【摘要】：鳶尾屬(Iris L.)是鳶尾科(Iridaceae)所有屬中涵蓋種類最多的一個(gè)屬,據(jù)統(tǒng)計(jì)目前已知的鳶尾屬植物共計(jì)300余種。鳶尾屬植物生長的氣候帶以北溫帶為主,分布的區(qū)域集中在亞洲、歐洲及北美洲。我國也是鳶尾屬植物的分布中心之一,已知的種有64個(gè),變種13個(gè),此外還有1個(gè)亞種和6個(gè)變型。鳶尾屬植物屬多年生草本類,花色豐富(有單色也有復(fù)色),花型多變(旗瓣與垂瓣形態(tài)、大小比例等多變),是聞名于世的宿根花卉之一。隨著鳶尾屬植物園藝品種的增多,其遺傳背景愈發(fā)復(fù)雜。本研究以鳶尾屬園藝品種和種共38份材料,利用形態(tài)學(xué)標(biāo)記結(jié)合ISSR和SRAP兩種分子標(biāo)記方法對(duì)其遺傳多樣性及親緣關(guān)系進(jìn)行了評(píng)價(jià)分析。另外,結(jié)合鳶尾屬植物的生產(chǎn)應(yīng)用,測(cè)定了鳶尾屬部分材料高溫、干旱脅迫下的生理響應(yīng)與分子機(jī)制,建立了鳶尾屬植物的耐熱、抗旱生理評(píng)價(jià)體系,并對(duì)實(shí)驗(yàn)材料的耐熱、抗旱能力進(jìn)行了評(píng)價(jià)。本研究獲得主要結(jié)果如下:1、鳶尾屬參試材料觀測(cè)的29個(gè)表型性狀中,變異系數(shù)差異較大。對(duì)表型性狀主成分分析得出,基于表型性狀分類時(shí)首先應(yīng)考慮花器官性狀的分類,其次是根據(jù)株型、營養(yǎng)器官生長以及花瓣上附屬物的情況分類。主成分分析的結(jié)果也反映出表型性狀的主成分貢獻(xiàn)率集中度不明顯,其中的累積貢獻(xiàn)率呈現(xiàn)緩慢的增長,據(jù)此推測(cè)其中主要是相關(guān)性強(qiáng)的變量較分散的原因,而性狀分別屬于多類,由此也說明鳶尾屬的表型性狀在演化中的多樣性十分明顯,這也是其存在著豐富性狀變異的主要原因。2、ISSR和SRAP這兩種標(biāo)記方法的多態(tài)性條帶比率均為100%,但SRAP較ISSR檢測(cè)出的條帶數(shù)高,兩種方法的多態(tài)性含量PIC值均為1,說明鳶尾屬種質(zhì)資源遺傳差異較大,在分子水平具有非常高的遺傳多樣性。另外,兩種分子標(biāo)記對(duì)鳶尾屬植物的遺傳多樣性分析具有較高的一致性,且ISSR和SRAP相結(jié)合的方法比單一的ISSR或SRAP分子標(biāo)記對(duì)鳶尾屬植物遺傳多樣性分析的結(jié)果與表型分類結(jié)果更為接近。采用ISSR和SRAP兩種方法相結(jié)合進(jìn)行分子標(biāo)記聚類,其遺傳距離矩陣進(jìn)行Mantel檢驗(yàn)得出相關(guān)性系數(shù)r=0.8052,可見兩種標(biāo)記方法結(jié)合的結(jié)果較為可靠,也能更好的反映出鳶尾屬植物的遺傳結(jié)構(gòu),可為其他1ii相關(guān)研究提供參考。3、前人很多研究表明鳶尾屬植物種間雜交存在有一定的障礙,雜交結(jié)實(shí)率很低或不結(jié)實(shí),而本研究中反應(yīng)出德國鳶尾‘Gala Madrid’和‘Nibelungen’在有髯鳶尾中遺傳距離與其他德國鳶尾品種較遠(yuǎn),且分子標(biāo)記的三種方式聚類中均與無髯鳶尾聚為一類,由此推測(cè)這兩種鳶尾很有可能具有無髯鳶尾的遺傳背景,可以將其作為鳶尾屬植物無髯與有髯鳶尾類種間雜交的優(yōu)勢(shì)親本進(jìn)行種質(zhì)創(chuàng)新。4、對(duì)雞冠狀附屬物亞屬在分類中存在的爭(zhēng)議,通過本研究支持其按照Rodionenko系統(tǒng)作為亞屬的分類地位存在。另外,琴瓣鳶尾亞屬的喜鹽鳶尾三種方法聚類后都與無附屬物亞屬的種聚在了一起,可見將喜鹽鳶尾作為琴瓣鳶尾單獨(dú)成為一個(gè)亞屬存在的意義不大,建議將其歸入無附屬物亞屬中。另外,無附屬物亞屬中馬藺、黃菖蒲、路易斯安那鳶尾都單獨(dú)聚為亞類群,與其它鳶尾關(guān)系較遠(yuǎn)單獨(dú)成一組,結(jié)合其形態(tài)學(xué)的差異,建議可將無附屬物亞屬分成幾個(gè)組。5、不同鳶尾品種(種)在高溫和干旱脅迫下,測(cè)定部分氧化酶和非酶活性物質(zhì)生理生化指標(biāo),利用主成分分析法結(jié)合隸屬函數(shù)法分別對(duì)測(cè)定指標(biāo)與耐熱、抗旱的相關(guān)性進(jìn)行評(píng)價(jià)。結(jié)果表明,電導(dǎo)傷害率(x1)、葉綠素(x3)、可溶性糖(x4)及丙二醛(x5)4個(gè)指標(biāo)與鳶尾屬植物抗旱能力強(qiáng)弱的相關(guān)性較高;SOD活性(x1)、POD活性(x2)、葉綠素(x3)、可溶性蛋白(x4)及丙二醛(x5)5個(gè)指標(biāo)與鳶尾屬植物耐熱能力相關(guān)性較高。進(jìn)一步采用逐步回歸法結(jié)合相關(guān)性高的生理生化指標(biāo)建立了鳶尾屬植物抗旱、耐熱評(píng)價(jià)最優(yōu)回歸方程�？购底顑�(yōu)回歸方程:D=-439.27+1.744x1+4.913x3+0.5822x4+2.6369x5,耐熱最優(yōu)方程:D=29.667-0.208x1-2.502 x2+0.405x3-1.742 x4-0.036 x5。依據(jù)抗旱、耐熱最優(yōu)回歸方程供試的10個(gè)鳶尾品種(種)抗旱性由強(qiáng)到弱的順序依次為‘金娃娃’、‘笛聲’、‘魂斷藍(lán)橋’、德國鳶尾、扁竹蘭、西伯利亞鳶尾、馬藺、‘黑騎士’、黃菖蒲和花菖蒲;耐熱性由強(qiáng)到弱的順序依次為:花菖蒲、黃菖蒲、馬藺、扁竹蘭、‘笛聲’、‘黑騎士’、‘魂斷藍(lán)橋’、‘金娃娃’、西伯利亞鳶尾和德國鳶尾。6、從觀賞性較高且同為德國鳶尾系列的4個(gè)德國鳶尾中分別篩選出2個(gè)耐熱性相對(duì)較差的‘金娃娃’與德國鳶尾通過Ca Cl2處理,以及2個(gè)耐熱性相對(duì)好的的品種‘笛聲’和‘黑騎士’通過Ca2+的拮抗劑La Cl3處理,測(cè)定Ca2+對(duì)于鳶尾高溫脅迫下的耐熱性的調(diào)控作用。結(jié)論是Ca Cl2較為明顯的提高了高溫脅迫下金娃娃與德國鳶尾葉片中的可溶性蛋白質(zhì)的濃度,這可能與Ca2+信號(hào)介導(dǎo)的熱激蛋白的產(chǎn)生有關(guān),使其在較短時(shí)間內(nèi)可保持對(duì)高溫的抗性,但高溫脅迫時(shí)間超過其耐受能力,則呈現(xiàn)葉片焦枯逐漸死亡;La Cl3作為Ca2+阻斷劑對(duì)笛聲和黑騎士葉片的耐熱性影響顯著;由此,推測(cè)笛聲和黑騎士依賴Ca2+信號(hào)介導(dǎo)來產(chǎn)生耐熱性。7、根據(jù)基因庫模式植物的熱激因子基因的保守區(qū)域設(shè)計(jì)引物序列,通過PCR克隆得到鳶尾熱激因子基因的片段;利用熒光實(shí)時(shí)定量PCR檢測(cè)不同鳶尾品種在不同溫度脅迫與不同高溫脅迫時(shí)間下熱激因子基因轉(zhuǎn)錄表達(dá)水平的變化。結(jié)果顯示耐熱性較高的笛聲和黑騎士熱激因子轉(zhuǎn)錄水平高于耐熱性差的金娃娃與德國鳶尾;同時(shí),Ca2+信號(hào)參與了這四個(gè)鳶尾品種葉片的熱激因子轉(zhuǎn)錄表達(dá)的調(diào)控。
[Abstract]:Iris L. is one of the most species of all genera of the family irisfamily (Iridaceae). According to the statistics, there are more than 300 species of iris. The climate zone of the genus Iris is mainly in the north temperate zone, which is distributed in Asia, Europe and North America. China is also one of the distribution centers of the Iris plants. There are 64 species, 13 varieties, and 1 subspecies and 6 variants. Iris is a perennial herb with rich color (monochromatic color) and one of the most famous perennial flowers in the world. With the increase of Iris plants, the genetic background of iris is more complex. In this study, the genetic diversity and relationship of 38 species of Iris species and species were evaluated by two molecular markers combined with morphological markers combined with ISSR and SRAP. In addition, the physiological responses and molecular machines under drought stress were measured in combination with the production and application of Iris plants. The system of heat resistance and drought resistance evaluation of Iris plants was established, and the heat resistance and drought resistance of the experimental materials were evaluated. The main results of this study were as follows: 1, the variation coefficient of the 29 phenotypic characters observed in iris was large. The principal component analysis of phenotypic traits was based on the first classification of phenotypic traits. First, the classification of flower organs should be considered, followed by the classification of the plant type, the growth of the vegetative organs and the appendages on the petals. The results of the principal component analysis also reflect that the concentration of the principal component contribution rate of the phenotypic traits is not obvious, and the cumulative contribution rate of the phenotypic traits is slow. Because of the more scattered reasons, the characters of the characters belong to multiple categories, which also shows that the diversity of the phenotypic traits of the iris is very obvious. This is the main reason for the existence of.2, ISSR and SRAP, which are all two of the polymorphic bands of 100%, but the number of bands detected by SRAP is higher than that of ISSR, and two methods are found. The PIC value of polymorphic content is 1. It shows that the genetic diversity of iris germplasm is large and has very high genetic diversity at the molecular level. In addition, the two molecular markers have high consistency in the genetic diversity analysis of Iris plants, and the method of combining ISSR and SRAP is better than single ISSR or SRAP molecular markers. The results of genetic diversity analysis of plants are more close to the results of phenotypic classification. The combination of two methods of ISSR and SRAP is used to cluster molecular markers, and the genetic distance matrix is used to test the correlation coefficient r=0.8052 by Mantel test. It can be seen that the results of the combination of the two methods are more reliable, and can also better reflect the remains of the iris plant. The transmission structure can provide reference for other 1ii related studies. A lot of previous studies have shown that there are some obstacles in the interspecific hybridization of iris, and the hybrid seed setting rate is very low or not strong. In this study, the genetic distance of the German iris' Gala Madrid 'and' Nibelungen 'in the beard kite is far from the other German iris varieties. The three types of molecular markers are all clustered with the beard iris. Therefore, it is presumed that these two kinds of iris are likely to have the genetic background of the beard irises. They can be used as the dominant parent of the hybrid irises between the irises and the bearded irises to carry out the germplasm innovation.4, and the contention of the subgenus of the coronal appendage of the chicken in the classification. According to this study, it supports the classification status of the subgenera according to the Rodionenko system. In addition, the three methods of halophilic iris in the genus Iris are clustered together with non appendage subgenera. It is obvious that it is not of great significance to make iris iris as a subgenus as a single subgenus. In addition, the subgenus of appendages, iris iris, Huang Changpu and Louisiana irises are separated into subgroups separately, and separate from other irises. According to their morphological differences, the subgenus of non appendages can be divided into several groups of.5, and different iris varieties (species) can determine partial oxidation under high temperature and drought stress. The physiological and biochemical indexes of enzymes and non enzyme active substances were evaluated by principal component analysis and membership function method respectively. The results showed that the 4 indexes of conductance damage rate (x1), chlorophyll (x3), soluble sugar (x4) and malondialdehyde (x5) were more related to the drought resistance of Iris Plants, SOD The 5 indexes of activity (x1), POD activity (x2), chlorophyll (x3), soluble protein (x4) and malondialdehyde (x5) were highly correlated with the heat tolerance of Iris plants. The optimum regression equation for drought resistance and heat tolerance evaluation of Iris plants was established by stepwise regression method combined with high correlation physiological and biochemical indexes. The optimal regression equation of drought resistance: D=-439.27+1.7 44x1+4.913x3+0.5822x4+2.6369x5, the optimum heat tolerance equation: the drought resistance of the 10 varieties (species) of D=29.667-0.208x1-2.502 x2+0.405x3-1.742 x4-0.036 x5., based on the optimum regression equation of drought resistance and heat resistance, is in sequence of "gold dolls", 'flute sound', 'soul blue bridge', German iris, flat bamboo orchid, and Siberia iris. Iris, 'Black Knight', Huang Changpu and flower calamus; the order of heat resistance from strong to weak: Flower calamus, Huang Changpu, iris, bamboo orchid, 'flute', 'Black Knight', 'soul blue bridge', 'gold doll', Siberia iris and German iris.6, respectively sifting from 4 German irises with a higher appreciation and German iris series. 2 relatively poor heat-resistant 'gold dolls' and German irises were treated by Ca Cl2, and 2 varieties with relatively good heat resistance were treated by Ca2+'s antagonist La Cl3. The result was that Ca2+ was used to regulate the heat resistance of iris under high temperature stress. The conclusion was that Ca Cl2 increased the high temperature obviously. The concentration of soluble protein in the gold dolls and the leaves of German iris may be related to the production of heat shock protein mediated by Ca2+ signal, which can keep the resistance to high temperature in a short time, but the time of high temperature stress exceeds its tolerance, while La Cl3 is a Ca2+ blocker to the flute and the sound of the flute. The heat resistance of the Black Knight leaves is significant. Therefore, it is speculated that the flute and the Black Knight rely on the Ca2+ signal to produce heat resistance.7. According to the conserved region of the heat shock factor gene of the gene bank model plant, the primer sequence is designed and the segments of the iris heat shock factor gene are obtained by PCR cloning, and the different iris are detected by real time fluorescence quantitative PCR The transcriptional level of heat shock factor gene in different temperature stress and high temperature stress time was observed. The results showed that the transcriptional level of the flute and the Black Knight heat shock factor was higher than that of the poor heat resistant gold dolls and German irises, and the Ca2+ signal was involved in the transcriptional expression of the heat shock factor of the leaves of the four irises Regulation and control.
【學(xué)位授予單位】：華中農(nóng)業(yè)大學(xué)
【學(xué)位級(jí)別】：博士
【學(xué)位授予年份】：2016
【分類號(hào)】：S682.19

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