本文選題：胡楊　切入點：異形葉　出處：《中央民族大學》2016年博士論文　論文類型：學位論文

【摘要】：本文研究對象為胡楊(Populus euphratica Oliv.),又稱幼發(fā)拉底河楊樹或沙漠楊樹,屬于楊柳科楊屬,是第三紀以來古地中海地區(qū)形成的最古老的植物。目前胡楊主要分布在中亞、西亞、北非到南歐(大約在北緯30°～50°之間)。胡楊是中亞和中國西北干旱荒漠地區(qū)唯一天然分布的喬木樹種,中國是世界上胡楊分布面積最廣、數(shù)量最多的國家,胡楊在我國主要分部在新疆維吾爾自治區(qū)塔里木盆地、輪臺縣和內(nèi)蒙古阿拉善盟額濟納地區(qū)。胡楊生長期漫長,由于風沙和干旱的影響,很多胡楊樹造型奇特、詭異,有句古語形容它“活三千年不死,死三千年不倒,倒三千年不朽”�！谖覈贁�(shù)民族文化中,胡楊被視為“神木”或“圣樹”,并作為死亡和復活的象征總是和鮮血與鳳凰緊密相連。胡楊在生長發(fā)育過程中表現(xiàn)出顯著的異形葉特征。胡楊幼樹葉形均為細長狀,隨著生長發(fā)育,植株上部的葉片逐漸變短變寬,最后形成兩側(cè)帶鋸齒狀的闊葉形。成年胡楊同一植株上的葉形有披針形、橢圓形、卵圓形、闊卵形、鋸齒闊卵形等多種形狀,這是一種植物適應異質(zhì)環(huán)境的葉片表型可塑性,是胡楊長期適應干旱地區(qū)生長環(huán)境的結(jié)果。本研究以胡楊葉表皮角質(zhì)膜為切入點,通過對內(nèi)蒙古額濟納地區(qū)八道橋(較為干旱)和七道橋(較為濕潤)胡楊異形葉片的形態(tài)特征、滲透調(diào)節(jié)、水分利用效率等生態(tài)特征、蠟質(zhì)晶體的形態(tài)結(jié)構(gòu)、胡楊長葉闊葉葉片表皮角質(zhì)膜中蠟質(zhì)、角質(zhì)的成分含量等生態(tài)及生理生化特征的比較,分析了胡楊異形葉對干旱環(huán)境的適應能力。同時,對不同水分條件下胡楊異形葉表皮蠟質(zhì)相關(guān)代謝基因的表達情況進行了對比分析。主要研究結(jié)果如下：(1)通過對比觀察干濕兩地的胡楊葉片形態(tài)我們發(fā)現(xiàn),胡楊幼樹均為長葉,成樹葉形多變,同種形態(tài)的葉片,在濕潤環(huán)境下,其表面積和體積均較大,葉片較薄；從葉形來講,長葉葉片單位體積的表面積更大,闊葉葉柄長度更大。從樹冠分布位置來看,長葉主要分布在樹冠中下層,闊葉主要分布在樹冠中上層。根據(jù)葉片表皮滲透性實驗,我們發(fā)現(xiàn)不論是在干旱環(huán)境還是相對濕潤環(huán)境,胡楊的長葉比闊葉表現(xiàn)出更高的葉綠素滲出速率和失水速率,說明長葉表皮比闊葉具有更高的滲透性,長葉的保水能力相對較弱,闊葉的保水能力相對較強。(2)胡楊不同形態(tài)葉片的表皮角質(zhì)膜微觀結(jié)構(gòu)存在較大差異,同時葉表皮蠟質(zhì)形態(tài)、晶體排列、分布以及葉片氣孔形態(tài)、數(shù)量與葉形和生長環(huán)境中的水分條件具有密切關(guān)系。與濕潤環(huán)境相比,干旱環(huán)境下的胡楊長葉具有更多的直立狀鱗片狀蠟質(zhì)晶體,而且排列更為密集、有序,而闊葉表皮蠟質(zhì)晶體融合成更多更厚的蠟被。在相同水分條件下,胡楊闊葉表皮蠟質(zhì)晶體融合成的蠟被比長葉面積更大,厚度更高。闊葉和干旱環(huán)境下的葉片氣孔有較明顯的下陷,而蠟被增厚造成氣孔位置相對下降是形成氣孔下陷的主要原因。相同水分條件下,胡楊闊葉表皮氣孔密度比長葉大；相同形態(tài)的葉片,干旱環(huán)境下葉片表皮的氣孔密度比濕潤環(huán)境大；相同類型的葉片遠軸面表皮氣孔密度比近軸面大。胡楊葉片表皮蠟質(zhì)晶體形態(tài)和氣孔形態(tài)、密度的差異是長期以來對干旱環(huán)境適應的結(jié)果。(3)通過對兩地胡楊異形葉表皮角質(zhì)膜中的蠟質(zhì)和角質(zhì)含量及成分進行對比分析,我們發(fā)現(xiàn)同一地點的胡楊闊葉比長葉表皮蠟質(zhì)和角質(zhì)含量高,干旱環(huán)境下生長的胡楊比濕潤環(huán)境下生長的胡楊葉片表皮蠟質(zhì)和角質(zhì)含量高,而蠟質(zhì)和角質(zhì)的組分比例也會隨著葉形和水分條件不同發(fā)生變化�？傮w來講,成樹闊葉的總蠟質(zhì)含量是長葉的1.3倍,而在干旱環(huán)境下,成樹闊葉的總蠟質(zhì)含量是成樹長葉的1.6倍。其中烷烴是蠟質(zhì)中差異最大的組分,在干旱環(huán)境下烷烴組分的含量遠高于其它組分,闊葉表皮蠟質(zhì)中烷烴含量達50%以上；在干旱環(huán)境下長葉的烷烴含量比濕潤環(huán)境下高出1.4倍,而干旱環(huán)境下闊葉的烷烴含量比在濕潤環(huán)境下高出1.2倍。我們在胡楊異形葉中共發(fā)現(xiàn)了7種角質(zhì)單體組分,其中最主要的成分是10,16-diOH C16:0酸,在所有檢驗樣本中占角質(zhì)單體總量一半以上,其它檢測到的組分包括：16-OHCi6：o酸C16：o酸C18:idioic酸,18-OHC18：1酸,C16:0dioic酸和18-OH C18:2酸。在濕潤環(huán)境下和干旱環(huán)境下,成樹闊葉比長葉分別多產(chǎn)生2.1倍和0.9倍角質(zhì)。長葉在干旱環(huán)境下比在濕潤環(huán)境下的角質(zhì)總含量多1.1倍,其中10,16-OH C16：0酸高出79.8%；闊葉在干旱環(huán)境下比在濕潤環(huán)境下的角質(zhì)總含量多31.5%,其中10,16-OHC16：0酸高出26.5%。另外,在兩種水分條件下闊葉角質(zhì)中C16：o酸和C18：1 dioic酸含量也高于長葉。(4)通過對不同水分條件下胡楊異形葉表皮蠟質(zhì)代謝基因的表達分析我們發(fā)現(xiàn),在干旱環(huán)境下胡楊葉中烷烴合成相關(guān)基因CER1以及ABC轉(zhuǎn)運蛋白和延長酶基因WBC11,WBC12比在濕潤環(huán)境下表達水平更高,從葉形上比較,闊葉CER1,CER4,KCS1和ATT1的表達量都高于長葉2倍以上(l0g2倍數(shù)),從環(huán)境水分條件來看,干旱環(huán)境下長葉CER1, WBC11、WBC12和WIN1的表達量都高于濕潤環(huán)境下長葉2倍以上(l0g2倍數(shù))。相對而言,干旱環(huán)境下闊葉CER1, CER26和MAH1的表達量高于濕潤環(huán)境下的闊葉2倍以上(log2倍數(shù))；干旱環(huán)境下,CER1, WBC11, WBC12和WIN1的表達量在長葉和闊葉中的表達量均較濕潤環(huán)境下有所提高。在兩種水分條件下,蠟質(zhì)代謝相關(guān)基因在闊葉中表達水平普遍較高,而且兩種葉形的葉片在干旱環(huán)境下蠟質(zhì)代謝相關(guān)基因表達水平普遍較高。在干旱環(huán)境下,胡楊葉片具有產(chǎn)生更多蠟質(zhì)的潛能。上述結(jié)果表明,胡楊葉片角質(zhì)膜在不同水分條件下、不同葉形葉片間的差異是胡楊對干旱環(huán)境適應性的重要體現(xiàn)。本研究為自然生境下的胡楊林及其生態(tài)環(huán)境的保護提供了有價值的數(shù)據(jù),同時為角質(zhì)膜和異形葉與植物的抗旱性關(guān)系研究提供了啟示。
[Abstract]:The study object of this article is Populus (Populus euphratica Oliv.), also called euphrates poplar or desert to poplar, Populus, is the formation of the ancient Mediterranean region since the third century of the oldest plants. The Populus mainly distributed in Central Asia, West Asia, North Africa to southern Europe (approximately at latitude 30 degrees to 50 degrees between Central Asia and China). Poplar is the northwest arid desert region is the only natural distribution of tree species, Populus China is the most widely distribution area in the world, the largest number of countries, in our country the main branch of Populus euphratica in the Tarim Basin in the Xinjiang Uygur Autonomous Region, Luntai county and Inner Mongolia Alashan Ejina area. The growth period is long, due to the impact of wind and drought a lot of poplar, strange shapes, strange, there is an old saying to describe it "live three thousand years dead, dead three thousand years, down three thousand years" immortal ". In China's ethnic culture, Hu Yang is regarded as "Shenmu" or "sacred tree", and as a symbol of death and resurrection is always closely connected with Phoenix and blood. Populus showed obvious characteristics of heteromorphic leaves during the growth of Populus euphratica. Leaf were slender, with the development of the upper leaves gradually become shorter and wider, the final form of on both sides with a jagged shape. Leaf leaved Populus euphratica in the same plant having lanceolate, elliptic, ovate, broadly ovate, serrated broad oval shape, this is the phenotypic plasticity of leaf of a plant to adapt to the heterogeneous environment, Hu Yang is long-term adaptation to arid area growth environment results in this study. Populus euphratica leaf epidermal cuticle as the breakthrough point, through to the Inner Mongolia Ejina area eight (drought) bridge and seven bridge (more humid) morphological characteristics of Populus euphratica leaf shaped osmotic adjustment, water use efficiency and ecological characteristics, The morphological structure of waxy crystal, waxy leaf leaf cuticle of Populus euphratica broadleaf film, ecological and physiological and biochemical characteristics of cutin composition, analyzes the adaptability of Populus euphratica heteromorphic leaves to drought environment. At the same time, the expression of Populus euphratica heteromorphic leaves of epicuticular wax related metabolic genes under different water conditions were analyzed the main results are as follows: (1) through the observation of Populus euphratica leaf morphology of wet and dry places we found that Populus euphratica were long leaf, a tree leaf blade is changeable, the same form, in the moist environment, the surface area and volume are larger, thinner leaves; from the leaf, leaf length per unit volume the surface area is larger, more broad leaf petiole length. From the view of crown distribution, mainly distributed in the lower leaf canopy, mainly distributed in the upper broadleaf canopy. According to leaf epidermal permeability. Check, we found both in the arid environment is still relatively moist environment, long leaf broad-leaved Populus showed chlorophyll exudation rate and water loss rate higher, long leaf epidermis has higher permeability than hardwood, leaf water retention capacity is relatively weak, broad leaf water retention capacity is relatively strong (2). There is a big difference between the cuticle microstructure morphology of Populus euphratica in different leaf, and leaf epicuticular wax morphology, crystal structure, distribution and leaf stomatal morphology, water conditions and the number of leaf and growth environment have close relationship. Compared with the moist environment, the arid environment of Populus euphratica leaf with crystal zhilizhuang flake more wax moreover, arranged more orderly, intensive, while broad-leaved epicuticular wax crystals merged into more thick wax. In the same water condition, Populus broad-leaved epicuticular wax crystals merged into the wax is longer than leaves The area is larger, the thickness is higher. Leaf stomata and broadleaf arid environment has obvious subsidence, and wax thickening caused by stomatal position relative decline is mainly due to the formation of sunken stomata. The same water condition, the stomatal density of Populus euphratica broadleaf than leaves; the same leaf morphology, stomatal density of leaf epidermis of drought under the moist environment than leaves; the same type of abaxial stomatal density than the adaxial surface. Leaf epidermis wax crystal morphology and pore morphology, density difference is to adapt to drought environment long-term results. (3) through the analysis and comparison of wax and cutin content and composition of two poplar shaped the leaf epidermal angle in plasma membrane, we found the same location than the leaf epicuticular wax of Populus euphratica broadleaf and cutin content is high, the leaf growth of Populus euphratica than humid environment in arid environment Epicuticular wax and cutin and wax and cutin content is high, the component ratio also with leaf shape and moisture change. Overall, a total wax content of broad-leaved tree is 1.3 times the length of leaves, while in the arid environment, the total wax content of broad-leaved tree is 1.6 times as Shuchang leaves. Which is the biggest difference in paraffin wax composition, content of component alkane in the arid environment is much higher than that of other components of epicuticular wax in broad-leaved alkane content reached more than 50%; the content of alkanes in the arid environment leaves is 1.4 times higher than that of the moist environment, and alkane content environment than broad-leaved drought in the moist environment is 1.2 times higher. We found 7 kinds of cutin monomer components in Populus euphratica heteromorphic leaves of the Communist Party of China, one of the most important components of 10,16-diOH C16:0 acid in all test samples accounted for more than half of the total cutin monomers, the other detected component package Including: 16-OHCi6:o acid C16:o acid C18:idioic acid, 18-OHC18:1 acid, C16:0dioic acid and 18-OH acid C18:2. In the moist environment and arid environment, tree leaves were more than broad-leaved produce 2.1 times and 0.9 times of cutin. Long leaf under drought conditions than the total cutin content in the moist environment of more than 1.1 times, including 10,16-OH C16:0 acid 79.8% higher; broad leaved in the arid environment than the total cutin content in the moist environment of more than 31.5%, which is higher than the 26.5%. 10,16-OHC16:0 acid, in addition, in the two water conditions in C16:o acid and C18:1 leaf cutin dioic acid content is higher than that of leaf. (4) the expression of Populus euphratica heteromorphic leaves of epicuticular wax metabolic genes under different water conditions analysis we found that in the arid environment of Populus euphratica leaf alkanes in synthesis related genes CER1 and ABC transporters and elongase gene WBC11, WBC12 expression level more than in the moist environment. High, from the leaf on the broad leaved CER1, CER4, expression of KCS1 and ATT1 were higher than that of leaves more than 2 times (l0g2 times), from the point of view of water condition, leaf CER1, arid environment WBC11, expression of WBC12 and WIN1 were higher than that in the moist environment leaves more than 2 times (l0g2 times number). In contrast, the arid environment of broad-leaved CER1, expression of CER26 and MAH1 was higher than that of the moist environment of hardwood is more than 2 times (log2 times); the arid environment, CER1, WBC11, WBC12 and WIN1 expression in leaf and broadleaf in volume were increased in the moist environment. The two water conditions, the expression level of higher wax metabolism related genes in the broad-leaved, and leaves of two leaf wax in the arid environment of metabolism related gene expression level is generally high. In the arid environment, has produced more leaf wax potential. The results showed that P. Leaf cuticle under different water conditions, the difference between the leaves of different leaf is an important manifestation of Populus to drought environment adaptability. This study provides valuable data for the protection of natural habitats of Populus euphratica forest and its ecological environment, at the same time as the cuticle and heteromorphic leaves provided the inspiration and research of drought resistance in plants.

【學位授予單位】：中央民族大學
【學位級別】：博士
【學位授予年份】：2016
【分類號】：S792.11

【參考文獻】

相關(guān)期刊論文 前10條

1 宋玉偉;楊偉林;;玉米幼苗葉表面蠟質(zhì)含量及成分對干旱脅迫的響應[J];玉米科學;2016年02期

2 宋玉;塔西甫拉提·特依拜;李崇博;侯艷軍;阿爾達克;張飛;;荒漠胡楊葉片含水率的高光譜反演[J];西北農(nóng)業(yè)學報;2016年02期

3 焦培培;井然;黃文娟;;胡楊不同發(fā)育期異形葉礦質(zhì)養(yǎng)分日內(nèi)動態(tài)特征研究[J];黑龍江生態(tài)工程職業(yè)學院學報;2016年01期

4 王海珍;韓路;徐雅麗;牛建龍;于軍;;灰胡楊葉片氣孔導度特征及數(shù)值模擬[J];林業(yè)科學;2016年01期

5 楊愛國;張玉玲;王利;董彬;紀鳳奎;;干旱脅迫下楊樹苗期葉片的阻抗譜參數(shù)和蒸騰速率的研究[J];遼寧林業(yè)科技;2016年01期

6 王海珍;徐雅麗;張翠麗;韓路;;干旱脅迫對胡楊和灰胡楊幼苗滲透調(diào)節(jié)物質(zhì)及抗氧化酶活性的影響[J];干旱區(qū)資源與環(huán)境;2015年12期

7 朱雙艷;齊軍倉;惠宏杉;廖樂;林立昊;華淼源;;大麥幼苗葉片表皮蠟質(zhì)組分及含量對外源ABA的響應及其對表皮透性的影響[J];西南農(nóng)業(yè)學報;2015年05期

8 張帆;白羽;王嘉歡;王美玲;孫瑜琳;王艷婷;景兵;王中華;肖恩時;;黑芝麻葉片表皮蠟質(zhì)成分及含量研究[J];西北農(nóng)業(yè)學報;2015年11期

9 白海霞;尚紅喜;石玉民;馬季;李景文;;極端干旱區(qū)胡楊葉與果實形態(tài)多樣性研究[J];林業(yè)調(diào)查規(guī)劃;2015年05期

10 趙春彥;司建華;馮起;魚騰飛;李培都;;風對極端干旱區(qū)胡楊蒸騰速率的影響[J];冰川凍土;2015年04期

相關(guān)博士學位論文 前10條

1 王文嬌;黃瓜蠟質(zhì)合成基因CsCER1和CsWAX2的克隆與功能驗證[D];中國農(nóng)業(yè)大學;2015年

2 馬濤;胡楊抗鹽的基因組遺傳基礎(chǔ)研究[D];蘭州大學;2014年

3 趙陽;黑河下游不同地下水位胡楊葉形態(tài)學生理學響應[D];蘭州大學;2012年

4 彭琦;與甘藍型油菜脂肪酸組分形成相關(guān)新基因的克隆[D];湖南農(nóng)業(yè)大學;2011年

5 薩如拉;額濟納天然胡楊林生態(tài)用水機理研究[D];內(nèi)蒙古農(nóng)業(yè)大學;2010年

6 王振宇;植物根源信號及葉片表皮層與耐旱性生理生態(tài)及分子生物學功能研究[D];蘭州大學;2009年

7 岳寧;胡楊異形葉生態(tài)適應的解剖及生理學研究[D];北京林業(yè)大學;2009年

8 周小云;水稻葉表皮蠟質(zhì)發(fā)育及蠟質(zhì)相關(guān)轉(zhuǎn)錄因子基因OsWTF1和OsWTF2的克隆與鑒定[D];湖南農(nóng)業(yè)大學;2007年

9 孫偉;蠟質(zhì)合成與轉(zhuǎn)運相關(guān)基因的功能研究[D];山東師范大學;2007年

10 劉普幸;額濟納旗胡楊徑向生長的水文氣候因子研究[D];蘭州大學;2007年

相關(guān)碩士學位論文 前10條

1 李加好;胡楊階段轉(zhuǎn)變過程枝、葉和花芽形態(tài)數(shù)量變化及生理特征研究[D];塔里木大學;2015年

2 馬持衡;擬南芥CER9基因調(diào)控種子油脂合成機理初步研究[D];中國科學院研究生院(武漢植物園);2015年

3 呂爽;胡楊幼苗地上地下生長特性對水土條件的響應[D];北京林業(yè)大學;2015年

4 鄭亞瓊;胡楊和灰葉胡楊克隆生長特征及與內(nèi)外資源的關(guān)系研究[D];塔里木大學;2014年

5 馮梅;胡楊葉形變化與個體發(fā)育階段的關(guān)系研究[D];塔里木大學;2014年

6 劉樹寶;基于穩(wěn)定同位素技術(shù)的荒漠河岸林胡楊水分來源研究[D];新疆農(nóng)業(yè)大學;2014年

7 涂文霞;塔里木河不同河段胡楊徑向生長與地表徑流的關(guān)系研究[D];新疆師范大學;2014年

8 徐靜;煙草葉片表皮蠟質(zhì)分析及組織培養(yǎng)環(huán)境對煙草葉片蠟質(zhì)層與保水性的影響[D];西北農(nóng)林科技大學;2014年

9 高麗娟;水分梯度下胡楊水分利用策略研究[D];新疆大學;2013年

10 王蕓;胡楊逆境光合生理及其甜菜堿醛脫氫酶基因動態(tài)表達研究[D];新疆大學;2013年

，

本文編號：1591022