變化的轉(zhuǎn)折點(diǎn),在此之前所有棚架山葡萄漿果的nad-mdh活性均高于立架,而之后則相反。nadp-me的活性呈現(xiàn)先增長后下降趨勢,轉(zhuǎn)折點(diǎn)是轉(zhuǎn)色中期,整個nadp-me活性在該時間點(diǎn)達(dá)到峰值,同時所有棚架山葡萄漿果的nadp-me活性均高于立架模式。除漿果膨大期和成熟前期,其余時間段立架mit-aco的活性均高于棚架。成熟前期是cyt-aco活性變化的轉(zhuǎn)折點(diǎn),在此之前,立架的cyt-aco活性均高于棚架。酒石酸和蘋果酸在山葡萄整個生長季中的變化趨勢較為相似。檸檬酸含量的變化趨勢相對不夠穩(wěn)定,出現(xiàn)略微波動。不同架式下其檸檬酸含量的變化也存在差異。琥珀酸含量的變化無固定模式可循,兩種不同架式其各自琥珀酸的含量也不盡相同。5.新型閉合y型架式對山葡萄漿果內(nèi)糖分組成及糖代謝相關(guān)酶影響的研究表明:在生長發(fā)育過程尤其是轉(zhuǎn)色期至成熟期內(nèi)vsp和y型架式山葡萄漿果中的代謝酶sps、ss-c、ss-s及sai活性之間和漿果內(nèi)葡萄糖、果糖、蔗糖和乳糖含量之間存在一定差異。架式對葡萄糖和果糖的影響主要集中在轉(zhuǎn)色期之后直到成熟期,但對乳糖含量并無太大影響。vsp架式對蔗糖含量的影響主要集中在成熟期而y型架式對蔗糖含量的影響主要集中在轉(zhuǎn)色中期至成熟期。轉(zhuǎn)色前期,vsp和y型架式能夠通過調(diào)節(jié)sps和ss-c的活性進(jìn)而調(diào)節(jié)蔗糖的濃度;轉(zhuǎn)色后期,ss-c活性的輕微增加而sai的活性未受影響。轉(zhuǎn)色后期,兩種架式中的ss-s和sps都能夠調(diào)節(jié)其他糖分,如葡萄糖、半乳糖、麥芽糖、阿拉伯糖及果糖。6.不同光呼吸抑制劑nahso3和inh對山葡萄光合特性及單糖(ms)和二糖(ds)含量影響的研究表明:三種濃度(150、250、350mg/l)的nahso3處理均對山葡萄光呼吸有強(qiáng)烈抑制作用并能顯著提高光合產(chǎn)物(可溶性固形物ss,ms+ds)的產(chǎn)量及品質(zhì),且效果優(yōu)于inh的三個處理,并以250mg/l濃度表現(xiàn)最優(yōu)。inh能夠提高山葡萄葉片的光合熒光特性,但對光合產(chǎn)量的提高效果不明顯。7.不同負(fù)載對山葡萄漿果糖和有機(jī)酸代謝影響的研究發(fā)現(xiàn):山葡萄品種北冰紅和雙紅均屬低蔗糖類型。二者的蔗糖含量在轉(zhuǎn)色前期最高,隨后在漿果發(fā)育過程中逐漸被消耗直到漿果完全成熟。除成熟期時雙紅高負(fù)載處理外,兩種品種各處理間漿果內(nèi)果糖的含量變化與葡萄糖含量的變化幾乎一致。中負(fù)載處理的兩種山葡萄品種均含有各自最高的葡萄糖及果糖含量。不同負(fù)載對山葡萄漿果內(nèi)有機(jī)酸有較大影響:酒石酸和蘋果酸含量隨著漿果的成熟而逐漸降低,兩種品種中負(fù)載處理在成熟期時均含有各自最低含量的檸檬酸、蘋果酸以及酒石酸;北冰紅和雙紅在各時期,其漿果內(nèi)的蘋果酸含量均高于酒石酸含量,且中負(fù)載的蘋果酸含量低于低負(fù)載和高負(fù)載處理。不同負(fù)載對檸檬酸和琥珀酸含量并無明顯影響。綜合分析可知,無論是北冰紅還是雙紅山葡萄品種,中負(fù)載處理是得到漿果內(nèi)最優(yōu)糖酸含量的最佳選擇。
[Abstract]:Sugar and organic acid content is a key factor in determining the quality of grape and Wine quality. In actual production, micro environment can be adjusted by using different grape cultivation measures for growth, improve grape quality, so as to obtain high quality Wine. In this study, in Anchan mountain grape varieties Beibinghong and double red as test material, the photosynthetic characteristics and the berries in sugar and organic acid content and related metabolic enzymes were studied, and the different type, different loads and different respiratory inhibitors treatment on different cultivation measures on correlation of grape berry sugar and organic acid metabolism and to provide support for the content. The scientific theory to improve the local grape berry quality and improve the grape production management model. The main results are as follows: 1. of grape leaf photosynthesis and fluorescence correlation index change The study found that: with the grape growth, net photosynthesis during berry expansion period and mountain grape leaf color of photosynthetic rate (Pn) value showed a rising trend, while the beginning maturity grape leaf Pn showed a slightly decreasing trend. Berry expansion period and stomatal conductance of veraising (Gs day) the trend is more close, and the transpiration rate of three key periods (Tr) change trend with Gs is almost the same. Berry expansion period respiration rate (Pr) value to maintain the highest level, and initial maturity stage of the lowest Pr. Grape maturity in intercellular CO2 concentration (Ci) value was significantly higher than the other the two period, and berry enlargement stage and color changing of Ci value had little difference. In the early development of grape berry fluorescence maximum leaf (Fm) diurnal variation more volatile, but Fm diurnal variation of maturity is relatively smooth. Berry expansion period mountain grape leaves near dusk is still in the high degree of electronic Transfer; mature Fm value declined slightly, although a slight decline but did not decrease to the minimum value, show that this time system II (PS II) fluorescence yield completely closed reaction center is still high, the electron transfer will have a higher level of maximum photochemical efficiency of.PS (Fv/Fm) and potential activity of PS II (Fv/Fo) the ratio in different time decreases sharply at the moment that grape leaves in PS II was hurt by the inhibition of photosynthetic primary reaction process and the effect of.2. on the color turning stage and mature stage of grape berry phloem in vitro non sugar unloading research system and Postphloem sugar transport pathways were found that the color change stage or maturity, non isolated mountain grape berries assimilation sugar unloading system input and maintain a high activity before noon, the main accumulation of sugar is glucose and fructose; sucrose in these two stages are kept in, and slowly down Drop trend. Color turning stage and mature stage of inhibitor treatment on grape berries, the "new berry Cup" method and high performance liquid chromatography (HPLC) method for the determination of berry sugar composition and content of grape berry complete assimilation input dynamic and non isolated berry assimilate unloading dynamics analysis revealed from grape berry Postphloem sugar transport pathway mainly in the apoplastic unloading pathway, the rate limiting critical period is mainly veraison and maturity of.3. glucose metabolism related enzyme activity in grape berries and dynamic changes of sugar content found in the whole growth season, the two type (Li frame and scaffolding) sucrose phosphate synthase grape berries (SPS) and synthesis direction of sucrose phosphate (SS-s) enzyme activity were far lower than the acid invertase (SAI) and the activity of SS-s, and increased with fruit development process from the physical point of view, Sai and SS-C activity and lower SPS and higher ss-s activity may be the main reason of mature grape berries with low sugar content, and the influence of posture on enzymes related to sugar metabolism was.4. on organic acid metabolism in grape berry development of enzyme and organic acid content dynamic change research found: scaffolding and stand mountain grape berries in CS activity than mature berry expansion period high. The change trend of PEPC stand and the scaffolding was almost the same, in addition to mature stage, PEPC activity in the rest of each period of each period of time are high than scaffolding frame model. The difference of NAD-IDH activity changes of two kinds of posture is very significant, its maturity the activity was higher than that of veraison berry expansion period. The change of nad-mdh activity is a turning point, before all the scaffolding mountain grape berry nad-mdh activity was higher than the vertical frame, and then the opposite.Nadp-me activity showed first increase After the decline, the turning point is the mid turn color, the activity of nadp-me at the time point and reached the peak at the same time, all scaffolding Vitis amurensis berry nadp-me activity were higher than that of vertical frame mode. In addition to Berry expansion period and mature period, the remaining time of the activity of mit-aco was higher than that of vertical frame scaffolding. Early maturing period is the turning point, the activity of cyt-aco change prior to this, the activity of cyt-aco was higher than that of vertical frame scaffolding. Tartaric acid and malic acid are similar in the whole growth trend of grape Ji Zhong. The change trend of citric acid content is not stable, slightly fluctuations. Different changes in the type of citric acid content is also different. The changes of the content of succinic acid fixed pattern, the research content of two kinds of different type of their respective succinic acid are not the same type.5. type Y type closure related enzyme effect on grape berry sugar composition and glucose metabolism Show: SS-C in the process of growth and development especially metabolizing enzymes SPS, colour changed stage to mature stage of VSP and Y type mountain in grape berries, glucose, ss-s and Sai activity and berries in fructose, there are some differences between the sucrose and lactose content. Effects of type of glucose and fructose are mainly concentrated in the turn after color until maturity, but the content of lactose have little effect on the effect of.Vsp type sucrose content mainly concentrated in the mature period and the effect of Y type on sucrose content mainly concentrated in the middle turn color to maturity. Color stage, the concentration of VSP and Y type frame type can through regulating SPS and SS-C activity and then adjust the sucrose; color changing period, slightly increased SS-C activity and Sai activity were not affected. Color change and SPS two ss-s later, the rack can be adjusted to other sugars, such as glucose, galactose and maltose, glucose and fructose in Arabia .6. and INH on photorespiration inhibitor NaHSO3 grape photosynthetic characteristics and monosaccharides (MS) and two (DS) of sugar content effects showed that three concentration (150250350mg/l) of the NaHSO3 treatments on grape photorespiration have an inhibitory effect and can significantly improve the photosynthetic products (soluble solids SS, ms+ds) the yield and quality, and the three treatment effect is better than that of INH, and the concentration of 250mg/l.Inh can be best to improve grape leaf fluorescence light, but photosynthetic yield increasing effect is not obvious.7. different load on mountain grape pulp fructose and affect the metabolism of organic acid. Mountain grape varieties Beibinghong and double red are low sucrose type. The sucrose content of the two highest early in the color turning, then during berry development is gradually consumed until berries fully mature. In addition to the mature red double high processing load, two varieties of each treatment The content change of change and the content of glucose in fructose between berries are nearly same. Two kinds of grape varieties of processing load contain their highest glucose and fructose content. Different load has a great influence on the grape berries in organic acid, tartaric acid and malic acid content decreased with the fruit mature, two kinds of load the treatment of citric acid contained the lowest content in their maturity varieties, malic acid and tartaric acid; Beibinghong and double red in each period, the content of malic acid in the berries were higher than tartaric acid content, low load and high load is lower than the content of malic acid load. And different load effect on citric acid and succinic acid. There is no comprehensive analysis shows that, whether Beibinghong or double Hongshan grape varieties, in the processing load is the best choice to get the optimal berry sugar and acid content.

【學(xué)位授予單位】：西北農(nóng)林科技大學(xué)
【學(xué)位級別】：博士
【學(xué)位授予年份】：2016
【分類號】：S663.1
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本文編號：1405004