前期水淹對(duì)水杉樹(shù)苗響應(yīng)干旱脅迫的影響

白林利，韓文嬌，李昌曉

(西南大學(xué) 生命科學(xué)學(xué)院 三峽庫(kù)區(qū)生態(tài)環(huán)境教育部重點(diǎn)實(shí)驗(yàn)室，重慶 400715)

前期水淹對(duì)水杉樹(shù)苗響應(yīng)干旱脅迫的影響

白林利，韓文嬌，李昌曉

(西南大學(xué) 生命科學(xué)學(xué)院 三峽庫(kù)區(qū)生態(tài)環(huán)境教育部重點(diǎn)實(shí)驗(yàn)室，重慶 400715)

水分脅迫；水杉；生理生化；消落帶；三峽庫(kù)區(qū)

三峽工程建成蓄水后，由于其“冬蓄夏排”的反季節(jié)水位調(diào)度管理方式，使庫(kù)岸消落帶處于被淹沒(méi)與出水暴露的周期性更替變化環(huán)境之中[1-2]。消落帶水分條件的周期性變化使原有植被消失殆盡。因此，修復(fù)與重建消落帶植被要求所栽種的植物不僅具有適應(yīng)水分飽和與過(guò)剩的能力，還要具有適應(yīng)水分虧缺與干旱的能力。水杉(Metasequoiaglyptostroboides)是三峽庫(kù)區(qū)庫(kù)岸消落帶典型的鄉(xiāng)土樹(shù)種，別稱(chēng)水沙，是中國(guó)特有孑遺珍貴樹(shù)種。水杉根系發(fā)達(dá)，耐寒且耐受多種水分逆境的能力較強(qiáng)，是亞熱帶地區(qū)平原綠化的優(yōu)良樹(shù)種，為園林植物造景中的精品[3]。目前，關(guān)于水杉的研究主要集中在基因結(jié)構(gòu)[4-6]、無(wú)性繁殖[7]、苗木營(yíng)養(yǎng)特性[3]、葉片化學(xué)成分[8-9]和光合作用[10-11]等方面。而關(guān)于水杉如何應(yīng)對(duì)水分脅迫，尤其是自然生長(zhǎng)或人工栽植于三峽庫(kù)岸的水杉，其所經(jīng)歷的水淹脅迫是否會(huì)對(duì)隨后的干旱脅迫耐受性產(chǎn)生影響，至今未見(jiàn)相關(guān)報(bào)道。

本研究以水杉樹(shù)苗為試驗(yàn)材料，通過(guò)模擬三峽庫(kù)區(qū)消落帶土壤水分的變化格局，研究水杉樹(shù)苗各項(xiàng)生理生化指標(biāo)在水分脅迫處理下的變化規(guī)律，探討其對(duì)水分脅迫的生理生化響應(yīng)及耐受程度，旨在揭示水分脅迫對(duì)水杉樹(shù)苗生理生化特性的傷害機(jī)制及植株自身的抗逆機(jī)制，為三峽庫(kù)區(qū)庫(kù)岸消落帶的植被恢復(fù)與重建提供候選植物材料。

1 材料與方法

1.1 材料及處理

考慮到三峽庫(kù)區(qū)庫(kù)岸防護(hù)林體系建設(shè)多采用2年生苗木，因此，本試驗(yàn)以2年生水杉(Metasequoiaglyptostroboides)樹(shù)苗為材料。2012-11-20將生長(zhǎng)基本一致的72株樹(shù)苗帶土盆栽，土壤性質(zhì)為：pH 8.26±0.04,有機(jī)質(zhì)(11.62±0.56) g/kg，全氮(1.11±0.04) g/kg，全磷(1.11±0.10) g/kg，全鉀(53.61±5.24) g/kg，堿解氮(76.70±3.78) mg/kg，有效磷(0.85±0.16) mg/kg，速效鉀(161.02±4.08) mg/kg，每盆1株(盆中央內(nèi)徑20 cm，盆高17 cm)，置于西南大學(xué)三峽庫(kù)區(qū)生態(tài)環(huán)境教育部重點(diǎn)實(shí)驗(yàn)室實(shí)驗(yàn)基地大棚(海拔249 m)內(nèi)進(jìn)行相同的光照和水分管理。2013-01-18開(kāi)始試驗(yàn)，此時(shí)水杉苗高為(97.05±1.53) cm。

1.2 試驗(yàn)設(shè)計(jì)

結(jié)合三峽庫(kù)區(qū)消落帶水位變化的實(shí)際情況，將試驗(yàn)分為3個(gè)階段。第1階段為淹水處理期，共設(shè)3個(gè)處理組，分別為對(duì)照組(Control,C)、半淹組(Half-submersion,HS)和全淹組(Full-submersion,FS)，每組試驗(yàn)苗木24株，共72株。其中，C組為常規(guī)供水，保持土壤含水量為田間持水量的75%～80%[12]；HS組苗盆放入水池中，池水保持淹沒(méi)至植物中段；FS組苗盆也放入水池中，但池水保持沒(méi)過(guò)植物頂端20 cm。75 d后，進(jìn)行第2階段試驗(yàn)。

第2階段為干旱處理期，將第1階段每組的24株苗木再隨機(jī)均分為2組，其中1組進(jìn)行輕度干旱脅迫處理，另1組則繼續(xù)保持第1階段的水分處理。故第2階段共有對(duì)照-干旱組(Control followed by drought,CD)、半淹-干旱組(Half-submersion followed by drought,HSD)、全淹-干旱組(Full-submersion followed by drought,FSD)及對(duì)照組(C)、半淹組(HS)、全淹組(FS)6個(gè)處理，每組苗木12株，共72株。輕度干旱處理組土壤含水量為田間持水量的47%～50%[13]。為監(jiān)測(cè)輕度干旱處理的效果，于清晨06:00-07:00測(cè)定水杉樹(shù)苗上部成熟葉的葉水勢(shì)(用美國(guó)Wescor公司生產(chǎn)的露點(diǎn)水勢(shì)儀Psypro測(cè)定)，以小于-0.5 MPa為標(biāo)準(zhǔn)(圖1)。60 d后，每組各取6株水杉葉片用于生理指標(biāo)的測(cè)定，其余6株用于后續(xù)試驗(yàn)(第3階段試驗(yàn))。

圖1 輕度干旱處理階段水杉的清晨葉水勢(shì)
Fig.1 Changes of predawn leaf water potential ofM.glyptostroboidesduring drought stress

第3階段為恢復(fù)處理期，將HS與FS處理組苗盆從水池中取出，所有處理組均參照對(duì)照組(C)進(jìn)行常規(guī)供水，時(shí)間為21 d。第3階段試驗(yàn)結(jié)束后，各組分別取6株水杉葉片用于生理指標(biāo)的測(cè)定。

1.3 生理指標(biāo)的測(cè)定

1.4 數(shù)據(jù)統(tǒng)計(jì)分析

利用SPSS 16.0軟件進(jìn)行單因素方差分析(One-way ANOVA)，揭示水分處理對(duì)水杉生理生化特性的影響；并用Tukey檢驗(yàn)法檢驗(yàn)每個(gè)指標(biāo)在同一階段不同處理間(P=0.05)的差異顯著性。

2 結(jié)果與分析

2.1 前期水淹對(duì)水杉響應(yīng)干旱脅迫的影響

2.1.1 滲透調(diào)節(jié)物質(zhì) 水淹及干旱脅迫對(duì)水杉樹(shù)苗滲透調(diào)節(jié)物質(zhì)的影響如圖2所示。

圖2顯示，與C組相比，HS、FS組水杉樹(shù)苗脯氨酸含量顯著升高，而CD、HSD、FSD組與C組之間均無(wú)顯著性差異；CD、HSD、FSD組可溶性糖含量顯著高于C組，但與HS組之間均無(wú)顯著性差異，而FS組顯著低于C組；與C組相比，HS、FS、CD組的可溶性蛋白含量分別增加了63%，88%，48%(P<0.05)，而HSD、FSD組與C組之間均無(wú)顯著性差異。就滲透調(diào)節(jié)物質(zhì)而言，CD、HSD、FSD組的脯氨酸、可溶性糖含量之間均無(wú)顯著性差異，說(shuō)明前期的水淹并未增加水杉樹(shù)苗后期對(duì)干旱脅迫的敏感性。

圖3顯示，HS組水杉樹(shù)苗的MDA、超氧根離子含量顯著高于其他處理組，與C組相比分別升高了96%，24%，這與FS組顯著低于其他處理組形成鮮明對(duì)照；不同的是，與C組相比，CD、HSD、FSD組的MDA含量均顯著降低，且CD、HSD、FSD組之間均無(wú)顯著性差異；而CD、HSD、FSD組的超氧根離子與C組均無(wú)顯著差異，說(shuō)明前期的水淹并未對(duì)水杉樹(shù)苗后期在干旱脅迫下的MDA、超氧根離子含量變化造成影響。

與C組相比，F(xiàn)S、CD、HSD組的SOD、POD活性均顯著升高；就SOD而言，HS組與C組之間無(wú)顯著性差異，F(xiàn)SD組顯著低于C組；HS 組的POD活性顯著低于其他各組。與C組相比，HS、FS、HSD、FSD組的ASP活性均顯著降低，而CD與C組之間無(wú)顯著性差異。HS、FS、CD、HSD、C組的CAT活性均無(wú)顯著性差異，而FSD組顯著高于其他各組。就抗氧化酶活性而言，CD、HSD組的SOD、POD、CAT活性之間均無(wú)顯著性差異，說(shuō)明對(duì)照干旱組與半淹干旱組前期的水淹并未影響水杉樹(shù)苗后期抗氧化酶對(duì)干旱脅迫的響應(yīng)。

2.2 干旱脅迫后水杉對(duì)復(fù)水的生理響應(yīng)

2.2.1 滲透調(diào)節(jié)物質(zhì) 由圖4可見(jiàn)，在恢復(fù)正常供水之后，各個(gè)處理組水杉樹(shù)苗的脯氨酸含量之間均無(wú)顯著性差異。HS組的可溶性糖含量在所有處理中居于最高，CD、FSD組與C組及HS組與HSD組之間可溶性糖含量無(wú)顯著性差異。HS組的可溶性蛋白含量在所有處理中居于最高，HS、HSD組與C組及CD組與FSD組之間可溶性蛋白含量均無(wú)顯著性差異。

3 討 論

3.1 水杉對(duì)水淹脅迫的響應(yīng)

滲透調(diào)節(jié)是植物適應(yīng)逆境的重要生理機(jī)制[18]。脯氨酸作為一種重要的非酶抗氧化物質(zhì)，在穩(wěn)定亞細(xì)胞結(jié)構(gòu)、清除活性氧自由基中具有重要作用[19]。本研究中，水淹脅迫下，HS、FS組水杉樹(shù)苗游離脯氨酸含量顯著增加，說(shuō)明水杉樹(shù)苗通過(guò)產(chǎn)生脯氨酸來(lái)適應(yīng)水分脅迫[20]，保護(hù)酶活性。同時(shí)，HS、FS組可溶性蛋白含量均顯著高于C組，說(shuō)明水淹期間水杉樹(shù)苗能夠維持正常的氮代謝過(guò)程，這也可能是水杉樹(shù)苗對(duì)水淹逆境的一種內(nèi)在生理適應(yīng)機(jī)制[21]。在本研究中，F(xiàn)S組水杉樹(shù)苗可溶性糖含量顯著降低，說(shuō)明全淹脅迫干擾了水杉樹(shù)苗正常的糖代謝，抑制了糖的合成并誘導(dǎo)糖的降解，從而使植株體內(nèi)的可溶性糖含量降低[22]；也可能是由于水杉樹(shù)苗從有氧到無(wú)氧轉(zhuǎn)換的過(guò)程中，需要消耗大量的糖來(lái)維持發(fā)酵代謝活動(dòng)的完成[23]。脯氨酸、可溶性糖與可溶性蛋白綜合作用，調(diào)節(jié)水杉樹(shù)苗的滲透勢(shì)，從而保護(hù)細(xì)胞膜免受損害[24]。

3.2 水杉對(duì)水淹后干旱脅迫的響應(yīng)

在三峽庫(kù)區(qū)水文變動(dòng)條件下，庫(kù)區(qū)消落帶植被在冬季會(huì)遭受水淹脅迫；而在夏季水位下降后，由于氣溫和降雨的關(guān)系又可能面臨短暫的輕度干旱脅迫。干旱是限制植物生長(zhǎng)的重要非生物因子[33]，了解鄉(xiāng)土樹(shù)種對(duì)干旱脅迫的生理響應(yīng)有益于植被恢復(fù)工作的進(jìn)行[34]。

滲透調(diào)節(jié)是植物對(duì)干旱的關(guān)鍵響應(yīng)之一[35]。在干旱脅迫條件下，游離脯氨酸的積累對(duì)植物的適應(yīng)能力起到關(guān)鍵作用，其是有效的滲透調(diào)節(jié)物質(zhì)之一[36]。本研究中，CD、HSD、FSD組水杉樹(shù)苗脯氨酸含量與C組無(wú)顯著性差異，這與Gao等[37]對(duì)高山松的研究結(jié)果相似，說(shuō)明水杉樹(shù)苗對(duì)干旱脅迫的耐受有一定的局限性[38]。而CD、HSD、FSD組可溶性糖含量顯著高于C組，說(shuō)明在干旱脅迫下，水杉樹(shù)苗通過(guò)升高可溶性糖含量來(lái)調(diào)節(jié)滲透勢(shì)，維持膨壓，從而使體內(nèi)各種與膨壓有關(guān)的生理過(guò)程可以正常進(jìn)行[39]。同時(shí)，CD、HSD、FSD組可溶性蛋白含量高于C或與C組相當(dāng)，說(shuō)明干旱并未干擾水杉樹(shù)苗的正常氮代謝[40]。從脯氨酸含量相對(duì)較低且在干旱脅迫后并未明顯增加可知，可溶性糖是水杉樹(shù)苗應(yīng)對(duì)水分脅迫的主導(dǎo)性滲透調(diào)節(jié)物質(zhì)，其可以降低水勢(shì)，提高植物的吸水與保水能力[41]。

本研究發(fā)現(xiàn)，雖然CD與HSD組水杉樹(shù)苗的SOD、POD活性均顯著高于C組，但CD與HSD組的SOD、POD活性無(wú)顯著差異；同樣，干旱脅迫提高了CD與HSD組的脯氨酸和可溶性糖含量，但CD與HSD組之間無(wú)顯著差異，說(shuō)明前期水淹并未影響水杉樹(shù)苗后期對(duì)干旱脅迫的生理生化響應(yīng)。

3.3 水杉對(duì)干旱脅迫后復(fù)水的響應(yīng)

一個(gè)物種對(duì)脅迫的忍耐性取決于脅迫時(shí)間、脅迫強(qiáng)度以及去除脅迫后的恢復(fù)能力[48]，恢復(fù)過(guò)程對(duì)物種緩解前期脅迫的影響至關(guān)重要。本試驗(yàn)第3階段，移除水分脅迫后，CD、HS、HSD、FSD組水杉樹(shù)苗的CAT、ASP活性顯著降低或與C組相當(dāng)，MDA含量顯著低于C組，與木果楝(Carapaguianensis)[46]的表現(xiàn)相似，表明水杉樹(shù)苗在恢復(fù)供水階段H2O2的形成量下降，對(duì)水分脅迫表現(xiàn)出良好的適應(yīng)能力[46]。

水分脅迫去除后，恢復(fù)的程度及幅度取決于脅迫的強(qiáng)度、持續(xù)時(shí)間以及受脅迫的物種種類(lèi)[49]。本試驗(yàn)中，在干旱脅迫下，各個(gè)處理組水杉樹(shù)苗的存活率均達(dá)100%。在去除干旱脅迫后，不同處理的水杉樹(shù)苗抗氧化酶活性及滲透調(diào)節(jié)物質(zhì)含量顯著降低或與C組相當(dāng)，這與Verma等[50]對(duì)麻瘋樹(shù)(Jatrophacurcas)的研究結(jié)果相似，其原因可能是前期的干旱脅迫時(shí)間過(guò)長(zhǎng)或是恢復(fù)時(shí)間不夠[51]，具體原因還有待于進(jìn)一步研究。

4 結(jié) 論

本研究發(fā)現(xiàn)，前期水淹并未影響水杉樹(shù)苗對(duì)后期干旱脅迫的生理生化響應(yīng)。水杉樹(shù)苗對(duì)土壤水分變化具有較強(qiáng)的生理生化響應(yīng)能力，不僅表現(xiàn)出耐水濕的特點(diǎn)，而且表現(xiàn)出一定的耐旱性。在水分脅迫下，水杉樹(shù)苗能有效誘導(dǎo)體內(nèi)的保護(hù)酶系統(tǒng)，清除體內(nèi)的活性氧等自由基，在一定程度上維持了植株正常的生理代謝活動(dòng)。同時(shí)，其還通過(guò)體內(nèi)滲透調(diào)節(jié)物質(zhì)的積累，降低了細(xì)胞的水勢(shì)，起到了滲透調(diào)節(jié)的作用。正是保護(hù)酶系活性的啟動(dòng)以及滲透調(diào)節(jié)物質(zhì)的產(chǎn)生，使得水杉樹(shù)苗的耐淹性和耐旱性得以提高，從而具有較強(qiáng)的滲透脅迫耐受能力。因此，在三峽庫(kù)區(qū)植被恢復(fù)建設(shè)中，可以考慮將水杉樹(shù)苗作為候選樹(shù)種之一。

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Effects of flooding on responses ofMetasequoiaglyptostroboidessaplings to drought

BAI Lin-li,HAN Wen-jiao,LI Chang-xiao

(KeyLaboratoryfortheEco-EnvironmentoftheThreeGorgesReservoirRegionoftheMinistryofEducation,CollegeofLifeSciences,SouthwestUniversity,Chongqing400715,China)

water stress;Metasequoiaglyptostroboides;physiology and biochemistry;the hydro-fluctuation belt;the Three Gorges Reservoir Area

2014-07-16

重慶市基礎(chǔ)與前沿研究計(jì)劃重點(diǎn)項(xiàng)目(CSTC2013JJB00004)；中央高?；究蒲袠I(yè)務(wù)費(fèi)專(zhuān)項(xiàng)(XDJK2013A011)；國(guó)家林業(yè)公益性行業(yè)科研專(zhuān)項(xiàng)(201004039)；留學(xué)回國(guó)人員科研啟動(dòng)基金項(xiàng)目(教外司留[2010-1561])

白林利(1990-)，女，山西呂梁人，在讀碩士，主要從事環(huán)境生態(tài)學(xué)研究。E-mail：895845358@qq.com

李昌曉(1972-)，男，重慶人，教授，博士生導(dǎo)師，主要從事生態(tài)修復(fù)研究。E-mail：lichangx@swu.edu.cn

時(shí)間:2015-04-13 12:59

10.13207/j.cnki.jnwafu.2015.05.023

S718.51+2.3;Q945.78

A

1671-9387(2015)05-0042-09

網(wǎng)絡(luò)出版地址:http://www.cnki.net/kcms/detail/61.1390.S.20150413.1259.023.html