熊強(qiáng)強(qiáng),鐘 蕾,沈天花,陳小榮,朱昌蘭,彭小松,傅軍如,賀浩華
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穗分化期旱澇急轉(zhuǎn)對雙季超級雜交稻物質(zhì)積累和產(chǎn)量形成的影響*
熊強(qiáng)強(qiáng),鐘 蕾,沈天花,陳小榮**,朱昌蘭,彭小松,傅軍如,賀浩華
(江西農(nóng)業(yè)大學(xué)農(nóng)學(xué)院/作物生理生態(tài)與遺傳育種教育部重點(diǎn)實(shí)驗(yàn)室/江西省超級稻工程技術(shù)研究中心/雙季稻現(xiàn)代化生產(chǎn)協(xié)同中心,南昌 330045)
采用桶栽方式,以超級雜交早稻品種(淦鑫203)、超級雜交晚稻品種(五豐優(yōu)T025)為材料,于穗分化初期設(shè)置不旱不澇(對照CK)、輕旱不澇(T1)、重旱不澇(T2)、不旱輕澇(T3)、不旱重澇(T4)、重旱輕澇(T5)、重旱重澇(T6)、輕旱輕澇(T7)和輕旱重澇(T8)處理,分析各處理產(chǎn)量及其構(gòu)成因素、干物質(zhì)積累與分配和凈光合速率等指標(biāo)。結(jié)果表明:(1)“旱”、“澇”及“旱澇急轉(zhuǎn)”使超級雜交早、晚稻單株產(chǎn)量不同程度下降,早稻T1、T2、T3、T4、T5、T6、T7和T8單株產(chǎn)量分別比CK降低8.89%、21.42%、8.33%、12.04%、19.65%、31.23%、15.39%和17.29%,晚稻分別降低6.21%、15.06%、7.77%、11.53%、17.40%、20.85%、13.46%和11.70%,早、晚稻以T2和T6處理產(chǎn)量下降較多,尤其T6處理產(chǎn)量降幅最大,表明單純前期輕旱(T1)和后期輕澇(T3)對產(chǎn)量的影響相對較小,而單純前期重旱(T2)和后期重澇(T4)對產(chǎn)量的影響相對較大,T2比T4對產(chǎn)量的影響更大,且重旱重澇處理下水稻單株產(chǎn)量存在一定程度的疊加減產(chǎn)效應(yīng)。產(chǎn)量構(gòu)成上,早、晚稻產(chǎn)量下降的主要原因是較大幅度降低了單株有效穗數(shù)、每穗總粒數(shù)、結(jié)實(shí)率和二次枝梗數(shù),各處理對產(chǎn)量構(gòu)成的影響規(guī)律與單株產(chǎn)量大致相同。(2)早、晚稻“旱”、“澇”及“旱澇急轉(zhuǎn)”等處理結(jié)束初期與CK比較,其莖、葉干物質(zhì)量均不同程度下降,其中早、晚稻成熟期T6處理穗干物質(zhì)量分別比CK下降33.40%、19.88%,表現(xiàn)為穗型變小。(3)“旱”、“澇”及“旱澇急轉(zhuǎn)”使早、晚稻葉片葉綠素含量(SPAD值)降低,其中早、晚稻均以T6處理的SPAD值降幅最大,且與CK一直呈顯著性差異。(4)至抽穗期,早、晚稻葉片凈光合速率均以T6處理最低,分別比CK下降37.57%、10.17%。研究結(jié)果說明,生長前期重旱后期急轉(zhuǎn)重澇對早、晚稻成熟期穗部干物質(zhì)量及總干物質(zhì)量的影響最大,重旱重澇急轉(zhuǎn)表現(xiàn)為疊加損傷效應(yīng)。
雙季超級雜交稻;穗分化期;旱澇急轉(zhuǎn);產(chǎn)量;干物質(zhì);光合速率
干旱、淹澇災(zāi)害,作為主要農(nóng)業(yè)自然災(zāi)害之一,長期影響著中國農(nóng)業(yè)生產(chǎn)[1-2]。江西等南方雙季稻區(qū)旱、澇災(zāi)害近幾十年來發(fā)生較頻繁,整體上表現(xiàn)為明顯的增長趨勢[3],特別是干旱后緊接著出現(xiàn)洪澇災(zāi)害,即“旱澇急轉(zhuǎn)”事件,造成水稻大面積減產(chǎn)甚至絕收時有發(fā)生[4]。旱澇急轉(zhuǎn)是指前期持續(xù)偏旱,接著因一場暴雨以上的強(qiáng)降水或雨量較大的連陰雨致使迅速轉(zhuǎn)旱為澇的天氣過程[5],該氣象災(zāi)害易造成農(nóng)作物大面積減產(chǎn)、減收,故引起人們極大關(guān)注[5-6]。統(tǒng)計(jì)表明,1960-2012年中國共有14a出現(xiàn)了“旱澇急轉(zhuǎn)”氣象災(zāi)害,約4a一遇,淮河流域總共發(fā)生23次“旱澇急轉(zhuǎn)”事件[4],其中2011年6月全國范圍遭受“旱澇急轉(zhuǎn)”自然災(zāi)害,對水稻生產(chǎn)造成了很大影響。近年來科研工作者有關(guān)穗分化期“旱澇急轉(zhuǎn)”對水稻產(chǎn)量及其構(gòu)成因子、有關(guān)生理特征特性等影響已有一些研究報道[7-9],也有從農(nóng)業(yè)工程(水利)等角度對水稻旱澇交替條件下產(chǎn)量形成方面的研究報道[10-11]。然而,有關(guān)不同程度“旱”、“澇”及“旱澇急轉(zhuǎn)”組合對水稻產(chǎn)量形成、生長發(fā)育及物質(zhì)積累的問題卻尚未引起足夠重視,有關(guān)不同程度“旱澇急轉(zhuǎn)”組合對水稻產(chǎn)量形成機(jī)制的研究不夠深入,關(guān)于旱后淹澇到底將產(chǎn)生疊加損傷還是拮抗作用爭議較大[12]。另一方面,近年來在南方雙季稻區(qū)推廣了一大批超級雜交稻品種,推動了增產(chǎn)增收及品種的更新?lián)Q代,這些超級稻品種物質(zhì)生產(chǎn)能力及產(chǎn)量潛力水平得到較大改善,但對水肥需求更大,對水分、氮素虧缺等更為敏感[13-14],探討不同程度“旱”、“澇”,尤其是“旱澇急轉(zhuǎn)”組合處理對雙季超級雜交稻產(chǎn)量及其干物質(zhì)積累與分配的影響尤為必要。
本研究選用江西等雙季稻區(qū)近年來大面積種植的超級雜交早稻品種淦鑫203,超級雜交晚稻品種五豐優(yōu)T025,選擇模擬該地常年發(fā)生“旱澇急轉(zhuǎn)”災(zāi)害天氣時雙季稻所處的生育階段,于幼穗分化初期設(shè)置不同程度的“旱”、“澇”及“旱澇急轉(zhuǎn)”組合處理,分析雙季超級雜交稻“旱”、“澇”及多種不同程度“旱澇急轉(zhuǎn)”組合下的水稻產(chǎn)量及其構(gòu)成、干物質(zhì)積累與分配和光合速率等指標(biāo)的差異,以期為該氣象災(zāi)害的抗災(zāi)減災(zāi)、水分合理利用提供科學(xué)依據(jù)。
1.1 試驗(yàn)材料
2個供試水稻品種均為秈型雜交組合,江西省主栽品種。淦鑫203為早稻類型,2009年被認(rèn)定為超級稻;五豐優(yōu)T025為晚稻類型,2010年被認(rèn)定為超級稻。
1.2 試驗(yàn)設(shè)計(jì)
1.2.1 栽培方法
試驗(yàn)于2015年在江西農(nóng)業(yè)大學(xué)科技園網(wǎng)室內(nèi)進(jìn)行,均采用桶栽方式。塑料桶桶高為24.0cm、上部外徑29.0 cm、底部內(nèi)徑23.5cm。栽培土壤取自稻田0-20cm耕層土壤,將其自然晾曬風(fēng)干,用FT-1000A土壤粉碎機(jī)(中國產(chǎn))粉碎過100目網(wǎng)篩,移栽前預(yù)先淹水2周。早稻3月20日播種,4月23日移栽,晚稻6月25日播種,7月23日移栽;統(tǒng)一于四葉一心期,選取長勢良好、一致的秧苗移栽,每桶栽插3穴,每穴1株苗;每桶裝風(fēng)干土10kg,拌施底肥施復(fù)合肥(N-P-K=15%-15%-15%)5g,移栽7d后每桶追施2g氯化鉀,2g尿素。移栽前大田統(tǒng)一管理,移栽至塑料桶后均按高產(chǎn)栽培方式進(jìn)行水分、病蟲害管理,待“旱”、“澇”及“旱澇急轉(zhuǎn)”處理結(jié)束后,均按水稻桶栽正常水分(保持水層3~5cm)及病蟲害防治管理。
1.2.2 試驗(yàn)設(shè)計(jì)
干旱、淹澇及“旱澇急轉(zhuǎn)”處理起止日期、持續(xù)天數(shù)主要根據(jù)長江流域雙季稻區(qū)常發(fā)生的“旱”、“澇”及“旱澇急轉(zhuǎn)”災(zāi)害天氣及前人研究[5,15-17]而設(shè)定,即模擬長江流域雙季稻區(qū)實(shí)際生產(chǎn)過程中旱、澇情況,重旱一般為干旱至土壤發(fā)白開裂,植株萎蔫干枯;重澇一般為淹澇6~8d。一般雙季早稻于5月下旬,雙季晚稻于8月下旬,此時稻株剛進(jìn)入穗分化初期。早稻于2015年5月23日、晚稻從8月21日開始進(jìn)行干旱處理,即倒去桶內(nèi)明水后移至防雨棚,安裝真空表式土壤濕度計(jì)(測定范圍0~85kPa,中國科學(xué)院某研究所)監(jiān)測土壤含水率,水稻自然落干。早稻幼穗分化期干旱處理第8天的土壤水勢超過最大測量值,晚稻幼穗分化期干旱處理第6天的土壤水勢超過最大測量值,繼續(xù)統(tǒng)一干旱至土壤發(fā)白開裂,植株呈萎蔫干枯狀(重旱),當(dāng)“旱澇急轉(zhuǎn)”處理達(dá)到設(shè)定的干旱級別后立即開始淹澇處理,將需要進(jìn)行淹澇試驗(yàn)的盆桶移至水深100cm高的水箱,受澇試驗(yàn)水箱中水為靜止、潔凈的自來水,受澇期間不換水,并完全浸沒稻株;對照(CK)一直保持3~5cm水層。試驗(yàn)共設(shè)9個處理,不旱不澇(CK)、輕旱不澇(T1)、重旱不澇(T2)、不旱輕澇(T3)、不旱重澇(T4)、重旱輕澇(T5)、重旱重澇(T6)、輕旱輕澇(T7)和輕旱重澇(T8),每處理重復(fù)3次,每個重復(fù)15桶,隨機(jī)區(qū)組排列。各處理持續(xù)天數(shù)和日期見表1。
表1 超級雜交早、晚稻幼穗分化期旱澇處理設(shè)置方案
注:CK為不旱不澇(對照);T1為輕旱不澇;T2為重旱不澇;T3為不旱輕澇;T4為不旱重澇;T5為重旱輕澇;T6為重旱重澇;T7為輕旱輕澇;T8為輕旱重澇。下同。
Note: CK is no drought in earlier stage and no floods in later stage(control); T1 is mild drought in earlier stage and no floods in later stage; T2 is severe drought in earlier stage and no floods in later stage; T3 is no drought in earlier stage and mild floods in later stage; T4 is no drought in earlier stage and severe floods in later stage; T5 is severe drought in earlier stage and mild floods in later stage abrupt alternation; T6 is severe drought in earlier stage and severe floods in later stage abrupt alternation; T7 is mild drought in earlier stage and mild floods in later stage abrupt alternation; T8 is mild drought in earlier stage and severe floods in later stage abrupt alternation. The same as below.
1.3 項(xiàng)目測定
1.3.1 凈光合速率
早、晚稻于“旱澇急轉(zhuǎn)”處理結(jié)束水稻生長至抽穗期,選擇晴朗無云的天氣,于上午9:00-11:00,每重復(fù)選取3~5株生長良好、長勢一致的水稻葉片掛牌標(biāo)記,用 CI-340便攜式光合測定儀(美國)測定倒2葉中部凈光合速率(net photosynthetic rate,Pn)。
1.3.2 SPAD值
早稻分別于“旱澇急轉(zhuǎn)”處理結(jié)束后1d(06-11)、8d(06-18)、12d(06-22),晚稻分別于“旱澇急轉(zhuǎn)”處理結(jié)束后2d(09-06)、8d(09-12)、13d(09-17),選取6~8株生長良好、長勢一致水稻的倒2葉葉片,采用SPAD-502葉綠素測定儀(浙江)測定葉片基部、中部和頂部的SPAD值,取其平均值。
1.3.3 干物質(zhì)
早稻于“旱澇急轉(zhuǎn)”處理結(jié)束初期即穗分化中后期(06-11)、抽穗期(06-22)、成熟期(07-14),晚稻于“旱澇急轉(zhuǎn)”處理剛結(jié)束初期即穗分化中后期(09-04)、抽穗期(09-16)、成熟期(10-18),每處理每重復(fù)取水稻3株,取樣后于實(shí)驗(yàn)室用清水清洗水稻植株,按器官部位莖、葉、穗分樣,在105℃殺青30min后再置于70℃烘箱烘至恒質(zhì)量,測定各器官干物質(zhì)量。
1.3.4 產(chǎn)量及產(chǎn)量結(jié)構(gòu)
成熟后各處理每重復(fù)取5株未損傷的水稻植株收獲考種,分別考察穗長、穗實(shí)粒數(shù)、穗空粒數(shù)、千粒重、一次枝梗數(shù)、二次枝梗數(shù)、單株有效穗數(shù)、每篼產(chǎn)量及生物量。計(jì)算每穗總粒數(shù)、結(jié)實(shí)率和收獲指數(shù)。
1.4 數(shù)據(jù)處理
使用SPSS17.0和Microsoft Excel 2003軟件對有關(guān)數(shù)據(jù)進(jìn)行分析和作圖。
2.1 “旱澇急轉(zhuǎn)”對超級雜交稻產(chǎn)量的影響
由表2可看出,與CK相比,僅前期干旱而后期不澇(T1、T2)和前期不旱僅后期澇(T3、T4)的處理中,無論早稻(淦鑫203)還是晚稻(五豐優(yōu)T025)單株產(chǎn)量及相應(yīng)的產(chǎn)量構(gòu)成性狀均有不同程度的降低;從數(shù)值上看,單純前期輕旱(T1)和后期輕澇(T3)對產(chǎn)量的影響相對較小,而且影響程度相當(dāng),早稻產(chǎn)量分別比對照(CK)減少8.89%和8.33%,晚稻產(chǎn)量分別比對照(CK)減少6.21%和7.77%,差異均不顯著;而單純前期重旱(T2)和后期重澇(T4)對產(chǎn)量的影響相對較大,早稻產(chǎn)量分別比對照(CK)減少21.42%和12.04%,晚稻產(chǎn)量分別比對照(CK)減少15.06%和11.53%。除了水稻一次枝梗數(shù),其它產(chǎn)量構(gòu)成性狀的影響規(guī)律與此大致相同,早稻T1、T2和T4處理的一次枝梗數(shù)均大于CK,晚稻T1、T2、T3和T4處理一次枝梗數(shù)均大于CK。
分析“旱澇急轉(zhuǎn)”情形對超級雜交稻產(chǎn)量的影響可見(表2),與CK相比,前期重旱狀態(tài)下后期急轉(zhuǎn)為輕澇的處理(T5)、前期重旱后期急轉(zhuǎn)重澇的處理(T6)中,無論早稻(淦鑫203)還是晚稻(五豐優(yōu)T025)單株產(chǎn)量及相應(yīng)的產(chǎn)量構(gòu)成均有更大程度的降低;重旱重澇(T6)早稻產(chǎn)量僅為CK的68.77%,是單純干旱(重旱)狀態(tài)下的87.52%,晚稻僅為CK的79.15%,是單純干旱(重旱)狀態(tài)下的93.17%;重旱轉(zhuǎn)輕澇(T5)早稻產(chǎn)量僅為CK的80.35%,是單純干旱(重旱)狀態(tài)下的102.25%,晚稻僅為CK的82.60%,是單純干旱(重旱)狀態(tài)下的97.25%;早、晚稻重旱重澇處理(T6)產(chǎn)量下降程度最大,表明重旱重澇處理下水稻單株產(chǎn)量存在一定程度的疊加減產(chǎn)效應(yīng),除千粒重和一次枝梗數(shù),其它產(chǎn)量構(gòu)成性狀的影響規(guī)律與此大致相同。
表2 各處理超級雜交早、晚稻產(chǎn)量及其構(gòu)成因素的比較
注:小寫字母表示處理間在0.05水平上的差異顯著性。下同。
Note:Lowercase indicates the difference significance among treatments at 0.05 level. TR is treatment, YP is yield per plant, GW is 1000-grain weight, NP is number of grains per panicle, SR is seed setting rate, EP is effective panicles per plant, PN is primary branch number, SN secondary branch number, EL is ear length, HI is harvest index. The same as below.
2.2 “旱澇急轉(zhuǎn)”對超級雜交稻地上部分物質(zhì)積累與分配的影響
從表3可看出,與CK相比,僅前期干旱而后期不澇(T1、T2)和前期不旱僅后期澇(T3、T4)的處理中,在處理結(jié)束初期,無論早稻(淦鑫203)還是晚稻(五豐優(yōu)T025)其莖、葉干物質(zhì)量均有不同程度的降低;抽穗階段主要進(jìn)行生殖生長,干物質(zhì)分配到生殖器官的比例逐漸升高,前期干旱而后期不澇(T1、T2)和前期不旱僅后期澇(T3、T4)的處理,無論早稻(淦鑫203)還是晚稻(五豐優(yōu)T025)穗部干物質(zhì)量均呈不同程度的降低,早稻單純前期輕旱(T1)和后期輕澇(T3)處理的穗部干物質(zhì)量分別比對照(CK)減少0.68%和45.25%,晚稻穗部干物質(zhì)量分別比對照(CK)減少3.43%和5.66%;早稻單純前期重旱(T2)和后期重澇(T4)處理的穗部干物質(zhì)量分別比對照(CK)減少3.39%和16.78%,晚稻穗部干物質(zhì)量分別比對照(CK)減少39.80%和5.05%,表明“旱”、“澇”水分脅迫處理對水稻莖、葉積累影響較大,且前期重旱比后期重澇對穗部干物質(zhì)量的影響更大一些,穗部干物質(zhì)量下降較多;成熟期時,各干旱處理,尤以單純前期重旱(T2)處理其穗部干物質(zhì)量下降較明顯,穗型變小,與CK相比,早稻穗部干物質(zhì)量下降24.90%,晚稻穗部干物質(zhì)量下降13.98%,并且單純前期重旱對總干物質(zhì)量的影響規(guī)律與此大致相同。
表3 各處理超級雜交早、晚稻各器官干物質(zhì)積累與分配的比較(g)
分析“旱澇急轉(zhuǎn)”情形對超級雜交稻干物質(zhì)分配的影響可見(表3),與CK相比,處理結(jié)束初期,前期重旱后期急轉(zhuǎn)重澇的處理(T6)中,無論早稻(淦鑫203)還是晚稻(五豐優(yōu)T025)其莖、葉干物質(zhì)量均有不同程度的降低,早稻莖、葉干物質(zhì)量分別比對照(CK)減少50.71%和31.08%,晚稻莖、葉干物質(zhì)量分別比對照(CK)減少33.48%和20.48%,且早、晚稻T6處理的莖、葉干物質(zhì)量較其它“旱澇急轉(zhuǎn)”處理莖、葉干物質(zhì)量下降最多;至抽穗期,晚稻前期輕旱急轉(zhuǎn)為輕澇的處理(T7)和前期輕旱急轉(zhuǎn)為重澇的處理(T8)其莖、葉干物質(zhì)量均高于CK,但穗部干物質(zhì)量分別比對照(CK)減少6.87%和15.35%,重旱重澇(T6)穗部干物質(zhì)量分別比對照(CK)減少58.99%,下降最為顯著;抽穗-成熟期階段為生產(chǎn)上水稻奪取高產(chǎn)的主要階段,水稻植株生產(chǎn)的干物質(zhì)主要轉(zhuǎn)移向穗部,“旱澇急轉(zhuǎn)”各處理穗干物質(zhì)量下降幅度較大,穗型變小,早稻T5、T6、T7和T8成熟期穗部干物質(zhì)量、總干物質(zhì)量分別比對照(CK)減少22.18%、33.40%、20.40%和19.90%,19.56%、32.10%、18.27%和20.88%,晚稻T5、T6、T7和T8成熟期穗部干物質(zhì)量、總干物質(zhì)量分別比對照(CK)減少14.61%、19.88%、13.22%和8.85%,12.46%、20.37%、14.30%和9.45%,且早、晚稻重旱重澇處理(T6)成熟期穗部干物質(zhì)量為單純重旱處理(T2)的87.52%、93.14%,總干物質(zhì)量也呈相應(yīng)的下降,由此可知,早、晚稻前期重旱后期急轉(zhuǎn)重澇的處理對水稻成熟期穗部干物質(zhì)量及總干物質(zhì)量影響最大。
2.3 “旱澇急轉(zhuǎn)”對超級雜交稻葉片SPAD值的影響
圖1可看出,與CK相比,僅前期干旱而后期不澇(T1、T2)或前期不旱僅后期澇(T3、T4)的處理中,在處理結(jié)束初期,無論早稻(淦鑫203)還是晚稻(五豐優(yōu)T025)葉綠素含量(SPAD值)均有不同程度的降低,早稻單純前期輕旱(T1)和后期輕澇(T3)處理的SPAD值分別比對照(CK)減少2.68%和0.61%,晚稻SPAD值分別比對照(CK)減少10.34%和11.73%,早稻單純前期重旱(T2)和后期重澇(T4)處理的SPAD值分別比對照(CK)減少4.43%和1.33%,晚稻葉綠素分別比對照(CK)減少9.90%和7.54%,表明水稻單純旱、澇脅迫下葉片受損,造成短時間內(nèi)SPAD值大幅下降,但隨著生育期推進(jìn)各處理與對照差距縮小。
分析“旱澇急轉(zhuǎn)”情形對超級雜交稻葉片葉綠素的影響可見(圖1),前期重旱后期急轉(zhuǎn)重澇的處理(T6)中,在處理結(jié)束初期,無論早稻(淦鑫203)還是晚稻(五豐優(yōu)T025)其SPAD值有較大程度的降低,且均達(dá)顯著性差異,早稻重旱重澇(T6)處理期SPAD值僅為CK的91.51%,晚稻重旱重澇(T6)處理其SPAD值僅為CK的88.20%,在處理后期早、晚稻重旱重澇(T6)葉片SPAD值在各處理中一直最低,且與CK差異均達(dá)顯著水平。表明水稻在重旱重澇急轉(zhuǎn)脅迫下葉片受損嚴(yán)重,造成葉綠素大幅降解,難以恢復(fù)至正常水平。
注:圖中柱狀數(shù)據(jù)為平均值,短線表示均方差。下同
Note: The column data in figure is average and the short lines represents the mean square deviation. The same as below
2.4 “旱澇急轉(zhuǎn)”對超級雜交稻葉片凈光合速率的影響
圖2可看出,與CK相比,僅前期干旱而后期不澇(T1、T2)或前期不旱僅后期澇(T3、T4)的處理中,在處理結(jié)束初期,無論早稻(淦鑫203)還是晚稻(五豐優(yōu)T025)其凈光合速率均有不同程度的降低,從數(shù)值上看,單純前期輕旱(T1)和后期輕澇(T3)對凈光合速率的影響相對較小,早稻凈光合速率分別比對照(CK)減少7.78%和23.72%,晚稻凈光合速率分別比對照(CK)減少5.81%和5.08%;而單純前期重旱(T2)和后期重澇(T4)對凈光合速率的影響相對較大,而且前期重旱比后期重澇對產(chǎn)量的影響更大一些,凈光合速率下降更多,早稻凈光合速率分別比對照(CK)減少30.36%和31.65%,晚稻凈光合速率分別比對照(CK)減少7.36%和7.80%。表明幼穗分化期單純重旱或者重澇對水稻葉片損傷更大,致使在抽穗期時凈光合速率下降程度更大。
分析“旱澇急轉(zhuǎn)”情形對超級雜交稻葉片凈光合速率的影響可見(圖2),前期重旱后期急轉(zhuǎn)為輕澇的處理(T5)和前期重旱后期急轉(zhuǎn)重澇的處理(T6),無論早稻(淦鑫203)還是晚稻(五豐優(yōu)T025)其凈光合速率均有更大程度的降低;重旱重澇處理(T6)早稻凈光合速率僅為CK的62.43%,兩者差異顯著,是重旱不澇處理(T2)的89.65%;晚稻僅為CK的91.77%,兩者差異顯著,是重旱不澇處理(T2)的96.97%;重旱轉(zhuǎn)輕澇處理(T5)早稻凈光合速率僅為CK的75.90%,是重旱不澇處理(T2)的108.99%,晚稻僅為CK的89.83%,是重旱不澇處理(T2)的99.06%,說明稻株重旱重澇處理比單純重旱和重旱輕澇處理的葉片凈光合速率下降更多,表明重旱重澇急轉(zhuǎn)處理對水稻葉片凈光合速率具有一定的疊加損傷作用,前期重旱后期急轉(zhuǎn)為輕澇對水稻影響相對較小。
3.1 穗分化期“旱澇急轉(zhuǎn)”對超級雜交稻產(chǎn)量及其構(gòu)成的影響
已有研究指出[16,18],水稻孕穗期是水分臨界期,對水分最為敏感,水稻幼穗分化期干旱處理導(dǎo)致水稻相關(guān)產(chǎn)量構(gòu)成因素發(fā)生變化,從而導(dǎo)致產(chǎn)量的下降。另有研究指出[15,17,19],淹澇脅迫致使水稻產(chǎn)量下降,不同生育期受澇水稻減產(chǎn)順序表現(xiàn)為孕穗期>拔節(jié)期>灌漿期,其中孕穗期受到淹澇脅迫對產(chǎn)量的影響最大,但其與淹澇深度和受澇天數(shù)有關(guān)。本試驗(yàn)結(jié)果顯示,幼穗分化期“旱”、“澇”處理使超級雜交早、晚稻單株產(chǎn)量不同程度降低,且單純重旱比單純輕旱、單純重澇比單純輕澇影響較大,單株產(chǎn)量下降較多,這與前人研究[15-17]結(jié)果基本一致。從產(chǎn)量構(gòu)成上看,夏瓊梅等[20]研究發(fā)現(xiàn),幼穗分化-齊穗期水分脅迫后產(chǎn)量下降的主要原因是有效穗數(shù)、總粒數(shù)、實(shí)粒數(shù)的降低,其中對實(shí)粒數(shù)影響最大,其次是總粒數(shù)。本試驗(yàn)發(fā)現(xiàn),經(jīng)過重旱處理的雙季超級雜交早、晚稻重旱不澇(T2)、重旱輕澇(T5)和重旱重澇(T6)處理單株產(chǎn)量下降幅度大,產(chǎn)量下降主要原因是單株有效穗數(shù)、每穗總粒數(shù)、結(jié)實(shí)率和每穗二次枝梗數(shù)降幅較大,這與夏瓊梅等研究結(jié)果基本一致。關(guān)于旱后淹澇急轉(zhuǎn)將產(chǎn)生疊加損傷或是拮抗作用有較大爭議[12],本試驗(yàn)結(jié)果顯示,前期重旱狀態(tài)下后期急轉(zhuǎn)為輕澇的處理(T5)、前期重旱后期急轉(zhuǎn)重澇的處理(T6)中,早、晚稻單株產(chǎn)量及相應(yīng)的產(chǎn)量構(gòu)成均有更大程度的降低,尤其早、晚稻重旱重澇急轉(zhuǎn)處理單株產(chǎn)量最低,同時說明重旱重澇處理下水稻單株產(chǎn)量存在一定程度的疊加減產(chǎn)效應(yīng),但前期重旱后期急轉(zhuǎn)為輕澇對水稻影響相對較小。造成疊加損傷效應(yīng)使產(chǎn)量下降的主要原因?yàn)槊克肟偭?shù)、結(jié)實(shí)率、單株有效穗數(shù)和二次枝梗數(shù)的降低。本試驗(yàn)還發(fā)現(xiàn),重旱處理后水稻葉片葉尖損傷較嚴(yán)重,接近1/3葉片枯萎死亡,緊接著重澇使水稻植株小蘗死亡較多,葉片損傷較嚴(yán)重。而僅進(jìn)行淹澇脅迫時,澇后植株生長(莖、節(jié)間)過快,劍葉伸出水面,有氧呼吸作用加強(qiáng),因此,不旱輕澇的產(chǎn)量下降程度較小,且淹澇時間越長對水稻單株產(chǎn)量影響越大,這與前人研究結(jié)果一致[21]。說明水稻在“旱”、“澇”及“旱澇急轉(zhuǎn)”脅迫下的生長發(fā)育將不同程度受到損害,尤其重旱重澇處理后,植株難以恢復(fù)而導(dǎo)致減產(chǎn),這也可從本試驗(yàn)中葉片SPAD值的變化得到印證,尤其是重旱重澇急轉(zhuǎn),至處理結(jié)束后很長一段時期內(nèi)其葉片SPAD值仍顯著低于對照CK,難以恢復(fù)至正常水平。與前人研究相比[22-23],本試驗(yàn)中千粒重受影響較小,有些處理還略有增加。造成研究結(jié)果差異的原因可能是“旱”、“澇”及“旱澇急轉(zhuǎn)”脅迫處理時期和持續(xù)時長不同,本研究為了模擬長江中下游“旱澇急轉(zhuǎn)”氣象災(zāi)害,至抽穗灌漿后期部分植株基本恢復(fù),特別是早稻輕旱輕澇,晚稻輕旱不澇和不旱輕澇3個處理。另外稻株千粒重的不同也可能因?yàn)槿哟嬖谝欢ǖ恼`差。然而,從本研究結(jié)果也可看出,即使在水稻幼穗分化期這樣一個對水分極為敏感的階段出現(xiàn)上述較為嚴(yán)重的“旱”、“澇”及“旱澇急轉(zhuǎn)”脅迫,也未出現(xiàn)絕收情況,可見在生產(chǎn)實(shí)際中遭遇旱災(zāi)后,應(yīng)積極采取抗旱保收措施,進(jìn)行補(bǔ)水保苗;而對于淹澇災(zāi)害,應(yīng)及時排水,同時輔以必要的災(zāi)后管理措施。由于本研究采用桶栽模擬長江中下游“旱澇急轉(zhuǎn)”氣象災(zāi)害進(jìn)行試驗(yàn),可能與大田下有差異,大田種植條件下“旱澇急轉(zhuǎn)”脅迫對水稻產(chǎn)量及其構(gòu)成的變化和機(jī)理還需深入研究,這對于減輕“旱澇急轉(zhuǎn)”對水稻的危害具有十分重要的意義。
3.2 穗分化期“旱澇急轉(zhuǎn)”對超級雜交稻地上部分物質(zhì)積累和分配的影響
水分是限制作物生產(chǎn)的重要因素,“旱”、“澇”及“旱澇急轉(zhuǎn)”導(dǎo)致植株損傷,從而影響生理生化過程和器官建成,進(jìn)而影響生長發(fā)育。大量研究表明[19, 24-25],水分脅迫會干擾植物的物質(zhì)生產(chǎn)。李樹杏等研究指出[24],水稻幼穗形成期經(jīng)過水分脅迫復(fù)水后,短歷時輕、重度脅迫的處理抽穗期莖鞘干物質(zhì)量高于對照。本試驗(yàn)結(jié)果顯示,晚稻輕旱不澇(T1)復(fù)水后在抽穗期莖干物質(zhì)量高于對照,但與對照CK差異顯著,與李樹杏等研究結(jié)果基本一致,其輕旱不澇(T1)處理的凈光合速率與對照CK無顯著性差異;早稻輕旱不澇處理(T1)則未出現(xiàn)類似結(jié)果,可能是由于早稻品種輕旱持續(xù)6d相對來說時間較長,旱后復(fù)水恢復(fù)時間延長,與此同時輕旱不澇(T1)處理的凈光合速率較對照降低5.81%,但無顯著性差異。抽穗-成熟期階段為生產(chǎn)上水稻奪取高產(chǎn)的主要階段,水稻植株生產(chǎn)的干物質(zhì)主要轉(zhuǎn)移至穗部,本研究發(fā)現(xiàn),在抽穗期,早稻重旱重澇(T6)處理穗部干物質(zhì)量分別比對照(CK)減少51.19%,晚稻重旱重澇(T6)處理穗部干物質(zhì)量分別比對照(CK)減少58.99%,重旱重澇處理穗部干物質(zhì)量下降程度最大,該結(jié)果與抽穗期時葉片凈光合速率得到印證,抽穗期早、晚稻重旱重澇(T6)處理凈光合速率較對照CK下降37.57%、10.17%,且均與CK差異顯著。黃元財?shù)萚25]研究指出,水分脅迫條件下,各器官中以葉片干物質(zhì)凈積累量的下降趨勢最為明顯,成熟期水稻穗部干物質(zhì)比率的大幅提高而莖鞘和葉片的比率下降,植株生產(chǎn)的干物質(zhì)主要運(yùn)向穗部。本試驗(yàn)結(jié)果還顯示,超級雜交早、晚稻在處理結(jié)束初期和成熟期,與對照CK相比,莖、葉干物質(zhì)量呈不同程度的下降,這與前人研究結(jié)論一致[25]。收獲指數(shù)是指籽粒產(chǎn)量與成熟期植株干物質(zhì)質(zhì)量的比率,其生理本質(zhì)反映了作物同化產(chǎn)物在籽粒和營養(yǎng)器官上的分配比例,其高低反映水稻營養(yǎng)生長與生殖生長是否協(xié)調(diào),光合產(chǎn)物的積累與分配是否合理的一種標(biāo)志[26]。抽穗-成熟階段為生產(chǎn)上水稻奪取高產(chǎn)的主要階段,水稻植株生產(chǎn)的干物質(zhì)主要轉(zhuǎn)移向穗部,而由于在穗分化期進(jìn)行了“旱”、“澇”及“旱澇急轉(zhuǎn)”處理,與對照CK相比,早、晚稻“旱澇急轉(zhuǎn)”各處理穗干物質(zhì)量降幅較大,穗型變小,早稻T5、T6、T7和T8成熟期穗部干物質(zhì)量、總干物質(zhì)量均呈不同程度的降低,說明穗分化期“旱”、“澇”及“旱澇急轉(zhuǎn)”處理不僅影響水稻物質(zhì)生產(chǎn)能力,也影響干物質(zhì)向籽粒的轉(zhuǎn)化與分配。本試驗(yàn)研究還表明,超級雜交早、晚稻有的水分干旱脅迫處理下抽穗期莖、葉物質(zhì)積累高于對照CK,可能與進(jìn)行了“旱”、“澇”及“旱澇急轉(zhuǎn)”處理后營養(yǎng)生長較快,生殖生長滯后有關(guān),但在成熟期總干物質(zhì)量(莖+葉+穗)和穗干物質(zhì)量均低于對照CK,尤以重旱重澇處理降幅最大,由此可知,早、晚稻前期重旱后期急轉(zhuǎn)重澇的處理對水稻成熟期穗部干物質(zhì)量及總干物質(zhì)量的影響最大,重旱重澇急轉(zhuǎn)表現(xiàn)為疊加損傷效應(yīng)。
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Effects of Drought-floods Abrupt Alternation During Panicle Differentiation Stage on Matter Accumulation and Yield Formation in Double-season Super Hybrid Rice
XIONG Qiang-qiang, ZHONG Lei, SHEN Tian-hua, CHEN Xiao-rong, ZHU Chang-lan, PENG Xiao-song, FU Jun-ru, HE Hao-hua
(College of Agronomy, Jiangxi Agricultural University/Key Laboratory of Crop Physiology, Ecology and Genetic Breeding, Ministry of Education/Research Center of Super Rice Engineering and Technology, Jiangxi Province/Collaboration Center for Double-season Rice Modernization Production, Nanchang 330045, China)
In this study, pot experiments were performed and super hybrid early rice(Ganxin 203) and late rice(Wufengyou T025) were selected as the material. During panicle differentiation initiation stage no drought in earlier stage and no floods in later stage(control, CK), mild drought in earlier stage and no floods in later stage(T1), severe drought in earlier stage and no floods in later stage(T2), no drought in earlier stage and mild floods in later stage(T3), no drought in earlier stage and severe floods in later stage(T4), severe drought in earlier stage and mild floods in later stage abrupt alternation(T5), severe drought in earlier stage and severe floods in later stage abrupt alternation(T6), mild drought in earlier stage and mild floods in later stage abrupt alternation(T7), mild drought in earlier stage and severe floods in later stage abrupt alternation(T8) were set. While the indices of the yield and its components, dry matter accumulation and distribution and net photosynthetic rate(Pn) were investigated. The results showed that, (1) the yield per plant in super hybrid early and late rice was decreased in different degrees under “drought”, “flood” and “drought-floods abrupt alternation”. Compared with CK, the early rice T1, T2, T3, T4, T5, T6, T7 and T8 yield per plant were decreased by 8.89%, 21.42%, 8.33%, 12.04%, 19.65%, 31.23%, 15.39% and 17.29%, respectively. The late rice were decreased by 6.21%, 15.06%, 7.77%, 11.53%, 17.40%, 20.85%, 13.46% and 11.70%, respectively. The yield of T2 and T6 treatments in early and late rice was decreased more, especially the yield of T6 treatment was largest decline, it was shown that only mild drought or mild floods on yield were less affected, but only severe drought or severe floods on yield were relatively large affected, and severe drought was greater affected than severe floods on yield. As for rice plants treated with severe drought in earlier stage and severe floods in later stage abrupt alternation, a certain degree of superimposition and reduction effect was existed. In the yield components, the main reasons for the yield decreased by effective panicles per plant, number of grains per panicle, seed setting rate and secondary branch number. The effects of yield and yield components were roughly the same among treatments. (2) Compare with CK, stem and leaf dry mass were decreased in different degrees in early and late rice under “drought”, “flood” and “drought-floods abrupt alternation”. Among the early and late rice at mature stage, the panicle dry mass of T6 treatment decreased by 33.40% and 19.88%, respectively, compared with CK, and the panicle shape becomes smaller. (3) The leaf chlorophyll content(SPAD value) in early and late rice was decreased under “drought”, “flood” and “drought-floods abrupt alternation”. SPAD value of T6 treatment in early and late rice were decreased remarkably, and with CK has a significant difference. (4) Leaf Pn of T6 treatment in early and late rice was the lowest, decreased by 37.57% and 10.17%, respectively, compared with CK. The results of the study were illustrated that severe drought in earlier stage and severe floods in later stage abrupt alternation had the maximum influence on panicle dry mass and total dry mass at mature stage in early and late rice, and severe drought in earlier stage and severe floods in later stage abrupt alternation showed a superposition damage effect.
Double-season super hybrid rice; Panicle differentiation stage; Drought-floods abrupt alternation; Yield; dry matter; Photosynthetic rate
2017-05-15
國家自然科學(xué)基金資助項(xiàng)目(314771441;30860136);江西省科技支撐項(xiàng)目(2010BNA03600);江西省教育廳項(xiàng)目(GJJ14283)
熊強(qiáng)強(qiáng)(1993-),博士生,主要從事水稻生理與遺傳育種研究。E-mail:xqq_qiang@163.com
10.3969/j.issn.1000-6362.2017.09.007
**通訊作者。E-mail:ccxxrr80@163.com
熊強(qiáng)強(qiáng),鐘蕾,沈天花,等.穗分化期旱澇急轉(zhuǎn)對雙季超級雜交稻物質(zhì)積累和產(chǎn)量形成的影響[J].中國農(nóng)業(yè)氣象,2017,38(9):597-608