韓云華，王彥榮，陶奇波

(草地農(nóng)業(yè)生態(tài)系統(tǒng)國(guó)家重點(diǎn)實(shí)驗(yàn)室 蘭州大學(xué)草地農(nóng)業(yè)科技學(xué)院，甘肅 蘭州 730020)

植物生產(chǎn)層

種子激素引發(fā)

韓云華，王彥榮，陶奇波

(草地農(nóng)業(yè)生態(tài)系統(tǒng)國(guó)家重點(diǎn)實(shí)驗(yàn)室 蘭州大學(xué)草地農(nóng)業(yè)科技學(xué)院，甘肅 蘭州 730020)

種子激素引發(fā)(Hormonal priming)是種子引發(fā)(Seed priming)的一種。通過(guò)激素引發(fā)可以有效改善種子萌發(fā)狀態(tài)，促進(jìn)幼苗生長(zhǎng)和產(chǎn)量提高，并增強(qiáng)植物的抗逆性。本文針對(duì)近年來(lái)植物激素引發(fā)的研究情況，介紹了激素引發(fā)在促進(jìn)種子萌發(fā)和幼苗生長(zhǎng)中的應(yīng)用，論述了其對(duì)植物生長(zhǎng)發(fā)育的有利影響。并從植物生理生化與分子生物學(xué)兩個(gè)方面闡述了種子激素引發(fā)的機(jī)理。同時(shí)，對(duì)影響引發(fā)效果的因素進(jìn)行了分析，指出種子引發(fā)的濃度與引發(fā)時(shí)間、種子引發(fā)后的回干條件是影響引發(fā)效果的最重要因素。未來(lái)的研究應(yīng)充分借助基因組學(xué)與蛋白質(zhì)組學(xué)等方法，深入探究種子激素引發(fā)的生理生化與分子生物學(xué)機(jī)理，并加強(qiáng)在草類(lèi)植物中的研究。

種子引發(fā)；激素；抗逆性；萌發(fā)；產(chǎn)量

種子引發(fā)(seed priming)是以水分為基礎(chǔ)的播前處理技術(shù)，通過(guò)控制種子吸水,激活種子萌發(fā)早期的新陳代謝過(guò)程，然后及時(shí)脫水，阻止其進(jìn)入完全萌發(fā)狀態(tài)[1]。根據(jù)引發(fā)物質(zhì)不同，種子引發(fā)的方法包括水引發(fā)(hydroproming)[2]、滲透引發(fā)(osmopriming)[3]、生物引發(fā)(biopriming)[4]、營(yíng)養(yǎng)引發(fā)(nutrient priming)[5]、化學(xué)物質(zhì)引發(fā)(chemical priming)[6]和激素引發(fā)(hormonal priming)[7]等(表1)。

國(guó)內(nèi)外有關(guān)引發(fā)的研究已經(jīng)有很多[8-16]，但是針對(duì)種子引發(fā)中激素的應(yīng)用及其原理還未見(jiàn)系統(tǒng)評(píng)述。本文從種子引發(fā)中激素的應(yīng)用、激素引發(fā)的生理生化基礎(chǔ)和影響激素引發(fā)等方面論述了近年來(lái)的成果。

激素作為一種參與種子萌發(fā)、植株生長(zhǎng)的信號(hào)小分子物質(zhì)，即使在植物體內(nèi)濃度極小甚至趨近于0時(shí)仍有非常重要的作用[9]。種子激素引發(fā)是利用一定濃度的激素進(jìn)行浸種，并精確控制溫度和時(shí)間，達(dá)到促進(jìn)萌發(fā)且不引起傷害的技術(shù)。種子激素引發(fā)的主要作用包括以下幾點(diǎn)：1)提高種子活力，即提高種子的發(fā)芽率、發(fā)芽速度以及整齊度；2)提高幼苗抗逆性；3)打破種子休眠；4)在一定程度上恢復(fù)種子活力。目前已有報(bào)道的激素種類(lèi)有赤霉素(GA)[7]、脫落酸(ABA)[17]、油菜素內(nèi)酯(BR)[18]、5-氨基乙酰丙酸(ALA)[19]、細(xì)胞分裂素(CTK)[20]等。通過(guò)播前激素引發(fā)，可有效改善種子質(zhì)量，提高抗逆性，促進(jìn)產(chǎn)量增加[9,21-22]，而且可以增加豆科植物種子維生素含量和營(yíng)養(yǎng)價(jià)值[23]。

表1 幾種常見(jiàn)種子引發(fā)技術(shù)的比較Table 1 Comparison of several common seed priming technology

1 種子激素引發(fā)的應(yīng)用

激素引發(fā)是種子引發(fā)技術(shù)的一種，廣泛應(yīng)用于農(nóng)業(yè)生產(chǎn)中，早在20世紀(jì)50年代，已有學(xué)者利用赤霉素引發(fā)豌豆(Pisumsativum)種子，以提高其出苗能力與幼苗長(zhǎng)勢(shì)[24];20世紀(jì)70年代亦有研究人員發(fā)現(xiàn)赤霉素引發(fā)可以提高洋蔥種子的萌發(fā)率[25]。經(jīng)過(guò)半個(gè)多世紀(jì)的發(fā)展，激素引發(fā)已經(jīng)廣泛應(yīng)用于農(nóng)林業(yè)生產(chǎn)。另外，由于激素參與調(diào)節(jié)植物生長(zhǎng)、發(fā)育和繁殖等生長(zhǎng)過(guò)程，激素引發(fā)還可以改善植物抗逆性，增加其產(chǎn)量。

1.1 改善種子萌發(fā)狀態(tài)

種子萌發(fā)過(guò)程伴隨著眾多生理生化過(guò)程，激素引發(fā)是在水引發(fā)的基礎(chǔ)上加入外源植物激素，達(dá)到打破休眠、促進(jìn)萌發(fā)和緩解老化活力喪失的作用。赤霉素類(lèi)激素(GAs)是目前引發(fā)中應(yīng)用最廣的一類(lèi)植物激素。近年來(lái)，在小麥(Triticumaestivum)[26]、大麥(Hordeumvulgare)[27]、燕麥(Avenasativa)[28]、番茄(Lycopersiconesculentum)[29]、大白菜(Brassicarapa)[30]、向日葵(Helianthusannuus)[31]、苜蓿(Medicagosativa)[32-33]等植物中，科研人員通過(guò)控制赤霉素濃度、引發(fā)時(shí)間和引發(fā)溫度等因素，發(fā)現(xiàn)GAs引發(fā)可增加種子吸水量，提高種子活力、發(fā)芽率、發(fā)芽指數(shù)、發(fā)芽速率，并顯著降低電導(dǎo)率、起始發(fā)芽時(shí)間、50%發(fā)芽時(shí)間和平均發(fā)芽時(shí)間。表2展示了幾種常見(jiàn)植物在激素引發(fā)下的萌發(fā)狀況。

除GAs外，其它一些植物激素也被證明具有促進(jìn)萌發(fā)的作用。雀麥(Bromusinermis)種子人工老化后，利用生長(zhǎng)素(NAA)引發(fā)，可提高出苗率、幼苗活力、幼苗生長(zhǎng)速率、根長(zhǎng)、芽長(zhǎng)和幼苗高度[35]。利用24-表油菜素內(nèi)酯(24-epibrassinolide)滾筒引發(fā)后的鈴椒(Capsicumannuum)種子表現(xiàn)出明顯的發(fā)芽和生長(zhǎng)優(yōu)勢(shì)[36]。

1.2 促進(jìn)幼苗生長(zhǎng)和提高產(chǎn)量

除影響種子萌發(fā)以外，激素引發(fā)的影響效果會(huì)延續(xù)至幼苗生長(zhǎng)甚至到產(chǎn)量形成時(shí)期。眾多研究結(jié)果表明，GAs引發(fā)可提高出苗率、出苗整齊度、根系長(zhǎng)度、幼苗根系活力、幼苗干重、葉片中葉綠素含量和植株高度[27-28,33,37-39]。但是，也有研究表明GA3引發(fā)效果不如水引發(fā)，利用GA3和水引發(fā)生菜(Iceberglettuce)種子，鹽脅迫處理15 d后，GA3處理的幼苗干重小于水引發(fā)處理[40]。

表2 幾種植物在激素引發(fā)下的萌發(fā)率Table 2 Germination percentage of several plants as influenced by hormone priming

其它植物激素也有類(lèi)似效果。如CTK可增加幼苗葉面積[30]。BR可降低種子電導(dǎo)率，增加幼苗葉綠素含量[41]。在200 mmol·L-1NaCl脅迫下，水楊酸引發(fā)后的甜高粱(Sorghumbicolor)種子可分別提高出苗率、出苗速率、葉綠素b和蛋白含量82%，130%、7.9%和1.9%。在37 ℃下，出苗率、出苗速率和根數(shù)增加了72.5%、108.5%和63.8%，同時(shí)丙二醛(MDA)含量降低了17.6%[42]。植物激素引發(fā)可提高種子產(chǎn)量。GA3可顯著提高小扁豆(Lensculinaris)產(chǎn)量因子和種子產(chǎn)量[35]，CTK引發(fā)后兩個(gè)品種的小麥產(chǎn)量均得到了提高[43-44](表3)。

表3 激素引發(fā)對(duì)幾種植物株高的影響Table 3 Effects of seed priming on plant height of several plants

1.3 抗逆應(yīng)用

激素引發(fā)可有效提高抗氧化酶系統(tǒng)活性，進(jìn)而提高幼苗抗逆特性[45,48]。GA3可以顯著提高干旱脅迫下油菜種子的發(fā)芽率，也可顯著提高幼苗抗旱性，引發(fā)后種苗鮮重和下胚軸長(zhǎng)度均有所增加[49]。研究發(fā)現(xiàn)，GA3引發(fā)可顯著提高耐鹽油菜品種在鹽脅迫條件下的表現(xiàn)，但是對(duì)不耐鹽品種油菜影響不顯著[50]。脫落酸(ABA)是GA的拮抗激素，可增加種子休眠。近些年研究發(fā)現(xiàn)，ABA引發(fā)可在一定程度上緩解小麥水淹和干旱脅迫[51]，也可提前出苗時(shí)間、提高出苗率、縮短出苗期[52]。

除了GAS和ABA以外，其它植物激素引發(fā)也會(huì)增加幼苗一定程度的抗逆能力。細(xì)胞分裂素(CTK)引發(fā)后，植株內(nèi)ABA含量降低，提高了小麥抗鹽能力[43]，在50%田間持水量下，BR引發(fā)后的植株有較高的葉水勢(shì)且積累更多的CO2，提高了水稻抗旱性[52]。ALA與KNO3配合引發(fā)紅辣椒，可顯著提高種子在低溫條件下(15 ℃)的表現(xiàn)，發(fā)芽率、發(fā)芽速率均得到顯著提高。種子貯藏一個(gè)月后(4 ℃或25 ℃)，引發(fā)效果依然很好[53]。

2 種子激素引發(fā)的生理生化基礎(chǔ)

2.1 細(xì)胞膜修復(fù)

細(xì)胞膜是植物細(xì)胞的基本功能單位，對(duì)細(xì)胞具有保護(hù)作用。種子活力喪失往往伴隨著細(xì)胞膜的損傷，而引發(fā)可在一定程度上對(duì)損傷的細(xì)胞膜進(jìn)行修復(fù)。研究發(fā)現(xiàn)，GA3引發(fā)后油菜種子電導(dǎo)率低于未被引發(fā)的，且植株細(xì)胞受到鹽脅迫損傷程度較低[54]。

2.2 改變細(xì)胞膜離子通透性

激素引發(fā)還可改變細(xì)胞膜對(duì)離子的選擇性，進(jìn)而增加植物抗逆特性。GA3引發(fā)可降低鹽脅迫下小麥芽部和根部的鈉離子含量，提高鈣離子和鉀離子含量，并提高水楊酸(SA)含量，降低ABA和聚胺類(lèi)物質(zhì)(腐胺Put和亞精胺Spd)含量，進(jìn)而提高其抗逆能力，促進(jìn)種子產(chǎn)量的提高[45]。水楊酸和抗壞血酸引發(fā)可降低小麥幼苗中鉀離子含量，增加可溶性糖含量[55]。生長(zhǎng)素(NAA)也有類(lèi)似的效果[56]。

2.3 激活抗氧化系統(tǒng)

激素引發(fā)可促進(jìn)植株內(nèi)抗氧化系統(tǒng)水平提升[32]。ABA引發(fā)的小麥種子受到干旱脅迫后，可顯著提高超氧化物歧化酶(SOD)和過(guò)氧化物酶(POD)活性，且細(xì)胞內(nèi)相對(duì)含水量顯著增加[57]。GA3引發(fā)后的油菜幼苗受到干旱脅迫后，中可溶性糖、可溶性蛋白和自由脯氨酸均顯著提高，而丙二醛(MDA)含量下降，抗氧化系統(tǒng)酶(如SOD，CAT，POD)水平升高[49]。

3 種子激素引發(fā)的分子生物學(xué)基礎(chǔ)

3.1 大分子物質(zhì)修復(fù)

DNA修復(fù)是種子萌發(fā)前重要的階段，可以避免種子萌發(fā)所需蛋白的錯(cuò)配，促進(jìn)種子迅速而整齊的萌發(fā)。研究者利用qRT-PCR激素研究了7個(gè)與DNA修復(fù)有關(guān)的基因(GTFⅡH2，MMZ3/UVE1C，RAD3，RecA-like1，RAD54，UDNAglycosylase和KU80)在種子引發(fā)后的表達(dá)情況，發(fā)現(xiàn)這些基因表達(dá)量會(huì)上調(diào)，但是受引發(fā)方法影響較大[58]。細(xì)胞學(xué)研究發(fā)現(xiàn)，ABA引發(fā)可觸發(fā)DNA修復(fù)機(jī)制，降低細(xì)胞第1次有絲分裂過(guò)程中染色體畸變頻率[59-60]。

3.2 促進(jìn)特定基因表達(dá)

引發(fā)后種子在逆境條件下可促進(jìn)某些特定基因表達(dá)，增強(qiáng)植物抗逆能力。外源ABA引發(fā)后，降低了cDNA克隆BnCAM1表達(dá)量，過(guò)量表達(dá)鈣調(diào)蛋白，促進(jìn)種子萌發(fā)早期新陳代謝酶類(lèi)合成，在非生物脅迫條件下(干旱、低溫和鹽脅迫)，使發(fā)芽時(shí)間提前2～7 d，且提高了發(fā)芽率[34]。研究發(fā)現(xiàn)，水孔蛋白基因BnPIP1編碼的蛋白與種子萌發(fā)早期營(yíng)養(yǎng)物質(zhì)酶代謝中水分專(zhuān)業(yè)有關(guān)，ABA引發(fā)可促進(jìn)基因BnPIP1表達(dá)上調(diào)，且基因表達(dá)時(shí)間提前，促進(jìn)種子萌發(fā)[61]。

4 影響種子激素引發(fā)的因素

種子引發(fā)涉及的生理生化、分子生物學(xué)過(guò)程很多。最直接影響激素引發(fā)的因素主要有引發(fā)時(shí)間、激素濃度、引發(fā)溫度和回干條件等。

4.1 引發(fā)時(shí)間和溫度

激素引發(fā)本質(zhì)上是液體引發(fā)的一種，且激素濃度都極低，溶液滲透勢(shì)高，種子易發(fā)生吸漲傷害。因此，要適當(dāng)控制引發(fā)時(shí)間和溫度，以便達(dá)到最佳的引發(fā)效果而不引起吸漲傷害。目前研究中引發(fā)時(shí)間普遍在6～24 h[26,28,32,36-37,40,55]。也有一些物種處理比較特殊，如用366 mg·kg-1GA3引發(fā)1 h顯著促進(jìn)白花蛇舌草(Hedyotisdiffusa)種子萌發(fā),提高干旱條件下種子的發(fā)芽率、發(fā)芽指數(shù)、活力指數(shù)并促進(jìn)幼苗的生長(zhǎng)[62]。Wagner等[63]在研究10種牧草種子引發(fā)試驗(yàn)中將種子在15 ℃下引發(fā)14 d。有報(bào)道稱(chēng)，引發(fā)在低溫下效果較好，且最佳溫度在15～20 ℃[61]。但是，也有研究采用高于20 ℃作為激素引發(fā)的溫度[38,64-66]。

4.2 激素濃度

濃度是影響引發(fā)效果的一個(gè)重要因素，但是不同物種間以及不同引發(fā)劑間最佳濃度各異。GAs是一類(lèi)最常用的種子引發(fā)激素，其濃度在不同物種間差異較大。鹽脅迫條件下，20 mg·L-1GA3引發(fā)小麥種子后發(fā)芽率最高[67]。在干旱脅迫條件下，用0.5 mmol·L-1GA3引發(fā)48 h處理北青蘭(Dracocephalumargunense)效果最好[68]。利用300 mg·L-1GA3引發(fā)油菜種子可顯著提高其幼苗抗旱性，提高幼苗鮮重和下胚軸長(zhǎng)[48]。

不同引發(fā)物的引發(fā)濃度也有較大差異。利用SA濃度50 mg·g-1引發(fā)春玉米，可顯著提高生物量、種子產(chǎn)量和收獲指數(shù)[40]；ABA 濃度 10～6 mol·L-1處理小麥種子效果較好[56]。50 mg·L-1的NAA干旱條件下高冰草(Thinopyrumponticum)發(fā)芽率增加了18%[44]，而NAA 150 mg·L-1引發(fā)對(duì)小麥生長(zhǎng)和種子產(chǎn)量增加最為有利[55]。

4.3 種子回干

實(shí)際應(yīng)用中，種子經(jīng)過(guò)激素引發(fā)后，一般需要經(jīng)過(guò)表面回干或者完全回干后儲(chǔ)藏，待田間條件合適后進(jìn)行播種。因此，引發(fā)后的回干參數(shù)相當(dāng)重要。對(duì)KNO3引發(fā)后的西瓜(Citrulluslanatus)種子設(shè)置了不同的回干處理，研究其對(duì)抗氧化酶活性和發(fā)芽率的影響，發(fā)現(xiàn)室內(nèi)緩慢回干效果最好[69]。研究發(fā)現(xiàn)，回干時(shí)間和回干溫度對(duì)羊草(Leymuschinensis)和無(wú)芒雀麥(Bromusinermis)的發(fā)芽率和發(fā)芽指數(shù)都有影響，羊草種子能忍受長(zhǎng)時(shí)間的低溫回干(20 ℃，20 d)，無(wú)芒雀麥種子能忍受短時(shí)間的高溫回干(30 ℃，1～10 d)[70]。

5 展望

種子的激素引發(fā)技術(shù)的研究已有數(shù)十年歷史，研究的內(nèi)容也從種子萌發(fā)、植株生長(zhǎng)和生產(chǎn)性能觀測(cè)深入到生理機(jī)制、信號(hào)通路和分子機(jī)理的研究。相對(duì)于蔬菜、花卉和農(nóng)作物而言，由于多年來(lái)對(duì)牧草種子的重視程度不夠，加之激素引發(fā)成本較高，牧草種子中激素引發(fā)的研究應(yīng)用還較少。牧草種子具有種類(lèi)繁多、特性各異、野生性狀強(qiáng)等特征，未來(lái)的研究應(yīng)集中在有應(yīng)用價(jià)值的栽培草種和野生草種中，在繼續(xù)篩選高效、廉價(jià)的植物激素的同時(shí)，加強(qiáng)牧草種子引發(fā)的生理機(jī)制和分子機(jī)理研究。同時(shí)，可借助蛋白組學(xué)和基因組學(xué)等研究方法，分析激素引發(fā)的種子和未引發(fā)種子在萌發(fā)、幼苗生長(zhǎng)、植株抗逆、產(chǎn)量形成和內(nèi)源激素信號(hào)轉(zhuǎn)導(dǎo)過(guò)程中的基因表達(dá)差異、代謝差異、生化過(guò)程差異，進(jìn)而對(duì)引發(fā)的機(jī)理形成更加深入的理解[71-75]。

雖然已經(jīng)出現(xiàn)了一些多種物質(zhì)配合引發(fā)種子的研究，如先利用PEG引發(fā)，再用GA3引發(fā)，效果要比二者同時(shí)使用和先GA3后PEG好[76]。但是大多數(shù)研究依然采用單一引發(fā)物質(zhì)或方法，采取多種引發(fā)物質(zhì)或者不同引發(fā)方法結(jié)合研究還較為少見(jiàn)。除此以外，當(dāng)前研究主要集中在引發(fā)后種子的萌發(fā)效果方面，有關(guān)引發(fā)后回干的相關(guān)研究較少。不同的回干方法對(duì)種子失水速率影響很大，進(jìn)而影響種子引發(fā)效果。因此，對(duì)引發(fā)后回干技術(shù)進(jìn)行研究對(duì)于提高引發(fā)效果有積極意義。

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(責(zé)任編輯 茍燕妮)

2016年11月國(guó)際市場(chǎng)主要畜產(chǎn)品與飼料價(jià)格分析

11月國(guó)際飼料和畜產(chǎn)品價(jià)格均跌漲互現(xiàn)。

一、大豆、豆粕、菜籽和豆粉市場(chǎng)價(jià)格上漲。玉米、高粱、苜蓿粉和棉籽餅市場(chǎng)價(jià)格下跌。

11月份美國(guó)大豆、豆粕、菜籽和豆粉市場(chǎng)平均價(jià)格分別為373.86、313.82、472.35和299.37美元·t-1，環(huán)比分別上漲2.86%、2.60%、4.65% 和1.50%。玉米、高粱、苜蓿粉和棉籽餅市場(chǎng)平均價(jià)格分別為135.73、147.51、213.95和268.75美元·t-1，環(huán)比分別下降1.40%、1.08%、2.75%、8.90%。

二、育肥牛、牛奶和羊肉市場(chǎng)價(jià)格上漲，瘦肉豬、羊羔肉、豬肉和牛肉市場(chǎng)價(jià)格下降，雞肉市場(chǎng)價(jià)格與10月持平。

11月份美國(guó)育肥牛、牛奶、瘦肉豬和牛肉市場(chǎng)平均價(jià)格分別為2.77、0.28、1.05和5.41美元·kg-1，其中育肥牛和牛奶市場(chǎng)價(jià)格環(huán)比分別上漲1.77%和4.44%，瘦肉豬和牛肉市場(chǎng)價(jià)格環(huán)比分別下降1.91%和0.23%；美國(guó)雞肉市場(chǎng)平均價(jià)格為2.45美元·kg-1，與10月份持平；新西蘭羊羔肉和羊肉市場(chǎng)平均價(jià)格分別為3.54和1.91美元·kg-1，其中羊羔肉市場(chǎng)價(jià)格環(huán)比下降6.09%，羊肉市場(chǎng)價(jià)格環(huán)比上漲0.58%； 歐盟豬肉市場(chǎng)平均價(jià)格為2.00美元·kg-1， 環(huán)比下降6.01%。

圖1 2016年11月國(guó)際市場(chǎng)主要飼料與畜產(chǎn)品價(jià)格

數(shù)據(jù)來(lái)源：國(guó)際市場(chǎng)商品價(jià)格網(wǎng) http://price.mofcom.gov.cn/；中國(guó)農(nóng)業(yè)信息http://www.agri.gov.cn/；雞肉 http://www.indexmundi.com/；羊羔肉、羊肉 http://www.interest.co.nz/rural；牛肉http://www.thebeefsite.com/；豬肉 http://www.thepigsite.com/；貨幣匯率：http://qq.ip138.com/hl.asp。

(蘭州大學(xué)草地農(nóng)業(yè)科技學(xué)院 王迎新 整理)

Advances of seed hormonal priming

Han Yun-hua, Wang Yan-rong, Tao Qi-bo

(State Key Laboratory of Grassland Agro-ecosystems, College of Pastoral Agriculture Science and Technology, Lanzhou University, Lanzhou 730020, China)

Hormonal priming is an important component of seed priming. It has great significance for seed germination, seedling growth, stress tolerance and yield increase. Based on the current research situation, we summarized the concept and application range of hormonal priming, illustrated the positive effects of hormonal priming on plant growth. Furthermore, biochemistry and molecular mechanism of seed hormonal priming were explained. In addition, we pointed out that hormonal concentration, priming time and drying conditions are the most important factors that constraint the priming effect. To improve the research in the future, priming mechanism should be intensively studied based on the genomics and proteome approach, and increase the use of hormonal priming in grass species.

seed priming; hormonal; stress tolerance; germination; yield

Han Yun-hua E-mail:hanyh@lzu.edu.cn

2015-12-22接受日期：2016-06-13

蘭州大學(xué)中央高?；究蒲袠I(yè)務(wù)費(fèi)專(zhuān)項(xiàng)資金(lzujbky-2015-40)

韓云華(1985-)，男，河北張家口人，講師，博士，主要從事牧草種子生產(chǎn)和種子生態(tài)研究。Email：hanyh@lzu.edu.cn

10.11829/j.issn.1001-0629.2015-0726

S330;Q945.34

A

1001-0629(2016)12-2494-09*

韓云華，王彥榮，陶奇波.種子激素引發(fā).草業(yè)科學(xué),2016,33(12)：2494-2502.

Han Y H,Wang Y R,Tao Q B.Advances of seed hormonal priming.Pratacultural Science，2016,33(12)：2494-2502.