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      蔬菜徒長(zhǎng)苗的形態(tài)及生理特征研究進(jìn)展

      2017-01-21 09:54:20王紅飛尚慶茂農(nóng)業(yè)部園藝作物生物學(xué)與種質(zhì)創(chuàng)制重點(diǎn)實(shí)驗(yàn)室中國(guó)農(nóng)業(yè)科學(xué)院蔬菜花卉研究所環(huán)渤海灣地區(qū)設(shè)施蔬菜優(yōu)質(zhì)高效生產(chǎn)協(xié)同創(chuàng)新中心北京100081
      中國(guó)蔬菜 2017年7期
      關(guān)鍵詞:胚軸徒長(zhǎng)幼苗

      王紅飛 尚慶茂(農(nóng)業(yè)部園藝作物生物學(xué)與種質(zhì)創(chuàng)制重點(diǎn)實(shí)驗(yàn)室,中國(guó)農(nóng)業(yè)科學(xué)院蔬菜花卉研究所,環(huán)渤海灣地區(qū)設(shè)施蔬菜優(yōu)質(zhì)高效生產(chǎn)協(xié)同創(chuàng)新中心,北京100081)

      蔬菜徒長(zhǎng)苗的形態(tài)及生理特征研究進(jìn)展

      王紅飛 尚慶茂*
      (農(nóng)業(yè)部園藝作物生物學(xué)與種質(zhì)創(chuàng)制重點(diǎn)實(shí)驗(yàn)室,中國(guó)農(nóng)業(yè)科學(xué)院蔬菜花卉研究所,環(huán)渤海灣地區(qū)設(shè)施蔬菜優(yōu)質(zhì)高效生產(chǎn)協(xié)同創(chuàng)新中心,北京100081)

      蔬菜集約化育苗以多孔連體式穴盤(pán)為容器,密度高,根系發(fā)育空間小,基質(zhì)緩沖能力弱,幼苗極易徒長(zhǎng)。徒長(zhǎng)苗形態(tài)及生理代謝迥異于正常苗,表現(xiàn)為下胚軸及節(jié)間顯著伸長(zhǎng),葉片開(kāi)展度增大、葉片變薄,根冠比降低,組織含水量提高,對(duì)生物和非生物逆境的適應(yīng)性減弱等。本文就蔬菜徒長(zhǎng)苗的形態(tài)特征、開(kāi)花習(xí)性、抗性表達(dá)、生理代謝及相關(guān)防控技術(shù)等進(jìn)行了簡(jiǎn)要綜述,旨在為蔬菜徒長(zhǎng)苗的防控及相關(guān)技術(shù)的開(kāi)發(fā)提供參考。

      徒長(zhǎng)苗;形態(tài)特征;開(kāi)花習(xí)性;生理特征;環(huán)境因子

      蔬菜集約化育苗以多孔連體式穴盤(pán)為容器,以人工混配輕型基質(zhì)替代土壤,具有節(jié)種、節(jié)能、省工,適于規(guī)?;?biāo)準(zhǔn)化生產(chǎn),便于機(jī)械化操作等優(yōu)點(diǎn),但在實(shí)際生產(chǎn)過(guò)程中存在幼苗密度大、植株間相互蔭蔽等問(wèn)題,容易形成徒長(zhǎng)苗。此外,為滿足周年生產(chǎn)需要,蔬菜集約化育苗常在氣候不適宜的季節(jié)進(jìn)行(尚慶茂,2011),夏季的陰雨高溫天氣、冬季的霧霾天氣頻繁出現(xiàn),這種弱光、高溫、高濕條件常會(huì)導(dǎo)致幼苗徒長(zhǎng),不利于壯苗的育成和優(yōu)質(zhì)豐產(chǎn)栽培(甘小虎 等,2012 a,2012 b)。

      幼苗徒長(zhǎng)已成為當(dāng)前的研究熱點(diǎn),國(guó)內(nèi)外眾多學(xué)者對(duì)幼苗徒長(zhǎng)機(jī)制進(jìn)行了研究,認(rèn)為植物激素在幼苗徒長(zhǎng)過(guò)程中發(fā)揮了關(guān)鍵作用。IAA、GA是較早發(fā)現(xiàn)的兩類(lèi)植物激素,植物體內(nèi)IAA、GA含量升高,幼苗株高顯著增加,除株高外的其他表型特征及幼苗體內(nèi)生理代謝等亦發(fā)生變化(Gray et al., 1998;Koini et al.,2009;Sun et al.,2012;Procko et al.,2014;Song et al.,2015)。本文就蔬菜徒長(zhǎng)苗的形態(tài)特征、開(kāi)花習(xí)性、生理特征、環(huán)境因子與徒長(zhǎng)苗的關(guān)系及幼苗徒長(zhǎng)防控技術(shù)進(jìn)行綜述,旨在為蔬菜徒長(zhǎng)苗的制御及相關(guān)技術(shù)的開(kāi)發(fā)提供參考。

      1 蔬菜徒長(zhǎng)苗的形態(tài)特征

      1.1 表型特征

      徒長(zhǎng)是指蔬菜幼苗因生長(zhǎng)條件不協(xié)調(diào)而產(chǎn)生的莖葉發(fā)育過(guò)旺的現(xiàn)象(單永輝,2006)。徒長(zhǎng)苗主要是由于縱向生長(zhǎng)加速引起的(梁鎮(zhèn)林,2000),株高顯著高于正常幼苗。徒長(zhǎng)苗的株高變化主要是由下胚軸長(zhǎng)度變化引起的(周學(xué)超,2010),下胚軸中段的長(zhǎng)度顯著高于上段和下段(Xiao et al.,2014,2016)。此外,徒長(zhǎng)苗的莖粗降低,株高/莖粗的比值升高(侯興亮 等,2002;王學(xué)文 等,2009;武曉玲 等,2014);徒長(zhǎng)苗的葉片顏色變淡,比葉面積增加,葉片厚度顯著降低(侯興亮 等,2002;王學(xué)文 等,2009;周學(xué)超,2010);葉柄變長(zhǎng),促使葉片靠上生長(zhǎng),葉柄與胚軸夾角變大,呈現(xiàn)出偏下生長(zhǎng)的特性(Zanten et al.,2009;Delker et al.,2014;Fankhauser & Batschauer,2016)。此外,徒長(zhǎng)苗的根系吸收能力也受到影響,根系長(zhǎng)度、側(cè)根數(shù)量顯著降低,根冠比、壯苗指數(shù)分別降至對(duì)照的66.67%和48.52%(武曉玲 等,2014;Prockoet al.,2014)。

      1.2 細(xì)胞及亞細(xì)胞結(jié)構(gòu)特征

      徒長(zhǎng)苗根莖葉等器官的形態(tài)變化,主要是由這些器官的組成細(xì)胞變化引起的,徒長(zhǎng)苗的下胚軸、葉片在細(xì)胞水平和亞細(xì)胞水平均發(fā)生顯著變化。徒長(zhǎng)苗的下胚軸細(xì)胞長(zhǎng)度顯著高于正常苗(Gendreau et al.,1997),且下胚軸中段細(xì)胞長(zhǎng)度顯著高于上段和下段(Xiao et al.,2016)。徒長(zhǎng)苗的下胚軸細(xì)胞呈矩形,表皮細(xì)胞表面較為平滑,細(xì)胞排列較為疏松,而正常幼苗的下胚軸細(xì)胞則為橢圓形,表皮細(xì)胞較小,呈多邊形,沿縱軸緊密排列(Qin et al.,2012)。徒長(zhǎng)苗葉片柵欄組織和海綿組織厚度均有所降低,但柵欄組織降低幅度大,細(xì)胞變短,排列疏松,而正常苗柵欄組織細(xì)胞排列整齊,呈長(zhǎng)柱形(王學(xué)文 等,2009;周學(xué)超,2010)。在亞細(xì)胞結(jié)構(gòu)水平,徒長(zhǎng)苗下胚軸細(xì)胞中的葉綠體變長(zhǎng),基粒和基質(zhì)片層數(shù)量減少,線粒體數(shù)量增多,淀粉粒消失(胡宏敏,2012);葉肉細(xì)胞液泡內(nèi)含物增多,葉綠體在細(xì)胞內(nèi)呈不規(guī)則分布,線粒體內(nèi)含物減少,部分線粒體出現(xiàn)膜破裂、空泡化和解體的現(xiàn)象(周學(xué)超,2010)。

      2 蔬菜徒長(zhǎng)苗的開(kāi)花習(xí)性

      蔬菜徒長(zhǎng)苗營(yíng)養(yǎng)生長(zhǎng)旺盛,消耗了大量營(yíng)養(yǎng)物質(zhì),導(dǎo)致組織C/N比例發(fā)生變化,進(jìn)而顯著影響開(kāi)花結(jié)實(shí)進(jìn)程(Casal,2013)。前人研究表明,徒長(zhǎng)苗的開(kāi)花時(shí)間提前,且開(kāi)花時(shí)間與抽生葉片數(shù)呈顯著正相關(guān)(Botto & Smith,2002;Blázquez et al.,2003;Balasubramanian et al.,2006)。一般情況下,擬南芥(Col)正常苗于第17片蓮座葉片展開(kāi)時(shí)進(jìn)行抽薹開(kāi)花,而徒長(zhǎng)苗提前至第10片蓮座葉片展開(kāi)時(shí)就抽薹開(kāi)花了(Koini et al.,2009)。黃瓜徒長(zhǎng)苗開(kāi)花習(xí)性也發(fā)生變化,第1雌花節(jié)位顯著升高,開(kāi)花時(shí)間較對(duì)照晚2~5 d(明村豪 等,2011)。擬南芥徒長(zhǎng)苗與黃瓜徒長(zhǎng)苗在開(kāi)花習(xí)性方面的差異,可能是由于物種、生長(zhǎng)條件不同造成的。

      3 蔬菜徒長(zhǎng)苗的生理特征

      3.1 水分代謝

      蔬菜徒長(zhǎng)苗的根系長(zhǎng)度、側(cè)根數(shù)量下降,根系活力受到抑制,由0.4 mg·g-1·h-1降低至0.2 mg·g-1·h-1,導(dǎo)致植物水分代謝及以水分代謝為載體的礦質(zhì)營(yíng)養(yǎng)代謝能力降低(周艷虹 等,2004;周學(xué)超,2010)。與正常苗相比,徒長(zhǎng)苗氣孔開(kāi)度降低,導(dǎo)致蒸騰速率下降,水分流失受到抑制,組織含水量增加,為正常苗的103.88%(侯興亮 等,2002;王學(xué)文 等,2009)。

      3.2 光合代謝

      徒長(zhǎng)苗的葉綠素和類(lèi)胡蘿卜素含量顯著下降,捕光能力降低,細(xì)胞色素質(zhì)琨(PQ)、細(xì)胞色素蛋白(Cyt)和鐵氧還素(Fd)等電子傳遞鏈組分含量顯著下降,光反應(yīng)受到抑制;此外,Rubisco羧化酶活性降低,導(dǎo)致葉肉細(xì)胞對(duì)CO2的利用能力下降,暗反應(yīng)受影響,碳水化合物合成量降低,植株干物質(zhì)積累量減少(周艷虹 等,2004;王祥寧 等,2007;明村豪 等,2011)。

      3.3 抗逆代謝

      徒長(zhǎng)苗葉面積、下胚軸長(zhǎng)度顯著增加,細(xì)胞膨大,細(xì)胞壁厚度降低,導(dǎo)致植物組織韌性降低,對(duì)生物和非生物逆境的適應(yīng)能力降低(Derbyshire et al.,2007;Irshad et al.,2008)。

      3.3.1 非生物脅迫 蔬菜徒長(zhǎng)苗可溶性固形物、可溶性糖、可溶性蛋白含量降低,滲透調(diào)節(jié)能力下降,不利于植物抵御高滲、低溫等非生物脅迫(徐磊 等,2009;明村豪 等,2011)。此外,徒長(zhǎng)苗體內(nèi)花青素、芥子油甙、酚類(lèi)物質(zhì)等次級(jí)代謝產(chǎn)物顯著降低(Izaguirre et al.,2006;Moreno et al.,2009;Xie et al.,2016)。花青素可以釋放H+,清除氧自由基,避免光抑制、膜脂過(guò)氧化等現(xiàn)象出現(xiàn),從而保護(hù)膜系統(tǒng)的完整性,提高對(duì)高溫、低溫、干旱、強(qiáng)光等非生物脅迫的適應(yīng)能力(Lorenc-Kuku?a et al.,2005;Albert et al.,2009)。

      3.3.2 生物脅迫 徒長(zhǎng)苗葉片及胚軸表面的凸起、絨毛、刺瘤等保護(hù)結(jié)構(gòu)密度降低。番茄徒長(zhǎng)苗絨毛數(shù)量減少,導(dǎo)致大量螨類(lèi)害蟲(chóng)在植株上繁殖并刺吸植物莖葉,受害葉片前期產(chǎn)生黃褐斑,后期失綠脫落(Nihoul,1993;Roberts & Paul,2006)。

      番茄灰霉菌和葡萄孢屬真菌等病原菌侵染植物組織后,能激發(fā)活性氧產(chǎn)生,而活性氧清除能力較差的徒長(zhǎng)苗受到致病菌青睞,發(fā)病率顯著高于正常苗(Iriti et al.,2004;Segmüller et al.,2008;Zhang et al.,2013),黃瓜徒長(zhǎng)苗的白粉病發(fā)病率約為正常苗的2.5倍(Wang et al.,2010)。芥子油甙和酚類(lèi)物質(zhì)具有揮發(fā)性,一方面能趨避有害昆蟲(chóng),另一方面能吸引有害昆蟲(chóng)的捕食者和寄生性天敵,以防蟲(chóng)害發(fā)生(Roberts & Paul,2006;Xia et al.,2009;Wang & Wu,2013)。番茄徒長(zhǎng)苗中綠原酸、蕓香苷等酚類(lèi)物質(zhì)含量降低,導(dǎo)致植株受毛毛蟲(chóng)啃食嚴(yán)重(Jansen & Stamp,1997)。

      徒長(zhǎng)苗體內(nèi)生長(zhǎng)素(IAA)、赤霉素(GA)、茉莉酸(JA)、水楊酸(SA)等含量變化顯著(Feng et al.,2008;Ballaré,2011;Agrawal et al.,2012;Fu et al.,2012),與細(xì)胞膨大相關(guān)的IAA和GA含量顯著增加,與植物抗逆能力相關(guān)的JA、SA含量降低。GA能誘導(dǎo)JA信號(hào)傳導(dǎo)關(guān)鍵因子DELLA蛋白泛素化降解,導(dǎo)致JA信號(hào)轉(zhuǎn)導(dǎo)中斷,JA能促使PR蛋白的合成量提高,并介導(dǎo)受病原菌侵染和有害昆蟲(chóng)啃食細(xì)胞的程序性死亡,從而提高植株抵御生物脅迫的能力(Hou et al.,2010;Ballaré et al.,2012;Campos et al.,2016)。

      4 環(huán)境因子與蔬菜幼苗徒長(zhǎng)

      4.1 單一環(huán)境因子

      光照強(qiáng)度與幼苗徒長(zhǎng)程度呈負(fù)相關(guān),光照越弱,徒長(zhǎng)越嚴(yán)重。當(dāng)光照強(qiáng)度降低至對(duì)照(1 100~1 300 μmol·m-2·s-1)的60%~70%時(shí),黃瓜幼苗便開(kāi)始徒長(zhǎng),徒長(zhǎng)苗高度達(dá)到對(duì)照的1.10倍;光照強(qiáng)度繼續(xù)降至對(duì)照的30%~40%時(shí),徒長(zhǎng)苗高度為對(duì)照的1.23倍(明村豪 等,2011)。紅光與遠(yuǎn)紅光對(duì)幼苗徒長(zhǎng)的調(diào)控作用相反,紅光抑制,遠(yuǎn)紅光促進(jìn),在R∶FR比值較低的條件下,幼苗徒長(zhǎng),下胚軸長(zhǎng)度超出對(duì)照1倍(Hersch et al.,2014)。晝夜節(jié)律作為重要的光信號(hào)因子,對(duì)植物的生長(zhǎng)發(fā)育起著重要調(diào)控作用,徒長(zhǎng)苗下胚軸長(zhǎng)度隨黑暗時(shí)間延長(zhǎng)而增加,極夜條件下徒長(zhǎng)苗下胚軸長(zhǎng)度為極晝條件下的6倍(Niwa et al.,2009)。

      作為影響植物生長(zhǎng)發(fā)育的另一重要環(huán)境因子,溫度在一定范圍內(nèi)升高時(shí),能促進(jìn)植物下胚軸伸長(zhǎng),但當(dāng)溫度超過(guò)植物適宜生長(zhǎng)范圍后,反而抑制下胚軸伸長(zhǎng)(Dafny-Yelin et al.,2008;Sun et al.,2012)。 在溫光適宜的范圍內(nèi),水分充足有利于壯苗的培育,但水分含量過(guò)高時(shí),容易造成幼苗徒長(zhǎng);水分含量過(guò)低,容易形成老化苗(王娟 等,2002)。

      4.2 復(fù)合環(huán)境因子

      溫、光、水作為植物賴以生存的環(huán)境條件,既能單獨(dú)發(fā)揮作用,又能協(xié)同調(diào)控植物生長(zhǎng)(Franklin,2009)。在低溫條件下,光照強(qiáng)度在0~100 μmol·m-2·s-1范圍內(nèi)增加時(shí),能顯著抑制幼苗下胚軸伸長(zhǎng);但當(dāng)溫度升高至27 ℃,光照強(qiáng)度在0~1 μmol·m-2·s-1范圍內(nèi)變化時(shí),則抑制下胚軸伸長(zhǎng),但光照強(qiáng)度在1~100 μmol· m-2·s-1范圍內(nèi)變化時(shí),能促進(jìn)下胚軸伸長(zhǎng)(Johansson et al.,2014)。研究表明,基質(zhì)含水量過(guò)高,能引起蔬菜幼苗徒長(zhǎng),在弱光條件下徒長(zhǎng)尤為嚴(yán)重(毛煒光 等,2007;徐磊 等,2009)。在調(diào)控幼苗生長(zhǎng)的過(guò)程中,溫度和水分互作顯著,二者協(xié)同作用條件下,幼苗生長(zhǎng)量增加,徒長(zhǎng)明顯(Vile et al.,2012)。

      5 蔬菜幼苗的徒長(zhǎng)防控技術(shù)

      近年來(lái),國(guó)內(nèi)外眾多學(xué)者對(duì)蔬菜幼苗徒長(zhǎng)機(jī)理進(jìn)行了研究,并取得了長(zhǎng)足進(jìn)展,研究結(jié)果表明,光敏色素互作蛋白PIFs在幼苗徒長(zhǎng)過(guò)程中發(fā)揮關(guān)鍵作用(Casal,2013;Procko et al.,2014)。PIFs是一類(lèi)bHLH轉(zhuǎn)錄因子,通過(guò)促進(jìn)植物激素合成,引起幼苗徒長(zhǎng)(Tao et al.,2008;Stavang et al.,2009;Franklin et al.,2011;Delker et al.,2014)。隨著下胚軸伸長(zhǎng)機(jī)理逐漸明確,生產(chǎn)管理人員加強(qiáng)了蔬菜生長(zhǎng)敏感期的環(huán)境管理,并針對(duì)調(diào)控下胚軸伸長(zhǎng)的關(guān)鍵位點(diǎn),研發(fā)了相應(yīng)的延緩型植物生長(zhǎng)調(diào)節(jié)劑,對(duì)幼苗徒長(zhǎng)防控效果明顯(Heins et al.,2000;宮萬(wàn)祥和丁克友,2007)。此外,許多行業(yè)專(zhuān)家試圖通過(guò)機(jī)械刺激防控蔬菜幼苗徒長(zhǎng),以替代植物生長(zhǎng)調(diào)節(jié)劑在蔬菜生產(chǎn)中的地位,推動(dòng)綠色、有機(jī)蔬菜的生產(chǎn)(Garner & Bjorkman,1996)。

      5.1 環(huán)境管理

      蔬菜幼苗生長(zhǎng)早期對(duì)環(huán)境條件敏感,外界的弱光、高溫及高濕等環(huán)境條件極易引發(fā)幼苗的徒長(zhǎng),此時(shí)要加強(qiáng)對(duì)環(huán)境的管理。

      光照強(qiáng)度是影響幼苗生長(zhǎng)的重要因素,連續(xù)的陰雨及霧霾等障礙天氣的出現(xiàn),常會(huì)導(dǎo)致幼苗徒長(zhǎng)。生產(chǎn)過(guò)程中,一般采用透光率較好的無(wú)滴膜及增加后墻反光幕等方法,增加光照強(qiáng)度,防止幼苗徒長(zhǎng)(楊艷春,2013)。此外,在溫室內(nèi)增加人工光源,可有效抑制下胚軸伸長(zhǎng),促進(jìn)植株根系生長(zhǎng),提高幼苗的抗逆性(祝聰宇 等,2017)。

      植物對(duì)溫度變化敏感,溫度稍高便可引起幼苗徒長(zhǎng),栽培管理過(guò)程中,要注意及時(shí)放風(fēng),以降低室內(nèi)溫度,夜間可適當(dāng)升溫,減小晝夜溫差,可有效抑制幼苗徒長(zhǎng)(Berghage,1998)。

      秋冬季光照相對(duì)較弱,要嚴(yán)格把控澆水量和澆水時(shí)間,定植前澆透水,定植后和緩苗期澆小水(楊艷春,2013),以防止幼苗徒長(zhǎng)。

      5.2 植物生長(zhǎng)調(diào)節(jié)劑

      利用植物生長(zhǎng)調(diào)節(jié)劑防控幼苗徒長(zhǎng)是一種簡(jiǎn)單而有效的方法,目前已在蔬菜幼苗培育過(guò)程中得到了廣泛應(yīng)用(王娟,2006)。生產(chǎn)上常用的植物生長(zhǎng)調(diào)節(jié)劑主要包括矮壯素(chlorocholine chloride,CCC)、烯效唑(U-niconazole,S3307)、多效唑(paclobutrazol,PP333)、比久(B9)等,處理方法主要包括浸種、種子包衣、葉面噴施、土壤或基質(zhì)混合處理等(Berova & Zlatev,2000;黃少華 等,2006;張靜 等,2007;劉東冉 等,2008),其主要通過(guò)抑制內(nèi)源生長(zhǎng)素(IAA)和赤霉素(GA)的合成,促進(jìn)IAA和GA的降解,抑制幼苗徒長(zhǎng)。

      5.3 機(jī)械調(diào)節(jié)

      機(jī)械刺激對(duì)幼苗生長(zhǎng)具有重要調(diào)控作用,可降低部分蔬菜作物莖及葉柄的伸長(zhǎng),從而抑制幼苗徒長(zhǎng)。目前常用的機(jī)械刺激包括:接觸刺激(brushing)、阻抑或阻壓(impedance)及振蕩處理等。

      接觸刺激是一種有效的機(jī)械調(diào)節(jié)方式,少量刺激便可有效抑制幼苗徒長(zhǎng),頻率過(guò)高可誘發(fā)機(jī)械傷害。前人研究表明,接觸刺激可有效降低番茄、黃瓜等蔬菜幼苗的植株高度,一天中的處理時(shí)間和頻率對(duì)株高沒(méi)有顯著影響,但處理天數(shù)與之顯著相關(guān),連續(xù)的接觸刺激處理可使番茄幼苗株高降低20%(Latimer,1991;Garner & Bjorkman,1996)。

      阻抑或阻壓主要是通過(guò)丙烯醇薄片、聚酯薄膜、纖維玻璃等材料阻壓幼苗頂部,防止幼苗徒長(zhǎng),但這種處理方式會(huì)導(dǎo)致幼苗莖彎曲,不利于機(jī)械化移栽(Piszczek & Jerzy,1987)。

      振蕩處理可有效控制番茄幼苗的生長(zhǎng),每天連續(xù)多次振蕩處理較連續(xù)振蕩處理效果更好,植株高度降低更多,但整體的處理效果不如接觸刺激(Latimer & Mitchell,1988)。

      6 展望

      幼苗徒長(zhǎng)是蔬菜集約化育苗過(guò)程中的常見(jiàn)問(wèn)題,目前防控上主要依賴植物生長(zhǎng)調(diào)節(jié)劑,但在育苗過(guò)程中生長(zhǎng)調(diào)節(jié)劑用量不容易把控,濃度過(guò)高易產(chǎn)生藥害,濃度過(guò)低則達(dá)不到防控效果,亟需研究與開(kāi)發(fā)新的徒長(zhǎng)防控技術(shù)及實(shí)用產(chǎn)品。為此,今后研究應(yīng)著重分析不同環(huán)境信號(hào)對(duì)幼苗徒長(zhǎng)的交互作用;深入了解細(xì)胞膨大與分裂調(diào)控機(jī)制及其在下胚軸伸長(zhǎng)過(guò)程中的作用;利用轉(zhuǎn)錄組學(xué)、蛋白質(zhì)組學(xué)及代謝組學(xué)相結(jié)合的方法,探明環(huán)境因子調(diào)控下胚軸伸長(zhǎng)的信號(hào)通路,分析不同環(huán)境信號(hào)的交叉調(diào)控位點(diǎn)及其生物學(xué)功能,這將是全面揭示幼苗徒長(zhǎng)機(jī)制及廣譜防控技術(shù)開(kāi)發(fā)的關(guān)鍵。

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      Research Progress on Morphological and Physiological Characteristics of Vegetable Leggy Seedlings

      WANG Hong-fei,SHANG Qing-mao*
      (Key Laboratory of Biology and Genetic Improvement of Horticultural Crops,Institute of Vegetables and Flowers,Chinese Academy of Agricultural Sciences,Collaborative Innovation Center of Protected Vegetable Surround Bohai Gulf Region,Beijing 100081,China)

      Intensive vegetable seedlings culture usually took with porous conjoined plug as container,which had high density,less space for root system development and weak buffer capacity of rhizosphere. It was very easy for seedlings to over grown. The morphological and physiological metabolism of leggy seedlings were different from the normal ones,including elongated hypocotyl and internode,enlarged leaf angle,reduced leaf thickness and root to shoot ratio,increased moisture content in tissue,weakened adaptability to biotic and abiotic stress. This review summarized the morphological characteristics of vegetable leggy seedlings,flowering habit,resistance expression,physiological metabolism and related preventing techniques,etc.,aiming at providing reference for preventing leggy seedlings and developing related technology.

      Leggy seedling;Morphological characteristic;Flowering habit;Physiological metabolism;Environmental factor

      王紅飛,女,博士研究生,專(zhuān)業(yè)方向:蔬菜種苗發(fā)育調(diào)控與繁育技術(shù),E-mail:wanghongfei0329@163.com

      *通訊作者(Corresponding author),尚慶茂,男,研究員,博士生導(dǎo)師,專(zhuān)業(yè)方向:蔬菜栽培生理及分子生物學(xué),E-mail:shangqingmao@ caas.cn

      2017-03-21;接受日期:2017-05-26

      國(guó)家自然科學(xué)基金項(xiàng)目(31172001),國(guó)家現(xiàn)代農(nóng)業(yè)產(chǎn)業(yè)技術(shù)體系建設(shè)專(zhuān)項(xiàng)(CARS-25),公益性行業(yè)(農(nóng)業(yè))科研專(zhuān)項(xiàng)(201303014),中國(guó)農(nóng)業(yè)科學(xué)院科技創(chuàng)新工程項(xiàng)目(CAAS-ASTIPIVFCAAS)

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