本文選題：鎘　切入點(diǎn)：蘋果砧木　出處：《沈陽(yáng)農(nóng)業(yè)大學(xué)》2017年博士論文　論文類型：學(xué)位論文

【摘要】：隨著人民生活水平的不斷提高,果品在人類膳食結(jié)構(gòu)中所占比例越來(lái)越大。我國(guó)是世界蘋果的主產(chǎn)國(guó),蘋果在我國(guó)居民水果消費(fèi)中占有重要地位。然而,污水灌溉及化肥農(nóng)藥的大量施用使得果園土壤重金屬污染問題日益突出,尤其是Cd污染。嫁接是果樹繁殖的重要方式,砧木是果樹生產(chǎn)的承接者,對(duì)Cd的吸收、轉(zhuǎn)運(yùn)能力直接影響果樹整體Cd耐性及接穗中Cd含量。因此,研究蘋果砧木對(duì)Cd的吸收、轉(zhuǎn)運(yùn)以及耐性差異,可為篩選低Cd富集耐性砧木提供理論依據(jù)。本試驗(yàn)首先研究了山定子、平邑甜茶、青州林檎和八棱海棠根系Cd2+吸收的時(shí)空動(dòng)態(tài)變化規(guī)律;隨后在生理和分子水平探討了Cd在蘋果砧木各組織轉(zhuǎn)運(yùn)和對(duì)Cd的耐性機(jī)理;接著研究褪黑素緩解Cd對(duì)蘋果砧木的毒害機(jī)理;最后探討了不同蘋果嫁接苗對(duì)Cd的吸收、轉(zhuǎn)運(yùn)和耐性差異。主要結(jié)果如下:1.通過比較4種蘋果砧木根系在50 μM Cd處理下Cd2+吸收的時(shí)空動(dòng)態(tài)變化規(guī)律和Cd2+在植物體的轉(zhuǎn)運(yùn),發(fā)現(xiàn)蘋果砧木白根根尖600 μm處Cd2+凈流量最大,隨著距根尖距離的延長(zhǎng),Cd2+凈流量逐漸降低。山定子根系和葉柄維管束Cd2+的凈流量低于其他3種蘋果砧木。代謝抑制劑和通道阻斷劑(正釩酸鈉、維拉帕米和NEM)處理顯著降低了 4種蘋果砧木根系Cd2+的凈流量。另外,環(huán)境Fe3+濃度也影響根系Cd2+的凈流量。除山定子外,Cd導(dǎo)致其他3種蘋果砧木凈光合速率和生物量增量減少。與其他3種蘋果砧木相比,Cd脅迫下山定子各組織Cd含量、Cd富集總量,BCF和Tf最小,參與Cd吸收和轉(zhuǎn)運(yùn)基因如HA4、FRO2-like、NRAMP1、NRAMP3和HMA4轉(zhuǎn)錄水平較低,參與Cd解毒相關(guān)基因如NAS1和MT2表達(dá)較強(qiáng)。由此表明,50 μM Cd處理下山定子對(duì)Cd吸收和轉(zhuǎn)運(yùn)的能力低于其他3種蘋果砧木。2.對(duì)4種蘋果砧木進(jìn)行較高濃度Cd(300 μM)處理45 d后,所有砧木幼苗的凈光合速率、葉綠素含量和生物量均受到了抑制,但對(duì)山定子的影響較小,且山定子整株耐性指數(shù)(TI)顯著高于其他3種蘋果砧木。4種蘋果砧木中,山定子根、木質(zhì)部和葉片中Cd含量最低、Cd富集總量和BCF最小。蘋果砧木能將Cd固持在細(xì)胞壁和液泡中,并將其轉(zhuǎn)化成果膠酸鹽或蛋白結(jié)合態(tài)以及難溶性磷酸鹽狀態(tài)來(lái)降低Cd的移動(dòng)性和毒性。4種蘋果砧木生理狀態(tài)如ROS、糖類和抗氧化系統(tǒng)的變化與對(duì)Cd的耐性相一致。與其他3種蘋果砧木相比,山定子根和韌皮部中ROS、根和葉片中H202以及根、木質(zhì)部和韌皮部中MDA含量最低,韌皮部中可溶性糖以及根和葉片淀粉含量最高,抗氧化酶活性和抗氧化物含量較高。說(shuō)明該試驗(yàn)條件下山定子對(duì)Cd的耐受性強(qiáng)于其他3種蘋果砧木,這與山定子各組織Cd含量、Cd的亞細(xì)胞分布和化學(xué)形態(tài),以及抗氧化系統(tǒng)有關(guān)。3.通過比較外源褪黑素對(duì)30μM Cd處理下山定子和青州林檎幼苗Cd吸收和轉(zhuǎn)運(yùn)、抗氧化系統(tǒng)及基因表達(dá)的影響,發(fā)現(xiàn)添加褪黑素后山定子和青州林檎根系Cd2+的凈吸收速率、葉片Cd含量以及Tf顯著降低,說(shuō)明褪黑素抑制了蘋果砧木對(duì)Cd的吸收和轉(zhuǎn)運(yùn),且對(duì)山定子的抑制作用更明顯。與30μMCd處理相比,30μMCd + 0.1μM褪黑素處理減少了蘋果砧木根、莖和葉中O2·-、H202和MDA含量,細(xì)胞器組分Cd含量和所占比例降低,以及Cd吸收相關(guān)基因如NARMAP1表達(dá)量降低;抗氧化酶如CAT、GPX、APX和GR活性升高,根和葉片中游離脯氨酸、根中可溶性酚和T-SH、莖和葉片中ASC,根和莖中GSH以及根中褪黑素含量增加,與Cd解毒相關(guān)基因如NAS1表達(dá)量上調(diào),說(shuō)明褪黑素能夠抑制蘋果砧木根系吸收和轉(zhuǎn)運(yùn)Cd的能力,提高細(xì)胞抗氧化水平,有效清除細(xì)胞內(nèi)的ROS,增強(qiáng)蘋果砧木對(duì)Cd的耐受性。但30μM Cd + 0.1 μM褪黑素處理下青州林檎根和葉片中CAT和GR、根中GPX和APX活性,以及根和葉片中游離脯氨酸、葉片可溶性酚和T-SH、莖和葉片GSH含量低于山定子;參與Cd吸收的基因如HA7和NRAMP1的轉(zhuǎn)錄水平高于山定子,與Cd解毒相關(guān)基因如NAS1表達(dá)量低于山定子。說(shuō)明褪黑素對(duì)緩解山定子Cd脅迫的作用更明顯。4.通過比較50μM Cd脅迫下4種蘋果嫁接苗對(duì)Cd的吸收和轉(zhuǎn)運(yùn)、抗氧化水平及基因表達(dá)的變化,發(fā)現(xiàn)Cd導(dǎo)致4種蘋果嫁接苗的生物量、光合速率和色素含量降低,但Cd對(duì)寒富/山定子的影響最小。寒富/山定子和富士/山定子根系和接穗Cd含量、Cd富集總量和BCF分別小于寒富/青州林檎和富士/青州林檎,說(shuō)明以山定子為砧木能有效減少Cd在蘋果嫁接苗中的富集。Cd脅迫下寒富/山定子木質(zhì)部、韌皮部和葉片Cd含量及水溶態(tài)Cd含量低于其他3種蘋果嫁接苗,根和葉片細(xì)胞壁組分Cd所占比例最大,細(xì)胞器組分Cd含量和比例最小,與Cd吸收和轉(zhuǎn)運(yùn)相關(guān)基因如HA7、FRO2-like、NRAMP1和NRAMP3表達(dá)量較低,說(shuō)明寒富/山定子根系固持Cd的能力較強(qiáng),能降低Cd在嫁接苗中的移動(dòng)速率。另外,Cd脅迫下寒富/山定子韌皮部和葉片ROS和MDA含量最低,葉片可溶性酚、ASC和GSH含量最高,與Cd解毒相關(guān)的基因如MT2表達(dá)量最高,說(shuō)明寒富/山定子對(duì)Cd的忍耐性強(qiáng)于其他3種蘋果嫁接苗。
[Abstract]:With the improvement of people's life, the fruit in the human diet in the proportion is increasing. China is the world's leading producers of apple, apple occupies an important position in China's residents consumption of fruit. However, the extensive application of sewage irrigation and fertilizer and pesticide that orchard soil heavy metal pollution problems have become increasingly prominent, especially Cd pollution. Grafting is an important way of fruit breeding, rootstock is to undertake the production of fruit trees, the absorption of Cd, directly affect the content of Cd and Cd in the whole fruit tolerance scion transport capacity. Therefore, the research on absorption of Apple Rootstock Cd, transport and tolerance, which can provide the theoretical basis for screening low Cd enrichment Tolerant Rootstock this test studied first baccata, Pingyi Tiancha, temporal variation. Roots Cd2+ absorption of Qingzhou LinQin and eight ribs; then in physiological and molecular level of Cd in Guo Zhenmu. The organization of transport and Cd tolerance mechanism; then study melatonin alleviates the poisoning mechanism of Cd on Apple Rootstock; finally discusses the absorption of different apple grafted on Cd, translocation and tolerance differences. The main results are as follows: 1. by comparing the 4 kinds of Apple Rootstock Roots in 50 M Cd treatment Cd2+ absorption dynamic changes of Cd2+ in plants and transport, found the net Cd2+ 600 m flow of Apple Rootstock root increased with white maximum distance from root tip, Cd2+ net flow decreased gradually. The net flow of Cd2+ root and petiole vascular bundle is lower than other 3 kinds of crab apple rootstocks. Metabolic inhibitors and channel blockers (orthovanadate Na, Vera Pammy and NEM) significantly decreased the net flow of 4 kinds of Apple Rootstock Roots of Cd2+. In addition, Fe3+ concentration also affects the net flow of root Cd2+. Except Cd baccata, lead to other 3 kinds of net photosynthetic rate of Apple Rootstocks The rate and biomass increment decreased. Compared with the other 3 kinds of apple rootstocks, Cd stress down tissue Cd content stator, Cd enrichment amount, BCF and Tf was the smallest, involved in Cd uptake and transport genes such as HA4, FRO2-like, NRAMP1, NRAMP3 and HMA4 transcription level is low, involved in Cd detoxification related genes such as NAS1 and MT2 expression strong. It showed that 50 M Cd processing ability of Cd stator down the absorption and transport of lower than the other 3 types of Apple Rootstock.2. high concentration Cd on 4 rootstocks (300 M) after 45 D, the net photosynthetic rate of all rootstock seedlings, chlorophyll content and biomass was inhibited however, the impact of Yama Joko and Yama Joko is small, the whole plant tolerance index (TI) Yama Joko root was significantly higher than the other 3 kinds of Apple Rootstock.4 apple rootstocks, and Cd content of the xylem and leaves the lowest, Cd and BCF. The minimum amount of Apple Rootstock Cd can be fixed in the cell wall and Vacuole, and convert the results of gel acid salt or protein bound and soluble state of Cd to reduce the mobility and toxicity of.4 Apple Rootstocks with physiological state changes such as ROS, sugar and antioxidant system consistent with the Cd tolerance. Compared with other 3 kinds of Apple Rootstock Roots and phloem cells in ROS, the root and leaf H202 and root MDA content in xylem and phloem of the lowest in the phloem of root and leaf soluble sugar and starch content is the highest, higher antioxidant enzyme activity and antioxidant content. The test under the condition of strong tolerance to the stator Cd than the other 3 apple rootstocks, and the the Cd content of the tissue cells, subcellular distribution and chemical forms of Cd, and the.3. antioxidant system by comparing the effects of exogenous melatonin on the 30 M Cd down the stator and Qingzhou LinQin seedling Cd uptake and translocation, antioxidant system and medium Because of the influence on the expression of melatonin, found that adding Malus baccata and Qingzhou Cd2+ net of mallus doumeri root absorption rate, leaf Cd content and Tf decreased significantly, indicating that melatonin inhibits the absorption and transport of Cd Apple Rootstock, and the inhibition of m.baccata was more obvious. Compared with 30 MCd, 30 MCd + 0.1 M melatonin treatment reduces the Apple Rootstock Roots, stems and leaves of O2, H202 and MDA content, Cd content and cell component proportion decreased, and the absorption of Cd related genes such as NARMAP1 expression decreased; antioxidant enzymes such as CAT, GPX, APX increased and GR activity, free proline in roots and leaves. Soluble phenol and T-SH in root, stem and leaf ASC, increase in root and stem and root GSH in melatonin content, and Cd toxin genes such as NAS1 up-regulated, indicating that melatonin can inhibit the ability of Apple Rootstock root uptake and translocation of Cd cells, improve the antioxidant The level of effective removal of intracellular ROS, enhance the tolerance of Apple Rootstocks on Cd. But 30 M Cd + 0.1 M melatonin treatment CAT and GR in roots and leaves of Malus in Qingzhou, GPX and APX activity in roots, and the roots and leaves of free proline, soluble phenol and T-SH, and stem leaf GSH content is lower than in Malus baccata; Cd absorption such as gene transcription of HA7 and NRAMP1 in the high mountain stator, and Cd detoxification related genes such as NAS1 low expression in the mountains. The stator of melatonin on Cd stress relieving m.baccata more obvious by comparing the.4. 50 M Cd under the stress of the absorption and transport of Cd 4 kinds of Apple grafted seedlings, the expression level and antioxidant genes, found that Cd causes 4 kinds of apple seedlings biomass, photosynthetic rate and pigment content decreased, but the influence of Cd on Hanfu Hanfu. The minimum / m.baccata / baccata and Fuji / Cd content and scion root cells, C D and BCF were less than the amount of Hanfu / Qingzhou / Qingzhou Fuji mallus doumeri and mallus doumeri, which can effectively reduce the Cd baccata as rootstock grafted seedlings in apple.Cd enrichment under the stress of Hanfu expressed in xylem, phloem and leaf Cd content and water soluble Cd content is lower than the other 3 kinds of apple seedlings, root and leaf cell wall components Cd the largest proportion of organelles component content of Cd and the ratio of the minimum, associated with the absorption and transportation of Cd genes such as HA7, FRO2-like, NRAMP1 and NRAMP3 low expression, indicating Hanfu / root Cd strong ability of holding cells, can reduce the rate of the mobile Cd in grafted seedlings in addition, under the stress of Cd / m.baccata Hanfu phloem and leaf ROS and MDA content was the lowest, soluble phenol, ASC and GSH were the highest, and Cd toxin genes such as the expression of MT2 was highest, that of Hanfu / strong tolerance to Cd in which he expressed 3 kinds of apple Grafted seedlings.

【學(xué)位授予單位】：沈陽(yáng)農(nóng)業(yè)大學(xué)
【學(xué)位級(jí)別】：博士
【學(xué)位授予年份】：2017
【分類號(hào)】：S661.1

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