冬小麥抗倒春寒性能鑒定方法和指標(biāo)的比較*

張樂樂，陳 翔，柯媛媛，柳彬彬，Muhammad Ahmad Hassan，張 妍，許 輝，李金才,2**

冬小麥抗倒春寒性能鑒定方法和指標(biāo)的比較*

張樂樂1，陳 翔1，柯媛媛1，柳彬彬1，Muhammad Ahmad Hassan1，張 妍1，許 輝1，李金才1,2**

（1.安徽農(nóng)業(yè)大學(xué)農(nóng)學(xué)院/農(nóng)業(yè)農(nóng)村部華東地區(qū)作物栽培科學(xué)觀測(cè)站，合肥 230036；2.江蘇省現(xiàn)代作物生產(chǎn)協(xié)同創(chuàng)新中心，南京 210095）

倒春寒是限制黃淮冬麥區(qū)小麥穩(wěn)產(chǎn)和豐產(chǎn)的主要因素之一，選育抗倒春寒小麥新品種是降低倒春寒危害的重要手段，建立簡(jiǎn)單易行、易推廣且全面系統(tǒng)的小麥抗倒春寒性能鑒定方法和指標(biāo)體系對(duì)保障糧食安全有著重要意義。本文系統(tǒng)總結(jié)了冬小麥抗倒春寒鑒定的方法和指標(biāo)，進(jìn)行了比較和評(píng)價(jià)，并提出鑒定原則和程序。（1）鑒定方法方面，主要包括直接鑒定法、間接鑒定法和綜合鑒定法等，直接鑒定法有田間自然鑒定法（引種栽培應(yīng)用最廣，但周期長(zhǎng)、工作量大）、人工模擬氣候室法（周期短、可重復(fù)性強(qiáng)，但對(duì)設(shè)備與技術(shù)要求較高）和生長(zhǎng)恢復(fù)法（冬前凍害鑒定較多，抗倒春寒鑒定方面較少，一般與其它鑒定方法配合使用）等；間接鑒定法有助倒春寒危害溫度閾值的研究，但目前指標(biāo)與抗寒性關(guān)系還未有明確定論，應(yīng)用受到限制；綜合鑒定法鑒定結(jié)果全面而準(zhǔn)確，但對(duì)技術(shù)、成本要求較高且工作量較大。因此，在鑒定過程中要結(jié)合實(shí)際情況合理搭配、綜合運(yùn)用。（2）鑒定指標(biāo)方面，主要包括形態(tài)與農(nóng)藝、生理生化、氣象與生態(tài)指標(biāo)等，形態(tài)與農(nóng)藝指標(biāo)一般用于田間自然鑒定法，常以數(shù)量、質(zhì)量、顏色和形態(tài)變化等作為參考項(xiàng)；生理生化指標(biāo)一般用于間接鑒定法，包括活性氧、滲透調(diào)節(jié)物質(zhì)、內(nèi)源激素含量、抗氧化酶活性以及光合與呼吸特性等；氣象與生態(tài)指標(biāo)常應(yīng)用于各種鑒定方法，氣溫、地溫、冠層溫度、半致死溫度和恢復(fù)生長(zhǎng)最低溫度常作為鑒定指標(biāo)，而光照、CO2、濕度和肥力等因素會(huì)影響結(jié)果準(zhǔn)確性。（3）關(guān)于鑒定原則與程序，本文提出了鑒定結(jié)果的地域性、指標(biāo)選用的針對(duì)性、鑒定方法與指標(biāo)的綜合性和實(shí)用性等鑒定原則，并以易災(zāi)期、品種敏感期、敏感器官、鑒定方法和指標(biāo)、低溫程度、田間/盆栽試驗(yàn)、數(shù)學(xué)分析方法、抗寒性能評(píng)價(jià)與分級(jí)為主要流程的鑒定程序。最后，從抗寒生理研究、新興技術(shù)鑒定研究與應(yīng)用和鑒定產(chǎn)品的研發(fā)等方面做了展望。

冬小麥；倒春寒；鑒定；指標(biāo)；方法

小麥（L）是世界三大糧食作物之一，全球約60%的人口以小麥為主食[1]。2050年世界人口預(yù)計(jì)達(dá)到97.3億，小麥產(chǎn)量年均增長(zhǎng)率要達(dá)到2.4%才能滿足全球人口增長(zhǎng)對(duì)糧食的需求，但目前全球小麥產(chǎn)量年均增長(zhǎng)率僅為0.9%[2?3]。同時(shí)隨著全球氣候變暖，極端低溫災(zāi)害事件頻發(fā)重發(fā)，嚴(yán)重影響了小麥的穩(wěn)產(chǎn)、豐產(chǎn)[4?5]。低溫災(zāi)害給中國(guó)、美國(guó)、澳大利亞以及部分歐洲國(guó)家的小麥生產(chǎn)造成了極大的損失[6?11]。據(jù)報(bào)道，1995?2010年美國(guó)堪薩斯州的低溫災(zāi)害事件發(fā)生次數(shù)多達(dá)41次，導(dǎo)致小麥減產(chǎn)超過538kg·hm?2[6]。澳大利亞昆士蘭和新南威爾士北部每年因低溫災(zāi)害導(dǎo)致小麥減產(chǎn)10%[7]，年均經(jīng)濟(jì)損失達(dá)1億澳元[8]。1980?2020年中國(guó)小麥主產(chǎn)區(qū)發(fā)生近20次大規(guī)模低溫災(zāi)害事件，其中尤以黃淮冬麥區(qū)的倒春寒（Late Spring Coldness，LSC）危害最為嚴(yán)重，該地區(qū)發(fā)生頻率可達(dá)40%，在2009、2013、2015、2018和2020年均有發(fā)生[9?11]。因此，倒春寒已成為限制中國(guó)小麥穩(wěn)產(chǎn)豐產(chǎn)與優(yōu)質(zhì)的主要農(nóng)業(yè)氣象災(zāi)害。

小麥倒春寒災(zāi)害主要指在拔節(jié)?孕穗期遭遇突然降溫天氣，造成幼穗受傷或死亡，部分小穗不結(jié)實(shí)甚至全穗不結(jié)實(shí)，從而導(dǎo)致小麥減產(chǎn)的一種農(nóng)業(yè)氣象災(zāi)害[12]。前人從光合作用[13]、呼吸作用[14]、活性氧代謝[15]、滲透調(diào)節(jié)[16]、內(nèi)源激素代謝[17]以及蛋白質(zhì)組學(xué)[18]和基因標(biāo)記技術(shù)[19]等方面對(duì)倒春寒危害機(jī)理[20]進(jìn)行了大量研究，但其致災(zāi)機(jī)理尚未明確。同時(shí)由于倒春寒的發(fā)生具有隨機(jī)性與不可預(yù)見性，且受害癥狀具有隱蔽性，從而導(dǎo)致災(zāi)前預(yù)防和災(zāi)后減災(zāi)措施難以及時(shí)到位，因此，小麥生產(chǎn)中應(yīng)對(duì)倒春寒的防災(zāi)減災(zāi)能力不強(qiáng)。選用抗倒春寒品種是小麥生產(chǎn)中降低倒春寒危害的重要手段，但由于缺少準(zhǔn)確有效的鑒定方法和指標(biāo)體系以及相關(guān)標(biāo)準(zhǔn)，使得抗倒春寒品種的選育面臨重大難題。因此，本研究通過總結(jié)和綜合評(píng)價(jià)前人研究的冬小麥品種抗倒春寒性能（Tolerance against Late Spring Coldness，TLSC）鑒定方法和指標(biāo)，提出相關(guān)鑒定原則和程序，并對(duì)未來的研究方向進(jìn)行展望，旨在為抗倒春寒小麥新品種選育和減災(zāi)保產(chǎn)提供理論參考。

1 冬小麥品種TLSC鑒定方法

1.1 鑒定方法分類

目前，冬小麥TLSC鑒定可以參考小麥品種抗寒性鑒定的一般方法，主要有三種，分別是直接鑒定法（DIM）、間接鑒定法（IIM）和綜合鑒定法（CIM），這些方法通常與各種數(shù)學(xué)分析方法（MAM）組合使用。各種方法的具體分類見圖1。

圖1 冬小麥品種TLSC鑒定方法分類

1.2 鑒定方法的優(yōu)劣勢(shì)分析

1.2.1 直接鑒定法

（1）田間自然鑒定法（FIM）

FIM是在倒春寒發(fā)生后對(duì)田間受害植株的器官或組織以一定的標(biāo)準(zhǔn)進(jìn)行評(píng)價(jià)和比較，根據(jù)作物受災(zāi)表現(xiàn)評(píng)價(jià)其抗寒性的一種直觀鑒定方法。該方法是引種栽培應(yīng)用最廣泛的一種鑒定方法，也是公認(rèn)最早、最有效的鑒定方法之一[21]。農(nóng)業(yè)農(nóng)村部標(biāo)準(zhǔn)《NY/T 1307?2007》[22]中提出的小麥抗寒性分級(jí)就是利用這一方法將冬麥區(qū)春季階段的小麥抗寒性分為5級(jí)，即1級(jí)（無凍害表征）、2級(jí)（葉尖受凍發(fā)黃）、3級(jí)（葉片凍死一半）、4級(jí)（葉片全枯）和5級(jí)（植株或大部分分蘗凍死）。喬玉強(qiáng)等[23]在倒春寒發(fā)生年份對(duì)同一品種進(jìn)行不同播期試驗(yàn)，調(diào)查了田間異型穗率、殘缺穗率和無籽穗率，結(jié)合其它相關(guān)指標(biāo)通過聚類分析得出同一品種在不同播期和不同地區(qū)的TLSC強(qiáng)弱表現(xiàn)均有差異，表明田間栽培管理措施和不同地區(qū)易災(zāi)期以及凍（冷）害敏感期等因素影響著品種的TLSC評(píng)價(jià)結(jié)果。FIM雖然能較好地鑒定冬小麥TLSC，但由于鑒定周期漫長(zhǎng)且工作量較大，一定程度上影響了抗倒春寒新品種選育和推廣效率。

（2）人工模擬氣候室法（ACCM）

ACCM是通過人工模擬大田低溫環(huán)境來進(jìn)行小麥抗寒性鑒定的一種常用方法。目前，利用該方法研究小麥抗寒生理機(jī)制和鑒定小麥抗寒性已成為主流。Liu等[24]利用ACCM在小麥拔節(jié)期和孕穗期進(jìn)行低溫脅迫模擬試驗(yàn)，發(fā)現(xiàn)低溫脅迫程度和持續(xù)時(shí)間顯著影響小麥的光合生產(chǎn)能力，進(jìn)而影響產(chǎn)量的形成。Zhang等[25]利用ACCM發(fā)現(xiàn)孕穗期低溫會(huì)誘導(dǎo)穗部脫落酸的積累，同時(shí)提升蔗糖合成酶的活性，進(jìn)而增加穗部蔗糖含量來提升抗寒能力，但會(huì)影響幼穗的正常發(fā)育。ACCM鑒定周期較FIM短，可重復(fù)性強(qiáng)，同時(shí)減少了環(huán)境與品種基因型互作的影響[26]。但該方法對(duì)設(shè)備要求較高，且對(duì)各種變量也有嚴(yán)格要求，尤其是對(duì)溫度參數(shù)的設(shè)置，需要對(duì)大量氣象數(shù)據(jù)進(jìn)行分析確定合理的降溫模式和水平。

（3）生長(zhǎng)恢復(fù)法（GRM）

GRM是通過觀察和測(cè)定小麥植株或組織受低溫脅迫后各項(xiàng)指標(biāo)的恢復(fù)水平來評(píng)判其抗寒性的一種鑒定方法。半致死溫度（LT50）是該方法中最常用到的指標(biāo)，多用于小麥冬前凍害的鑒定[27?28]，但單獨(dú)利用GRM鑒定冬小麥品種TLSC的情況較少。GRM常與ACCM、IIM等鑒定方法配合使用，通過小麥各項(xiàng)指標(biāo)的恢復(fù)水平對(duì)其TLSC進(jìn)行鑒定。

1.2.2 間接鑒定法（IIM）

IIM是采用化學(xué)和生物技術(shù)研究作物群體、個(gè)體、組織器官與細(xì)胞的生理生態(tài)變化，通過分析各項(xiàng)指標(biāo)與抗寒性之間的關(guān)系來鑒定小麥抗寒性的一種方法。倒春寒發(fā)生后，通過測(cè)定可溶性糖等滲透調(diào)節(jié)物質(zhì)的含量、活性氧含量與抗氧化酶活性、含水量與電導(dǎo)率、光合特性、呼吸特性等相關(guān)指標(biāo)[29?39]，以及對(duì)小麥?zhǔn)軆黾?xì)胞的顯微與超微結(jié)構(gòu)觀察[40]等來鑒定小麥抗寒性的方法均屬于IIM。該方法通過量化各項(xiàng)指標(biāo)準(zhǔn)確反映冬小麥的TLSC，有助于倒春寒危害的溫度閾值研究。但由于機(jī)理復(fù)雜，目前部分測(cè)定指標(biāo)與抗寒性關(guān)系還未有明確定論，該方法的應(yīng)用受到限制。

1.2.3 綜合鑒定法（CIM）

CIM是將多種鑒定方法與鑒定指標(biāo)相結(jié)合，利用統(tǒng)計(jì)分析方法，綜合鑒定或評(píng)價(jià)小麥抗寒性的一種方法。李桐等[41]利用了ACCM、FIM和IIM綜合鑒定冬小麥品種，通過各種方法相互驗(yàn)證研究發(fā)現(xiàn)室內(nèi)快速降溫處理結(jié)合超氧化物歧化酶（SOD）活性及抗壞血酸、過氧化氫和丙二醛（MDA）含量的分析，能有效鑒定出小麥品種抗寒性的強(qiáng)弱。因此，利用ACCM結(jié)合GRM、FIM和IIM等多種方法和多種指標(biāo)綜合鑒定冬小麥TLSC，其鑒定結(jié)果可能更加全面而準(zhǔn)確，但該方法對(duì)技術(shù)、成本要求較高且工作量較大。

1.2.4 數(shù)學(xué)分析方法

主成分分析法（Principal Component Analysis Method）[42]、聚類分析法（Cluster Analysis Method）[43]、極點(diǎn)排序法（Polar Ordination Method）、隸屬函數(shù)法（Membership Function Method）[44?45]、加權(quán)法（Weighting Method）以及模糊評(píng)判法（Fuzzy Evaluation Method）[42]等是TLSC鑒定常用的數(shù)學(xué)分析方法。主成分分析法通過降維和去噪，加快對(duì)樣本有價(jià)值信息的處理速度，常被用來篩選TLSC鑒定的關(guān)鍵性指標(biāo)。馬德栗等[46]利用主成分分析法對(duì)湖北省1951?2010年的倒春寒災(zāi)害事件建立了評(píng)估模型，并對(duì)其危害程度做了等級(jí)劃分。隸屬函數(shù)法、加權(quán)法和模糊評(píng)判法在小麥TLSC鑒定的使用過程中需設(shè)定各指標(biāo)的權(quán)重值，相對(duì)于主成分分析法，其主觀性較強(qiáng)。此外，鑒定中也常用到一些數(shù)學(xué)模型，例如累計(jì)低溫度日（Accumulated Cooling Degree Days，ACDD，℃·d）也能作為數(shù)學(xué)分析方法較好地量化不同低溫脅迫強(qiáng)度和持續(xù)時(shí)間的綜合效應(yīng)[1,47]。

2 冬小麥品種TLSC鑒定指標(biāo)

2.1 鑒定指標(biāo)分類

鑒定冬小麥品種的TLSC，不僅要選擇合適的鑒定方法，還要選用合適的鑒定指標(biāo)。小麥遭受倒春寒危害后，植株內(nèi)部會(huì)發(fā)生一系列復(fù)雜的生理生化變化，外部形態(tài)也會(huì)表現(xiàn)出不同的受害特征，從而影響產(chǎn)量和品質(zhì)。目前，國(guó)內(nèi)外學(xué)者根據(jù)冬小麥植株的內(nèi)外變化及產(chǎn)量等相關(guān)特征，從形態(tài)與農(nóng)藝、生理生化以及氣象與生態(tài)等方面來研究和鑒定冬小麥TLSC（圖2）。

2.2 鑒定指標(biāo)的適用范圍與篩選

（1）形態(tài)與農(nóng)藝指標(biāo)

形態(tài)與農(nóng)藝指標(biāo)一般用于田間自然鑒定法，它包括形態(tài)、穗部結(jié)實(shí)特性以及產(chǎn)量等指標(biāo)，且常以數(shù)量、質(zhì)量、顏色和形態(tài)變化等作為參考項(xiàng)。研究表明[48?49]，倒春寒會(huì)導(dǎo)致小麥葉片發(fā)黃、皺縮甚至枯萎，農(nóng)業(yè)農(nóng)村部標(biāo)準(zhǔn)《NY/T 1307?2007》[22]將小麥葉片受害癥狀作為TLSC分級(jí)的重要指標(biāo)。小麥遭遇倒春寒后，莖的生長(zhǎng)點(diǎn)受到損傷會(huì)導(dǎo)致節(jié)間變色、粗糙、裂開、塌陷或節(jié)擴(kuò)大等癥狀[50]。大量研究表明[51?55]倒春寒對(duì)小麥穗部危害更大，在拔節(jié)后期至孕穗期主要影響幼穗小花的發(fā)育[51]，導(dǎo)致結(jié)實(shí)率大幅降低，從而造成減產(chǎn)。由于同一麥穗不同穗位的小花發(fā)育順序不同[52]，如果該時(shí)期遭遇倒春寒，會(huì)導(dǎo)致退化、不孕小花和小穗數(shù)增加[53?54]，出現(xiàn)異形穗、殘缺穗或無效穗[23]等癥狀。胡新等[55]利用加權(quán)平均法構(gòu)建殘穗指數(shù)來評(píng)價(jià)冬小麥TLSC，同樣發(fā)現(xiàn)殘穗指數(shù)越大，危害發(fā)生越嚴(yán)重。

圖2 冬小麥品種TLSC鑒定指標(biāo)分類

因此，在選擇TLSC鑒定指標(biāo)時(shí)，要充分考慮小麥品種在多年田間試驗(yàn)中的形態(tài)與農(nóng)藝指標(biāo)變化，進(jìn)行田間調(diào)查并統(tǒng)計(jì)出現(xiàn)頻率最高、影響最大的指標(biāo)作為鑒定關(guān)鍵指標(biāo)。此外，鑒定時(shí)還要選擇受倒春寒危害最嚴(yán)重器官的相關(guān)指標(biāo)作為參考。

（2）生理生化指標(biāo)

活性氧、滲透調(diào)節(jié)物質(zhì)、內(nèi)源激素含量、抗氧化酶活性以及光合與呼吸特性常被作為TLSC鑒定的生理生化指標(biāo)，該類指標(biāo)一般用于間接鑒定法。研究表明，幼穗、葉片與根系的可溶性糖、游離脯氨酸、可溶性蛋白、MDA含量及過氧化物酶、SOD等抗氧化酶活性與相關(guān)內(nèi)源激素含量等均可作為小麥TLSC的鑒定生理生化指標(biāo)[27,29?32]。此外，自由水/束縛水比值、電解質(zhì)滲透率和電導(dǎo)率等也常被用作TLSC鑒定的生理生化指標(biāo)[33?34]。光合和呼吸特性方面，光合速率、蒸騰速率、葉片氣孔導(dǎo)度、胞間CO2濃度及最大光化學(xué)效率（Fv/Fm）也可作為鑒定冬小麥品種TLSC的指標(biāo)[35?38]。Kurimoto等[39]研究認(rèn)為，低溫下植物呼吸穩(wěn)態(tài)的保持和較高的ATP生產(chǎn)效率有助于維持正常的生長(zhǎng)發(fā)育，表明呼吸特性相關(guān)指標(biāo)用于冬小麥品種TLSC鑒定也具有可行性。

由于倒春寒分為0℃以上的冷害和0℃以下的凍害兩種類型，利用生理生化指標(biāo)鑒定時(shí)要考慮這兩種脅迫水平生理生化響應(yīng)機(jī)制的差異，從而篩選出適宜的鑒定指標(biāo)。倒春寒發(fā)生后，小麥生理生化指標(biāo)的變化比農(nóng)藝指標(biāo)更為顯著，對(duì)測(cè)定環(huán)節(jié)的技術(shù)要求更高，都可能造成結(jié)果的準(zhǔn)確度和實(shí)用性降低。因此，鑒定時(shí)利用相對(duì)受害值作為評(píng)價(jià)指標(biāo)，其結(jié)果更具代表性。

（3）氣象與生態(tài)指標(biāo)

前人研究常以氣溫作為冬小麥品種TLSC鑒定的指標(biāo)，但實(shí)際上地溫會(huì)補(bǔ)償?shù)蜏靥鞖馑鶐淼呢?fù)面影響，致使小麥生長(zhǎng)的環(huán)境溫度要比氣溫高。因此，冠層溫度更能代表小麥生長(zhǎng)的環(huán)境溫度[56]，將冠層溫度作為鑒定的氣象指標(biāo)更為切合實(shí)際。在具體的鑒定應(yīng)用中，如GRM還常利用LT50和恢復(fù)生長(zhǎng)最低溫度作為冬小麥品種TLSC的鑒定指標(biāo)。

鑒定冬小麥品種TLSC時(shí)，還易受光照強(qiáng)度、田間CO2濃度、空氣濕度和土壤肥力等因素的影響，導(dǎo)致結(jié)果的準(zhǔn)確性降低。此外，鑒定中還需考慮群體與個(gè)體及不同器官之間的差異，明確三者與氣象和生態(tài)指標(biāo)之間的關(guān)系，對(duì)大田生產(chǎn)上鑒定冬小麥品種TLSC更具實(shí)際意義。

3 冬小麥品種TLSC鑒定原則及程序

3.1 鑒定原則

（1）鑒定結(jié)果具有地域性。由于不同地區(qū)倒春寒易災(zāi)期不同，同一品種在不同地區(qū)可能表現(xiàn)出不同的TLSC，小麥的敏感期與易災(zāi)期重合將對(duì)小麥造成更為嚴(yán)重的危害。

（2）指標(biāo)選用具有針對(duì)性。小麥主莖與分蘗的生育時(shí)期并不完全同步，抗倒春寒能力也不相同，因此，鑒定時(shí)要注意區(qū)分是主莖還是分蘗帶來的差異，還要統(tǒng)一鑒定的生育時(shí)期。此外，由于冬小麥不同生育時(shí)期各個(gè)器官的抗寒能力不同，在鑒定時(shí)還應(yīng)重點(diǎn)考慮不同器官敏感性的差異。

（3）鑒定方法和指標(biāo)具有綜合性。冬小麥品種TLSC是一個(gè)復(fù)雜的生物學(xué)數(shù)量性狀，其鑒定方法和指標(biāo)之間并不是相對(duì)獨(dú)立的，單獨(dú)使用其中任何一種單項(xiàng)指標(biāo)或單個(gè)方法，所得的結(jié)果都具有片面性[57?58]。鑒定時(shí)還需結(jié)合相關(guān)氣象指標(biāo)同步進(jìn)行，注重地溫和冠層溫度對(duì)鑒定結(jié)果的影響[59]。

（4）鑒定時(shí)需排除交叉逆境脅迫的干擾。在鑒定冬小麥品種TLSC過程中要嚴(yán)格控制干旱等交叉逆境脅迫的干擾，以免影響鑒定結(jié)果的準(zhǔn)確性。

（5）鑒定方法和指標(biāo)要具有實(shí)用性。鑒定指標(biāo)和方法的研究和選用，要服務(wù)于冬小麥抗倒春寒新品種選育，符合“簡(jiǎn)單易行、易推廣、低成本”原則。

3.2 鑒定程序

鑒定冬小麥品種TLSC時(shí)，需要結(jié)合各地氣候條件和栽培管理方式，選用合適的鑒定方法和指標(biāo)，針對(duì)不同氣候的小麥產(chǎn)區(qū)建立不同的鑒定方法和指標(biāo)體系，以便選育出適合當(dāng)?shù)氐目沟勾汉缕贩N。目前，小麥育種中鮮有TLSC鑒定標(biāo)準(zhǔn)程序，因此，結(jié)合前人研究和鑒定原則，總結(jié)出一套冬小麥品種TLSC鑒定程序如圖3。該鑒定程序通過氣象易災(zāi)期和品種敏感期分析，結(jié)合適宜的鑒定方法和指標(biāo)對(duì)冬小麥品種TLSC進(jìn)行綜合鑒定，對(duì)國(guó)家和地方推廣品種均有一定參考價(jià)值。

圖3 冬小麥品種TLSC鑒定程序

4 新興技術(shù)在冬小麥品種TLSC鑒定中的應(yīng)用

近年來，農(nóng)業(yè)遙感技術(shù)、表型組學(xué)、作物模型、蛋白質(zhì)組學(xué)以及基因標(biāo)記技術(shù)飛速發(fā)展，對(duì)農(nóng)業(yè)生產(chǎn)的快速發(fā)展起到了巨大的推動(dòng)作用[60?73]。遙感技術(shù)[60?61]被認(rèn)為是大范圍探測(cè)作物物候狀況的可靠工具，利用智能手機(jī)、無人機(jī)或者相機(jī)對(duì)冬小麥進(jìn)行近地表遙感圖像監(jiān)測(cè)，評(píng)估極端天氣事件對(duì)作物的影響，將遙感探測(cè)到的植被指數(shù)與氣象信息相結(jié)合，可作為天氣變化對(duì)作物產(chǎn)量影響的評(píng)價(jià)和鑒定方法，成本低且快速高效，可及時(shí)做出預(yù)防或應(yīng)對(duì)。還有學(xué)者[62?63]將高時(shí)空分辨率遙感數(shù)據(jù)與作物模型相結(jié)合，獲取農(nóng)田作物生長(zhǎng)發(fā)育實(shí)時(shí)狀況的信息。作物模型方面，國(guó)際上小麥生理模型的研究發(fā)展迅速[64?65]，其中CERES-Wheat[66]、WOFOST[67]、FROSTOL[68]等模型可進(jìn)行低溫模擬并已在生產(chǎn)上推廣應(yīng)用。蛋白質(zhì)組學(xué)方面，低溫脅迫下植物體內(nèi)大量基因表達(dá)，誘導(dǎo)合成低溫脅迫響應(yīng)蛋白以抵御低溫對(duì)植物的破壞[69]。Janmohammadi等[70]研究表明，低溫脅迫初期光合作用代謝通路中的放氧增強(qiáng)蛋白、NADH脫氫酶和脫氫抗壞血酸還原酶等蛋白下調(diào)。基因標(biāo)記方面，可通過QTL等分子標(biāo)記技術(shù)[71]結(jié)合傳統(tǒng)育種技術(shù)，使育種家無需等待田間試驗(yàn)中倒春寒的發(fā)生就可選擇耐寒關(guān)鍵基因[72]，從而加快小麥抗倒春寒新品種的選育。薛輝等[73]研究發(fā)現(xiàn)，1B染色體上的AX-109492586、3A染色體上的AX-110064042具有較好的耐倒春寒特性，可用于開發(fā)CAPS標(biāo)記。但目前來看，新興技術(shù)在冬小麥品種TLSC上的應(yīng)用還較少，且與小麥抗寒性能方面的交叉研究不足。

5 總結(jié)與展望

5.1 深入冬小麥抗寒生理研究，完善TLSC鑒定方法和指標(biāo)體系

冬小麥抗寒生理研究是完善TLSC鑒定方法與指標(biāo)的重要基礎(chǔ)，這些研究包括冬小麥倒春寒發(fā)生的致災(zāi)溫度閾值、敏感生育時(shí)期、敏感部位以及生理、生化和生態(tài)變化等。今后需繼續(xù)深入小麥倒春寒災(zāi)害的災(zāi)變過程和致災(zāi)機(jī)理的研究，從而加快推進(jìn)國(guó)家和地方冬小麥品種TLSC鑒定標(biāo)準(zhǔn)的制定和推廣。

5.2 加快推進(jìn)新興技術(shù)在冬小麥品種TLSC鑒定上的研究與應(yīng)用

隨著現(xiàn)代信息技術(shù)的快速發(fā)展，大數(shù)據(jù)技術(shù)與作物模型、表型組學(xué)、遙感技術(shù)及分子生物技術(shù)的結(jié)合是冬小麥品種TLSC鑒定研究的創(chuàng)新突破點(diǎn)。CRISPR/Cas9和基因堆積等分子生物技術(shù)在育種工作中的應(yīng)用已逐漸成熟，可對(duì)小麥抗倒春寒基因進(jìn)行定位和編輯[74]。目前，國(guó)內(nèi)外學(xué)者已經(jīng)鑒定出一批抗寒基因和抗寒蛋白，未來應(yīng)該對(duì)已鑒定出的抗倒春寒基因或蛋白質(zhì)進(jìn)行重點(diǎn)篩選，驗(yàn)證其作為冬小麥品種TLSC鑒定指標(biāo)的可靠性，推進(jìn)分子鑒定指標(biāo)體系的構(gòu)建，加快研究成果轉(zhuǎn)化到實(shí)際生產(chǎn)應(yīng)用和抗倒春寒新品種選育工作中。

5.3 加快研究和開發(fā)冬小麥品種TLSC鑒定產(chǎn)品

目前，育種機(jī)構(gòu)常需采用多年復(fù)雜的傳統(tǒng)方法來鑒定小麥新品種的TLSC，成本大，周期長(zhǎng)，效率低，使新品種的推廣受到阻礙，亟需一種既能提升鑒定效率又能降低成本的TLSC鑒定產(chǎn)品。這種產(chǎn)品既可以是鑒定儀器，也可以是鑒定服務(wù)，這對(duì)推進(jìn)抗倒春寒小麥新品種的推廣種植意義非凡。此外，鑒定產(chǎn)品的研究和開發(fā)既要服務(wù)于新品種選育，還要服務(wù)于生產(chǎn)實(shí)際。對(duì)農(nóng)民來說，倒春寒災(zāi)前預(yù)防，災(zāi)后及時(shí)采取減災(zāi)措施，選用簡(jiǎn)便易行、快速和成本低廉的方法和產(chǎn)品更具實(shí)際意義。如應(yīng)用抗寒劑提升小麥TLSC也是有效的防災(zāi)減災(zāi)手段，利用現(xiàn)有的研究基礎(chǔ)開發(fā)成本低廉的抗寒劑與配套的農(nóng)機(jī)裝備應(yīng)是今后研究的重點(diǎn)方向之一。因此，加強(qiáng)研究并開發(fā)服務(wù)于育種工作的抗寒性鑒定高端技術(shù)產(chǎn)品和服務(wù)于生產(chǎn)中防災(zāi)減災(zāi)的實(shí)用新型技術(shù)產(chǎn)品，對(duì)提高中國(guó)小麥生產(chǎn)的防災(zāi)、減災(zāi)能力，促進(jìn)小麥生產(chǎn)提質(zhì)減損增效，保障國(guó)家糧食安全具有重要意義。

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An Inclusive Comparison of Identification Methods and Indices of Winter Wheat Tolerance against Late Spring Coldness

ZHANG Le-le1, CHEN Xiang1, KE Yuan-yuan1, LIU Bin-bin1, MUHAMMAD Ahmad Hassan1, ZHANG Yan1, XU Hui1, LI Jin-cai1,2

(1.College of Agronomy，Anhui Agricultural University/Crop Cultivation Science Observatory in East China of the Ministry of Agriculture and Rurad Affairs，Hefei 230036，China；2.Collaborative Innovation Center of Modern Crop Production in Jiangsu，Nanjing 210095)

Late spring coldness is an important factor limiting the stable and high yield of wheat in Huang-huai winter wheat region. New wheat varieties with tolerance to late spring coldness is an important strategy to reduce the damage of late spring coldness. It is of great significance to establish a simple, easy to popularize and comprehensive evaluation methods and indices system of wheat tolerance to late spring coldness for ensuring food security. In this paper, the methods and indices for identification of winter wheat tolerance to late spring coldness were summarized, compared and evaluated, and the identification principles and procedures were also put forward. (1) The identification methods of winter wheat tolerance to late spring coldness mainly consisted of direct identification method, indirect identification method and comprehensive identification method. Direct identification methods consisted of field natural identification method, artificial climate chamber method and growth recovery method. The field natural identification method had the characteristics of most widely used in the introduction and cultivation, but it had long cycle and heavy workload. Artificial climate chamber method had the characteristics of short cycle and strong repeatability, but it had high requirements for equipment and technology. The growth recovery method was widely used in the identification of freezing injury before winter, but it was less used in the identification of tolerance to late spring coldness. The indirect identification method was helpful to study the temperature threshold of late spring coldness, but the relationship between the indices and cold resistance had not been clear, so its application was limited. The comprehensive identification method had the characteristics of comprehensive and accurate appraisal results, but it had high requirements for technology, cost and workload. Therefore, in the identification process, it was necessary to combine the actual situation, reasonable collocation and comprehensive use of a variety of methods. (2) The identification indices mainly consisted of morphological and agronomic, physiological and biochemical, meteorological and ecological indices. Morphological and agronomic indices were generally used in field natural identification, and the quantity, quality, color and morphological changes were often used as reference items. Physiological and biochemical indices were generally used for indirect identification, including reactive oxygen species, osmotic adjustment substances, endogenous hormone content, antioxidant enzyme activity, photosynthetic and respiratory characteristics, etc. Meteorological and ecological indices were often used in various identification methods. Air temperature, ground temperature, canopy temperature, half lethal temperature and minimum temperature of restoration growth were often used as identification indices, while factors such as light, CO2, humidity and fertility could affect the accuracy of results. (3) Appraisal principles and procedures. This paper put forward the identification principles, such as the regionality of identification results, the pertinence of indices selection, the comprehensiveness and practicability of identification methods and indices, etc. The identification procedures were mainly composed of disaster prone period, variety sensitive period, sensitive organs, identification methods and indices, low temperature degree, field or pot experiment, evaluation and classification of cold resistance by mathematical analysis method and writing report. Finally, the prospects of cold resistance physiology research, identification research and application of emerging technologies and identification product development were made.

Winterwheat; Late spring coldness; Identification; Index; Method

10.3969/j.issn.1000-6362.2021.02.006

張樂樂,陳翔,柯媛媛,等.冬小麥抗倒春寒性能鑒定方法和指標(biāo)的比較[J].中國(guó)農(nóng)業(yè)氣象,2021,42(2):146-157

2020?12?02

“十三五”國(guó)家重點(diǎn)研發(fā)計(jì)劃課題（2017YFD0300408）；安徽省自然科學(xué)基金（2008085QC122）；安徽省科技重大專項(xiàng)（202003b06020021）

李金才，教授，博導(dǎo)，主要從事作物生理生態(tài)研究，E-mail:ljc5122423@126.com

張樂樂，E-mail：zhanglele912@163.com