干旱和鹽害對植物的影響和植物響應研究國內(nèi)外文獻綜述目錄TOC\o"1-3"\h\u148471.1干旱和鹽害現(xiàn)狀 195241.1.1干旱現(xiàn)狀 1274041.1.2鹽害現(xiàn)狀 1226721.2干旱和鹽害對植物的影響 247521.2.1干旱對植物的影響 2240291.2.2鹽害對植物的影響 2197721.3植物對干旱和鹽處理的響應 250501.3.1氣孔調(diào)節(jié) 2105301.3.2滲透調(diào)節(jié) 3312831.3.3清除活性氧 3121171.3.4根系調(diào)節(jié) 4169931.3.5激素調(diào)節(jié) 41.1干旱和鹽害現(xiàn)狀1.1.1干旱現(xiàn)狀干旱是指土壤水分散失多于吸收的氣候現(xiàn)象，高溫和降水偏少等ADDINEN.CITE<EndNote><Cite><Author>Basu</Author><Year>2016</Year><RecNum>72</RecNum><DisplayText><styleface="superscript">[13]</style></DisplayText><record><rec-number>72</rec-number><foreign-keys><keyapp="EN"db-id="9v2e5rpw1f5zzoewzxn5r5rys929ap0paztt"timestamp="1616581639">72</key><keyapp="ENWeb"db-id="">0</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Basu,S.</author><author>Ramegowda,V.</author><author>Kumar,A.</author><author>Pereira,A.</author></authors><secondary-authors><author>Basu,S.</author><author>Ramegowda,V.</author><author>Kumar,A.</author><author>Pereira,A.</author></secondary-authors></contributors><auth-address>Crop,Soil,andEnvironmentalSciences,UniversityofArkansas,Fayetteville,Arkansas,72701,USA.</auth-address><titles><title>Plantadaptationtodroughtstress</title><secondary-title>F1000Res</secondary-title></titles><periodical><full-title>F1000Res</full-title></periodical><volume>5</volume><edition>2016/07/22</edition><keywords><keyword>Adaptation</keyword><keyword>Droughttolerance</keyword><keyword>droughtresistance</keyword><keyword>grainyield</keyword><keyword>photosynthesis</keyword><keyword>rice</keyword></keywords><dates><year>2016</year></dates><isbn>2046-1402(Print) 2046-1402(Linking)</isbn><accession-num>27441087</accession-num><urls><related-urls><url>/pubmed/27441087</url></related-urls></urls><custom2>PMC4937719</custom2><electronic-resource-num>10.12688/f1000research.7678.1</electronic-resource-num></record></Cite></EndNote>[13]都能夠造成干旱現(xiàn)象。依據(jù)國家規(guī)定的《氣象干旱等級標準》，預測干旱的警示信號顏色設置為橙色和紅色。雖然現(xiàn)在科技高速發(fā)展，但是干旱問題依舊是我們難以克服的自然災害。隨著人口增多和商業(yè)經(jīng)濟的發(fā)展，用水量日益增多，進而加劇了干旱現(xiàn)象。因此，干旱難題應受到大多數(shù)人的關注和重視。全球的干旱面積占陸地的1/3，并且嚴重影響農(nóng)作物的生長。有數(shù)據(jù)表明，干旱現(xiàn)象引發(fā)的作物減產(chǎn)超過50%ADDINEN.CITE<EndNote><Cite><Author>張麗莉</Author><Year>2015</Year><RecNum>186</RecNum><DisplayText><styleface="superscript">[14]</style></DisplayText><record><rec-number>186</rec-number><foreign-keys><keyapp="EN"db-id="9v2e5rpw1f5zzoewzxn5r5rys929ap0paztt"timestamp="1616588881">186</key></foreign-keys><ref-typename="中文文獻">40</ref-type><contributors><authors><author>張麗莉</author><author>王慶祥</author><author>石瑛</author><author>祁雪</author></authors><secondary-authors><author>張麗莉</author><author>王慶祥</author><author>石瑛</author><author>祁雪</author></secondary-authors></contributors><auth-address>沈陽農(nóng)業(yè)大學農(nóng)學院;東北農(nóng)業(yè)大學農(nóng)學院;</auth-address><titles><title>干旱脅迫對馬鈴薯葉肉和莖部細胞超微結(jié)構的影響</title><secondary-title>沈陽農(nóng)業(yè)大學學報</secondary-title></titles><periodical><full-title>沈陽農(nóng)業(yè)大學學報</full-title></periodical><pages>91-95</pages><volume>46</volume><number>01</number><keywords><keyword>馬鈴薯</keyword><keyword>干旱脅迫</keyword><keyword>超微結(jié)構</keyword><keyword>葉肉</keyword><keyword>莖部</keyword></keywords><dates><year>2015</year></dates><isbn>1000-1700</isbn><call-num>21-1134/S</call-num><urls></urls><remote-database-provider>Cnki</remote-database-provider></record></Cite></EndNote>[14]，已經(jīng)變成一個不可忽視的問題ADDINEN.CITEADDINEN.CITE.DATA[15]。干旱可以造成植物形態(tài)、生理生化及體內(nèi)分子水平的變化，甚至破壞植物的光合系統(tǒng)和代謝系統(tǒng)，最后引起植物死亡。1.1.2鹽害現(xiàn)狀鹽堿化是由多種原因引起的土壤水分蒸發(fā)，形成土壤結(jié)塊的現(xiàn)象。依據(jù)聯(lián)合國糧農(nóng)組織統(tǒng)計數(shù)據(jù)，鹽堿化影響全球超過8億公頃土地面積以及約占2成有效使用土地面積。FAO組織推測21世紀50年代，由于世界人口增加，須加快糧食生產(chǎn)速度以至多產(chǎn)生70%的糧食才能滿足基本需要。中國鹽堿化土地面積幾乎有1億公頃，主要包括大西北、東三省和華北地區(qū)的缺水區(qū)域。我國能開發(fā)利用的鹽堿土地面積占中國現(xiàn)有田地的10%ADDINEN.CITE<EndNote><Cite><Author>王佳麗</Author><Year>2011</Year><RecNum>165</RecNum><DisplayText><styleface="superscript">[16]</style></DisplayText><record><rec-number>165</rec-number><foreign-keys><keyapp="EN"db-id="9v2e5rpw1f5zzoewzxn5r5rys929ap0paztt"timestamp="1616587299">165</key></foreign-keys><ref-typename="中文文獻">40</ref-type><contributors><authors><author>王佳麗</author><author>黃賢金</author><author>鐘太洋</author><author>陳志剛</author></authors><secondary-authors><author>王佳麗</author><author>黃賢金</author><author>鐘太洋</author><author>陳志剛</author></secondary-authors></contributors><auth-address>南京大學地理與海洋科學學院國土資源與旅游學系;安徽農(nóng)業(yè)大學經(jīng)管學院土地資源管理系;</auth-address><titles><title>鹽堿地可持續(xù)利用研究綜述</title><secondary-title>地理學報</secondary-title></titles><periodical><full-title>地理學報</full-title></periodical><pages>673-684</pages><volume>66</volume><number>05</number><keywords><keyword>鹽堿地</keyword><keyword>可持續(xù)利用</keyword><keyword>技術研發(fā)</keyword><keyword>農(nóng)戶行為</keyword><keyword>科學方法</keyword></keywords><dates><year>2011</year></dates><isbn>0375-5444</isbn><call-num>11-1856/P</call-num><urls></urls><remote-database-provider>Cnki</remote-database-provider></record></Cite></EndNote>[16]。因此將鹽堿土地轉(zhuǎn)變成農(nóng)田，將大大增加有效田地的使用面積。鹽堿土地中的鹽主要為氯化鈉和硫酸鈉等無機鹽，嚴重影響植物種子的發(fā)芽和幼苗生長，尤其是高鹽環(huán)境下的小麥和水稻幾乎都不能正常生長，嚴重影響我國糧食總量。因此，為了提高糧食總量，研究植物中的抗鹽基因為其提供了有效方法。1.2干旱和鹽害對植物的影響1.2.1干旱對植物的影響土壤含水量約束著植物的生長發(fā)育，不同等級的干旱都會引起植物滲透失水，而植物會通過自身調(diào)節(jié)途徑來抵抗干旱現(xiàn)象。植物能迅速地感知不利環(huán)境，進而調(diào)節(jié)脅迫相關基因的表達，最終使植物發(fā)生多種反應來適應干旱現(xiàn)象。植物的失水反應會打破離子平衡，引起氧化還原電位升高，產(chǎn)生活性氧，甚至破壞體內(nèi)的大分子物質(zhì)ADDINEN.CITE<EndNote><Cite><Author>Anjum</Author><Year>2011</Year><RecNum>52</RecNum><DisplayText><styleface="superscript">[17]</style></DisplayText><record><rec-number>52</rec-number><foreign-keys><keyapp="EN"db-id="9v2e5rpw1f5zzoewzxn5r5rys929ap0paztt"timestamp="1616581478">52</key><keyapp="ENWeb"db-id="">0</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Anjum,S.A.</author><author>Wang,L.</author><author>Farooq,M.</author><author>Khan,I.</author><author>Xue,L.</author></authors><secondary-authors><author>Anjum,S.A.</author><author>Wang,L.</author><author>Farooq,M.</author><author>Khan,I.</author><author>Xue,L.</author></secondary-authors></contributors><titles><title>MethylJasmonate-InducedAlterationinLipidPeroxidation,AntioxidativeDefenceSystemandYieldinSoybeanUnderDrought</title><secondary-title>JournalofAgronomyandCropScience</secondary-title></titles><periodical><full-title>JournalofAgronomyandCropScience</full-title></periodical><pages>296-301</pages><volume>197</volume><number>4</number><section>296</section><dates><year>2011</year></dates><isbn>09312250</isbn><urls></urls><electronic-resource-num>10.1111/j.1439-037X.2011.00468.x</electronic-resource-num></record></Cite></EndNote>[17]。如細胞膜脂質(zhì)發(fā)生氧化反應，產(chǎn)生丙二醛ADDINEN.CITEADDINEN.CITE.DATA[18,19]。干旱現(xiàn)象可破壞植物的細胞結(jié)構，導致植物非正常生長，如植株矮小、葉片枯萎等ADDINEN.CITE<EndNote><Cite><Author>Farooq</Author><Year>2009</Year><RecNum>227</RecNum><DisplayText><styleface="superscript">[20]</style></DisplayText><record><rec-number>227</rec-number><foreign-keys><keyapp="EN"db-id="9v2e5rpw1f5zzoewzxn5r5rys929ap0paztt"timestamp="1616632236">227</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Farooq,M.</author><author>Wahid,A.</author><author>Kobayashi,N.</author><author>Fujita,D.</author><author>Basra,Sma</author></authors></contributors><titles><title>Plantdroughtstress:effects,mechanismsandmanagement</title><secondary-title>AgronomyforSustainableDevelopment</secondary-title></titles><periodical><full-title>AgronomyforSustainableDevelopment</full-title></periodical><pages>185-212</pages><volume>29</volume><number>1</number><dates><year>2009</year></dates><urls></urls></record></Cite></EndNote>[20]。植物對干旱的響應可通過植物形態(tài)來判斷，如根組織和葉片組織。比如，抗旱性水稻可通過增加根部組織中的導管數(shù)量，提高水分使用效率。另外，根組織還可通過增加主根長度和側(cè)根數(shù)量來提高植物的吸收和運輸水分的能力ADDINEN.CITE<EndNote><Cite><Author>Henry</Author><Year>2011</Year><RecNum>113</RecNum><DisplayText><styleface="superscript">[21]</style></DisplayText><record><rec-number>113</rec-number><foreign-keys><keyapp="EN"db-id="9v2e5rpw1f5zzoewzxn5r5rys929ap0paztt"timestamp="1616581933">113</key><keyapp="ENWeb"db-id="">0</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Henry,Amelia</author><author>Gowda,VeereshR.P.</author><author>Torres,RolandoO.</author><author>McNally,KennethL.</author><author>Serraj,Rachid</author></authors><secondary-authors><author>Henry,Amelia</author><author>Gowda,VeereshR.P.</author><author>Torres,RolandoO.</author><author>McNally,KennethL.</author><author>Serraj,Rachid</author></secondary-authors></contributors><titles><title>Variationinrootsystemarchitectureanddroughtresponseinrice(Oryzasativa):PhenotypingoftheOryzaSNPpanelinrainfedlowlandfields</title><secondary-title>FieldCropsResearch</secondary-title></titles><periodical><full-title>FieldCropsResearch</full-title></periodical><pages>205-214</pages><volume>120</volume><number>2</number><section>205</section><dates><year>2011</year></dates><isbn>03784290</isbn><urls></urls><electronic-resource-num>10.1016/j.fcr.2010.10.003</electronic-resource-num></record></Cite></EndNote>[21]。此外，植物可以通過葉片向內(nèi)卷曲包藏氣孔和氣孔凹陷，來降低水分蒸騰速率ADDINEN.CITE<EndNote><Cite><Author>Gray</Author><Year>2004</Year><RecNum>74</RecNum><DisplayText><styleface="superscript">[22]</style></DisplayText><record><rec-number>74</rec-number><foreign-keys><keyapp="EN"db-id="9v2e5rpw1f5zzoewzxn5r5rys929ap0paztt"timestamp="1616581654">74</key><keyapp="ENWeb"db-id="">0</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Gray,J.E.</author><author>Hetherington,A.M.</author></authors><secondary-authors><author>Gray,J.E.</author><author>Hetherington,A.M.</author></secondary-authors></contributors><auth-address>DepartmentofMolecularBiologyandBiotechnology,UniversityofSheffield,SheffieldS102TN,UK.</auth-address><titles><title>Plantdevelopment:YODAthestomatalswitch</title><secondary-title>CurrBiol</secondary-title></titles><periodical><full-title>CurrBiol</full-title></periodical><pages>R488-90</pages><volume>14</volume><number>12</number><edition>2004/06/19</edition><keywords><keyword>*Adaptation,Physiological</keyword><keyword>ArabidopsisProteins/genetics/metabolism/*physiology</keyword><keyword>CellDifferentiation/physiology</keyword><keyword>*GeneExpressionRegulation,Plant</keyword><keyword>MAPKinaseKinaseKinases/genetics/metabolism/*physiology</keyword><keyword>Models,Biological</keyword><keyword>Mutation/genetics</keyword><keyword>PlantLeaves/genetics/*physiology</keyword><keyword>*PlantPhysiologicalPhenomena</keyword><keyword>Plants/*embryology</keyword><keyword>SerineEndopeptidases/metabolism</keyword></keywords><dates><year>2004</year><pub-dates><date>Jun22</date></pub-dates></dates><isbn>0960-9822(Print) 0960-9822(Linking)</isbn><accession-num>15203025</accession-num><urls><related-urls><url>/pubmed/15203025</url></related-urls></urls><electronic-resource-num>10.1016/j.cub.2004.06.019</electronic-resource-num></record></Cite></EndNote>[22]。1.2.2鹽害對植物的影響鹽害會抑制植物正常生長發(fā)育和降低生產(chǎn)效率ADDINEN.CITE<EndNote><Cite><Author>Hussain</Author><Year>2015</Year><RecNum>89</RecNum><DisplayText><styleface="superscript">[23]</style></DisplayText><record><rec-number>89</rec-number><foreign-keys><keyapp="EN"db-id="9v2e5rpw1f5zzoewzxn5r5rys929ap0paztt"timestamp="1616581762">89</key><keyapp="ENWeb"db-id="">0</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Hussain,M.Iftikhar</author><author>Lyra,Dionyssia-Angeliki</author><author>Farooq,Muhammad</author><author>Nikoloudakis,Nikolaos</author><author>Khalid,Nauman</author></authors><secondary-authors><author>Hussain,M.Iftikhar</author><author>Lyra,Dionyssia-Angeliki</author><author>Farooq,Muhammad</author><author>Nikoloudakis,Nikolaos</author><author>Khalid,Nauman</author></secondary-authors></contributors><titles><title>Saltanddroughtstressesinsafflower:areview</title><secondary-title>AgronomyforSustainableDevelopment</secondary-title></titles><periodical><full-title>AgronomyforSustainableDevelopment</full-title></periodical><volume>36</volume><number>1</number><dates><year>2015</year></dates><isbn>1774-0746 1773-0155</isbn><urls></urls><electronic-resource-num>10.1007/s13593-015-0344-8</electronic-resource-num></record></Cite></EndNote>[23]。鹽處理時，植物的滲透勢短期內(nèi)迅速升高，伴隨著過量氧化物質(zhì)和無機離子的積累對植物造成嚴重的損害ADDINEN.CITE<EndNote><Cite><Author>Munns</Author><Year>2008</Year><RecNum>50</RecNum><DisplayText><styleface="superscript">[24]</style></DisplayText><record><rec-number>50</rec-number><foreign-keys><keyapp="EN"db-id="9v2e5rpw1f5zzoewzxn5r5rys929ap0paztt"timestamp="1616581465">50</key><keyapp="ENWeb"db-id="">0</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Munns,R.</author><author>Tester,M.</author></authors><secondary-authors><author>Munns,R.</author><author>Tester,M.</author></secondary-authors></contributors><auth-address>CSIROPlantIndustry,Canberra,ACT,Australia.rana.munns@csiro.au</auth-address><titles><title>Mechanismsofsalinitytolerance</title><secondary-title>AnnuRevPlantBiol</secondary-title></titles><periodical><full-title>AnnuRevPlantBiol</full-title></periodical><pages>651-81</pages><volume>59</volume><edition>2008/05/01</edition><keywords><keyword>Hordeum/physiology</keyword><keyword>Oryza/physiology</keyword><keyword>OsmoticPressure</keyword><keyword>OxidativeStress</keyword><keyword>Photosynthesis</keyword><keyword>*PlantPhysiologicalPhenomena</keyword><keyword>PlantShoots/metabolism</keyword><keyword>Plants/genetics</keyword><keyword>SignalTransduction</keyword><keyword>Sodium/metabolism</keyword><keyword>SodiumChloride/metabolism</keyword><keyword>Thermodynamics</keyword></keywords><dates><year>2008</year></dates><isbn>1543-5008(Print) 1543-5008(Linking)</isbn><accession-num>18444910</accession-num><urls><related-urls><url>/pubmed/18444910</url></related-urls></urls><electronic-resource-num>10.1146/annurev.arplant.59.032607.092911</electronic-resource-num></record></Cite></EndNote>[24]，這種響應在高濃度鹽處理時尤其明顯。此外，鹽處理會通過降低土壤水勢來阻礙植物吸水，進而增加植物脫水的概率ADDINEN.CITEADDINEN.CITE.DATA[25-27]。此反應可由少量鹽分引起，并且發(fā)生速度較快ADDINEN.CITEADDINEN.CITE.DATA[28,29]。土地鹽堿化會大幅度地減少植物量，并且其問題日益突出ADDINEN.CITEADDINEN.CITE.DATA[30,31]。一些研究提到，在鹽處理時植物體內(nèi)的水分會從原生質(zhì)體流向質(zhì)外體，降低葉片細胞間隙中的鹽濃度，進而減少無機離子對細胞的傷害ADDINEN.CITEADDINEN.CITE.DATA[32]。對植物造成鹽害的主要為鈉離子和氯離子，會改變植物的滲透勢，并對植物造成離子毒害ADDINEN.CITE<EndNote><Cite><Author>Munns</Author><Year>2011</Year><RecNum>73</RecNum><DisplayText><styleface="superscript">[33]</style></DisplayText><record><rec-number>73</rec-number><foreign-keys><keyapp="EN"db-id="9v2e5rpw1f5zzoewzxn5r5rys929ap0paztt"timestamp="1616581644">73</key><keyapp="ENWeb"db-id="">0</key></foreign-keys><ref-typename="BookSection">5</ref-type><contributors><authors><author>Munns,Rana</author></authors><secondary-authors><author>Munns,Rana</author></secondary-authors></contributors><titles><title>PlantAdaptationstoSaltandWaterStress</title><secondary-title>PlantResponsestoDroughtandSalinityStress-DevelopmentsinaPost-GenomicEra</secondary-title><tertiary-title>AdvancesinBotanicalResearch</tertiary-title></titles><pages>1-32</pages><dates><year>2011</year></dates><isbn>9780123876928</isbn><urls></urls><electronic-resource-num>10.1016/b978-0-12-387692-8.00001-1</electronic-resource-num></record></Cite></EndNote>[33]。高濃度鹽處理，抑制了植物根通過滲透作用從土壤中吸水的反應。此外，這種情況還會導致氣孔關閉ADDINEN.CITE<EndNote><Cite><Author>Asrar</Author><Year>2017</Year><RecNum>87</RecNum><DisplayText><styleface="superscript">[34]</style></DisplayText><record><rec-number>87</rec-number><foreign-keys><keyapp="EN"db-id="9v2e5rpw1f5zzoewzxn5r5rys929ap0paztt"timestamp="1616581748">87</key><keyapp="ENWeb"db-id="">0</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Asrar,Hina</author><author>Hussain,Tabassum</author><author>Hadi,SyedaMidhatSabahat</author><author>Gul,Bilquees</author><author>Nielsen,BrentL.</author><author>Khan,M.Ajmal</author></authors><secondary-authors><author>Asrar,Hina</author><author>Hussain,Tabassum</author><author>Hadi,SyedaMidhatSabahat</author><author>Gul,Bilquees</author><author>Nielsen,BrentL.</author><author>Khan,M.Ajmal</author></secondary-authors></contributors><titles><title>SalinityinducedchangesinlightharvestingandcarbonassimilatingcomplexesofDesmostachyabipinnata(L.)Staph</title><secondary-title>EnvironmentalandExperimentalBotany</secondary-title></titles><periodical><full-title>EnvironmentalandExperimentalBotany</full-title></periodical><pages>86-95</pages><volume>135</volume><section>86</section><dates><year>2017</year></dates><isbn>00988472</isbn><urls></urls><electronic-resource-num>10.1016/j.envexpbot.2016.12.008</electronic-resource-num></record></Cite></EndNote>[34]，影響光合作用，從而產(chǎn)生過量的活性氧物質(zhì)ADDINEN.CITE<EndNote><Cite><Author>Pinheiro</Author><Year>2011</Year><RecNum>70</RecNum><DisplayText><styleface="superscript">[35]</style></DisplayText><record><rec-number>70</rec-number><foreign-keys><keyapp="EN"db-id="9v2e5rpw1f5zzoewzxn5r5rys929ap0paztt"timestamp="1616581624">70</key><keyapp="ENWeb"db-id="">0</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Pinheiro,C.</author><author>Chaves,M.M.</author></authors><secondary-authors><author>Pinheiro,C.</author><author>Chaves,M.M.</author></secondary-authors></contributors><auth-address>InstitutodeTecnologiaQuimicaeBiologica,UniversidadeNovadeLisboa,AvdaRepublica-EAN,2780-157Oeiras,Portugal.</auth-address><titles><title>Photosynthesisanddrought:canwemakemetabolicconnectionsfromavailabledata?</title><secondary-title>JExpBot</secondary-title></titles><periodical><full-title>JExpBot</full-title></periodical><pages>869-82</pages><volume>62</volume><number>3</number><edition>2010/12/22</edition><keywords><keyword>Arabidopsis/genetics/*metabolism</keyword><keyword>Droughts</keyword><keyword>GeneExpressionRegulation,Plant</keyword><keyword>Hordeum/genetics/*metabolism</keyword><keyword>*Photosynthesis</keyword><keyword>PlantGrowthRegulators/metabolism</keyword><keyword>Water/*metabolism</keyword></keywords><dates><year>2011</year><pub-dates><date>Jan</date></pub-dates></dates><isbn>1460-2431(Electronic) 0022-0957(Linking)</isbn><accession-num>21172816</accession-num><urls><related-urls><url>/pubmed/21172816</url></related-urls></urls><electronic-resource-num>10.1093/jxb/erq340</electronic-resource-num></record></Cite></EndNote>[35]。已在多種植物中報道了鹽處理會嚴重影響光合系統(tǒng)的正常工作，比如高粱ADDINEN.CITEADDINEN.CITE.DATA[36]和向日葵ADDINEN.CITE<EndNote><Cite><Author>Taher</Author><Year>2018</Year><RecNum>53</RecNum><DisplayText><styleface="superscript">[37]</style></DisplayText><record><rec-number>53</rec-number><foreign-keys><keyapp="EN"db-id="9v2e5rpw1f5zzoewzxn5r5rys929ap0paztt"timestamp="1616581485">53</key><keyapp="ENWeb"db-id="">0</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Taher,Mehdi</author><author>Beyaz,Ramazan</author><author>Javani,Marieh</author><author>Gürsoy,Mehtap</author><author>Yildiz,Mustafa</author></authors><secondary-authors><author>Taher,Mehdi</author><author>Beyaz,Ramazan</author><author>Javani,Marieh</author><author>Gürsoy,Mehtap</author><author>Yildiz,Mustafa</author></secondary-authors></contributors><titles><title>Morphologicalandbiochemicalchangesinresponsetosalinityinsunflower(HelianthusannusL.)cultivars</title><secondary-title>ItalianJournalofAgronomy</secondary-title></titles><periodical><full-title>ItalianJournalofAgronomy</full-title></periodical><pages>141-147</pages><section>141</section><dates><year>2018</year></dates><isbn>2039-6805 1125-4718</isbn><urls></urls><electronic-resource-num>10.4081/ija.2018.1096</electronic-resource-num></record></Cite></EndNote>[37]。除了破壞光合系統(tǒng)，鹽處理也會造成細胞膜的損傷，產(chǎn)生許多細胞膜氧化物質(zhì)丙二醛，從而激活體內(nèi)防御系統(tǒng)ADDINEN.CITE<EndNote><Cite><Author>田曉艷</Author><Year>2009</Year><RecNum>164</RecNum><DisplayText><styleface="superscript">[38]</style></DisplayText><record><rec-number>164</rec-number><foreign-keys><keyapp="EN"db-id="9v2e5rpw1f5zzoewzxn5r5rys929ap0paztt"timestamp="1616587240">164</key></foreign-keys><ref-typename="中文文獻">40</ref-type><contributors><authors><author>田曉艷</author><author>劉延吉</author><author>張蕾</author><author>張弘</author><author>劉沛含</author></authors><secondary-authors><author>田曉艷</author><author>劉延吉</author><author>張蕾</author><author>張弘</author><author>劉沛含</author></secondary-authors></contributors><auth-address>遼寧石油化工大學環(huán)境與生物工程學院;沈陽農(nóng)業(yè)大學生物科學技術學院;</auth-address><titles><title>鹽脅迫對景天三七保護酶系統(tǒng)、MDA、Pro及可溶性糖的影響</title><secondary-title>草原與草坪</secondary-title></titles><periodical><full-title>草原與草坪</full-title></periodical><pages>11-14</pages><number>06</number><keywords><keyword>景天三七</keyword><keyword>鹽脅迫</keyword><keyword>MDA</keyword><keyword>保護酶系統(tǒng)</keyword><keyword>Pro</keyword><keyword>可溶性糖</keyword></keywords><dates><year>2009</year></dates><isbn>1009-5500</isbn><call-num>62-1156/S</call-num><urls></urls><remote-database-provider>Cnki</remote-database-provider></record></Cite></EndNote>[38]。植物中的抗氧化酶用于清除體內(nèi)的ROS，保護細胞ADDINEN.CITE<EndNote><Cite><Author>Tiffin</Author><Year>1959</Year><RecNum>223</RecNum><DisplayText><styleface="superscript">[39]</style></DisplayText><record><rec-number>223</rec-number><foreign-keys><keyapp="EN"db-id="9v2e5rpw1f5zzoewzxn5r5rys929ap0paztt"timestamp="1616591021">223</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Tiffin,L.O.</author><author>Brown,J.C.</author></authors><secondary-authors><author>Tiffin,L.O.</author><author>Brown,J.C.</author></secondary-authors></contributors><titles><title>AbsorptionofIronfromIronChelatebySunflowerRoots</title><secondary-title>Science</secondary-title><alt-title>Science(NewYork,N.Y.)</alt-title></titles><periodical><full-title>Science</full-title></periodical><pages>274-5</pages><volume>130</volume><number>3370</number><edition>1959/07/31</edition><dates><year>1959</year><pub-dates><date>Jul31</date></pub-dates></dates><isbn>0036-8075(Print) 0036-8075</isbn><accession-num>17813680</accession-num><urls></urls><electronic-resource-num>10.1126/science.130.3370.274-a</electronic-resource-num><remote-database-provider>NLM</remote-database-provider><language>eng</language></record></Cite></EndNote>[39]。1.3植物對干旱和鹽處理的響應1.3.1氣孔調(diào)節(jié)氣孔為植物所獨有的器官，在控制植物呼吸和調(diào)控蒸騰速率中起作用，它由腎形或啞鈴形的保衛(wèi)細胞組成ADDINEN.CITEADDINEN.CITE.DATA[40]。兩種形態(tài)的保衛(wèi)細胞，都是利用細胞壁的厚度不均來控制氣孔開閉。當保衛(wèi)細胞吸水時，保衛(wèi)細胞壁薄部位的伸縮性能好，向外彎曲或兩邊膨脹，氣孔打開。相反情況下，氣孔關閉。因此，植物保衛(wèi)細胞參與調(diào)節(jié)蒸騰速率的過程ADDINEN.CITEADDINEN.CITE.DATA[41]。所以，當植物保衛(wèi)細胞吸水而水勢升高時，植物氣孔打開，促進蒸騰作用，維持植物體的水分平衡。相反，當保衛(wèi)細胞失水而水勢降低時，氣孔關閉，減少植物體內(nèi)水分進一步散失ADDINEN.CITE<EndNote><Cite><Author>Lee</Author><Year>2012</Year><RecNum>11</RecNum><DisplayText><styleface="superscript">[42]</style></DisplayText><record><rec-number>11</rec-number><foreign-keys><keyapp="EN"db-id="9v2e5rpw1f5zzoewzxn5r5rys929ap0paztt"timestamp="1616581246">11</key><keyapp="ENWeb"db-id="">0</key></foreign-keys><ref-typename="JournalArticle">17</ref-type><contributors><authors><author>Lee,S.C.</author><author>Luan,S.</author></authors><secondary-authors><author>Lee,S.C.</author><author>Luan,S.</author></secondary-authors></contributors><auth-address>SchoolofBiologicalSciences(BK21program),Chung-AngUniversity,Seoul,Korea.</auth-address><titles><title>ABAsignaltransductionatthecrossroadofbioticandabioticstressresponses</title><secondary-title>PlantCellEnviron</secondary-title></titles><periodical><full-title>PlantCellEnviron</full-title></periodical><pages>53-60</pages><volume>35</volume><number>1</number><edition>2011/09/20</edition><keywords><keyword>AbscisicAcid/*metabolism</keyword><keyword>Droughts</keyword><keyword>GeneExpressionRegulation,Plant/physiology</keyword><keyword>PlantGrowthRegulators/*metabolism</keyword><keyword>*PlantPhysiologicalPhenomena</keyword><keyword>PlantProteins/genetics/metabolism</keyword><keyword>PlantStomata</keyword><keyword>Plants/genetics/metabolism</keyword><keyword>SignalTransduction/*physiology</keyword><keyword>Stress,Physiological/*physiology</keyword></keywords><dates><year>2012</year><pub-dates><date>Jan</date></pub-dates></dates><isbn>1365-3040(Electronic) 0140-7791(Linking)</isbn><accession-num>21923759</accession-num><urls><related-urls><url>/pubmed/21923759</url></related-urls></urls><electronic-resource-num>10.1111/j.1365-3040.2011.02426.x</electronic-resource-num></record></Cite></EndNote>[42]。而氣孔密度/導度與植物對水脅迫信號的響應有關ADDINEN.CITEADDINEN.CITE.DATA[43]。當植物處于缺水狀態(tài)下，植物會迅速積累脫落酸，進而誘導保衛(wèi)細胞內(nèi)涌入大量的鈣離子。此時因保衛(wèi)細胞的滲透勢升高而失水，導致氣孔導度減小，提高植物的抗旱性ADDINEN.CITEADDINEN.CITE.DATA[44,45]。1.3.2滲透調(diào)節(jié)通過滲透調(diào)節(jié)來應對干旱和鹽脅迫已經(jīng)得到大家廣泛的認同，并且成為研究植物抗旱性應用最熱門的機制之一ADDINEN.CITE<EndNote><Cite><Author>黎裕</Author><Year>1994</Year><RecNum>182</RecNum><DisplayText><styleface="superscript">[46]</style></DisplayText><record><rec-number>182</rec-number><foreign-keys><keyapp="EN"db-id="9v2e5rpw1f5zzoewzxn5r5rys929ap0paztt"timestamp="1616588324">182</key></foreign-keys><ref-typename="中文文獻">40</ref-type><contributors><authors><author>黎裕</author></authors><secondary-authors><author>黎裕</author></secondary-authors></contributors><auth-address>中國農(nóng)業(yè)科學院作物品種資源研究所!北京,100081</auth-address><titles><title>植物的滲透調(diào)節(jié)與其它生理過程的關系及其在作物改良中的應用</title><secondary-title>植物生理學通訊</secondary-title></titles><periodical><full-title>植物生理學通訊</full-title></periodical><pages>377-383</pages><number>05</number><keywords><keyword>滲透調(diào)節(jié)</keyword><keyword>氣孔導度</keyword><keyword>水分虧缺</keyword><keyword>氣孔調(diào)節(jié)</keyword><keyword>籽粒產(chǎn)量</keyword><keyword>光合活性</keyword><keyword>水分脅迫</keyword><keyword>作物改良</keyword><keyword>生理過程</keyword></keywords><dates><year>1994</year></dates><isbn>0412-0922</isbn><call-num>31-1350/Q</call-num><urls></urls><remote-database-provider>Cnki</remote-database-provider></record></Cite></EndNote>[46]。滲透調(diào)節(jié)一般發(fā)生在水分脅迫過程中，誘導其發(fā)生是一個緩慢的過程。植物滲透調(diào)節(jié)的物質(zhì)包括簡單離子和有機物質(zhì)。簡單離子包括鉀離子、氯離子和鈣離子等，此外鈣離子還可在信號傳遞中起作用。另一類是有機化合物：脯氨酸、可溶性糖和甜菜堿等ADDINEN.CITE<EndNote><Cite><Author>朱維琴</Author><Year>2003</Year><RecNum>185</RecNum><DisplayText><styleface="superscript">[47]</style></DisplayText><record><rec-number>185</rec-number><foreign-keys><keyapp="EN"db-id="9v2e5rpw1f5zzoewzxn5r5rys929ap0paztt"timestamp="1616588546">185</key></foreign-keys><ref-typename="中文文獻">40</ref-type><contributors><authors><author>朱維琴</author><author>吳良歡</author><author>陶勤南</author></authors><secondary-authors><author>朱維琴</author><author>吳良歡</author><author>陶勤南</author></secondary-authors></contributors><auth-address>浙江大學環(huán)境與資源學院,浙江大學環(huán)境與資源學院,浙江大學環(huán)境與資源學院浙江杭州310029,浙江杭州310029,浙江杭州310029</auth-address><titles><title>干旱逆境下不同品種水稻葉片有機滲透調(diào)節(jié)物質(zhì)變化研究</title><secondary-title>土壤通報</secondary-title></titles><periodical><full-title>土壤通報</full-title></periodical><pages>25-28</pages><number>01</number><keywords><keyword>干旱脅迫</keyword><keyword>聚乙二醇(PEG)</keyword><keyword>滲調(diào)物質(zhì)</keyword><keyword>水稻</keyword></keywords><dates><year>2003</year></dat