李夢(mèng)莎, 閻秀峰

(東北林業(yè)大學(xué)鹽堿地生物資源環(huán)境研究中心, 東北油田鹽堿植被恢復(fù)與重建教育部重點(diǎn)實(shí)驗(yàn)室, 哈爾濱 150040)

1980年Ueda和Kato從苦艾(Artemisiaabsinthium)中分離并鑒定了具有促進(jìn)衰老生理活性的茉莉酸甲酯(methyl jasmonate, MeJA)[1]，從而引發(fā)了人們對(duì)茉莉酸信號(hào)分子的廣泛關(guān)注。迄今已知，具有信號(hào)分子生理功能的至少包括茉莉酸(jasmonic acid, JA)以及茉莉酸甲酯和茉莉酸-異亮氨酸復(fù)合物(jasmonoyl-isoleucine, JA-Ile)等茉莉酸衍生物，人們將它們統(tǒng)稱為茉莉酸類化合物(jasmonates, JAs)。研究表明，茉莉酸信號(hào)分子參與植物生長(zhǎng)發(fā)育眾多生理過(guò)程的調(diào)控，尤其是作為環(huán)境信號(hào)分子能有效地介導(dǎo)植物對(duì)病原菌、食草動(dòng)物及非生物脅迫等的防御反應(yīng)，誘導(dǎo)一系列防御基因的表達(dá)、防御反應(yīng)化學(xué)物質(zhì)的合成，并調(diào)節(jié)植物的“免疫”和應(yīng)激反應(yīng)。本文從環(huán)境信號(hào)分子角度介紹茉莉酸信號(hào)的啟動(dòng)、傳遞和生物學(xué)功能。

1 茉莉酸信號(hào)的啟動(dòng)

1.1 信號(hào)感知與轉(zhuǎn)導(dǎo)

最近幾十年，人們對(duì)外界生物脅迫和非生物脅迫信號(hào)如何被植物感知并啟動(dòng)茉莉酸信號(hào)的生物合成進(jìn)行了大量的研究，其中在番茄中的研究較多。1991年P(guān)earce等人在番茄(Lycopersiconesculentum)中發(fā)現(xiàn)了應(yīng)答昆蟲(chóng)食害等機(jī)械損傷的系統(tǒng)素(systemin)[2]，這是一種由18個(gè)氨基酸組成的多肽信號(hào)分子，來(lái)源于由200個(gè)氨基酸組成的前體蛋白——前系統(tǒng)素(prosystemin)[3]。番茄受到機(jī)械損傷后，前系統(tǒng)素水解為系統(tǒng)素，可通過(guò)質(zhì)外體(apoplast)運(yùn)輸?shù)狡渌?xì)胞，與細(xì)胞表面受體SR160(富含亮氨酸重復(fù)單位的蛋白)結(jié)合，最終激活茉莉酸信號(hào)途徑[4- 5]。除了創(chuàng)傷信號(hào)外，人們?cè)诜阎羞€發(fā)現(xiàn)病菌與真菌激發(fā)子誘發(fā)的寡聚糖信號(hào)，最終也激活茉莉酸信號(hào)途徑。推測(cè)寡聚糖的作用機(jī)制與系統(tǒng)素有相似的途徑，但具體的誘導(dǎo)機(jī)制還不清楚[6- 7]。

近幾年，在擬南芥(Arabidopsisthaliana)中也發(fā)現(xiàn)了與系統(tǒng)素具有相同功能的多肽——由23個(gè)氨基酸組成的AtPEP1。與系統(tǒng)素的產(chǎn)生類似，機(jī)械損傷或者病原體侵染促使前體蛋白PROPEP1(由92個(gè)氨基酸組成)水解為AtPEP1，AtPEP1與質(zhì)膜上的受體PEPR1(富含亮氨酸重復(fù)單位的酶)結(jié)合，最終激活茉莉酸信號(hào)途徑[8]。

系統(tǒng)素、AtPEP1與受體結(jié)合誘發(fā)茉莉酸信號(hào)途徑的過(guò)程比較復(fù)雜，目前知道最主要的過(guò)程是激活質(zhì)膜上的磷脂酶，之后磷脂酶作用于膜上的磷脂釋放亞麻酸(茉莉酸合成前體)[9]。不過(guò)，系統(tǒng)素激活磷脂酶活化的機(jī)制還不清楚。目前為止，人們發(fā)現(xiàn)了幾種可以被系統(tǒng)素、AtPEP1誘導(dǎo)的磷脂酶，包括番茄中的PLA2和擬南芥中的DAD1、DGL、PLD，且這幾種磷脂酶作用機(jī)制相似[10- 12]。

1.2 茉莉酸類化合物的合成

針對(duì)茉莉酸類化合物在植物體內(nèi)的合成，近一二十年人們已經(jīng)研究了多種單子葉和雙子葉植物，其中對(duì)模式植物擬南芥和番茄開(kāi)展的工作最多。到目前為止，茉莉酸類化合物的生物合成途徑已經(jīng)比較清楚，途徑中的各種酶已基本完成功能解析，并且關(guān)注茉莉酸合成途徑與其他代謝途徑關(guān)聯(lián)關(guān)系的研究逐漸增多。

關(guān)于茉莉酸類化合物生物合成的研究進(jìn)展，國(guó)內(nèi)學(xué)者已有較好的綜述[13- 14]，這里不再詳細(xì)闡述。在擬南芥中，至少存在兩條合成茉莉酸的途徑，即從亞麻酸(linolenic acid, 18:3)開(kāi)始的十八烷途徑和從十六碳三烯酸(16∶3)開(kāi)始的十六烷途徑。兩條途徑都涉及葉綠體、過(guò)氧化物酶體和細(xì)胞質(zhì)3個(gè)反應(yīng)場(chǎng)所，從不飽和脂肪酸到12-氧植物二烯酸(12-oxo-phytodienoic acid, 12-OPDA)或去甲基-含氧植物二烯酸(dn-OPDA)的合成發(fā)生在葉綠體中，而后到茉莉酸的轉(zhuǎn)化發(fā)生在過(guò)氧化物酶體中，對(duì)茉莉酸的修飾在細(xì)胞質(zhì)中完成[15]。

茉莉酸(JA)經(jīng)各種化學(xué)修飾會(huì)代謝成不同的結(jié)構(gòu)，但目前為止人們僅發(fā)現(xiàn)茉莉酸甲酯(MeJA)、茉莉酸-異亮氨酸復(fù)合物(JA-Ile)與順式-茉莉酸(cis-jasmone, CJ)這3種茉莉酸衍生物在環(huán)境誘導(dǎo)的茉莉酸信號(hào)途徑中起作用[16]。

2 茉莉酸信號(hào)的傳遞

茉莉酸類化合物在植物響應(yīng)生物與非生物脅迫的反應(yīng)中扮演著重要角色，在植物的防御過(guò)程中起著調(diào)控全局的作用。例如，在由創(chuàng)傷信號(hào)引發(fā)的防御反應(yīng)中，通常既有傷口附近的局部防御反應(yīng)，也有未受傷部位的系統(tǒng)獲得性抗性(systemic acquired resistance, SAR)，乃至相鄰植株產(chǎn)生的誘導(dǎo)防御反應(yīng)。在這些防御反應(yīng)中，人們發(fā)現(xiàn)了茉莉酸信號(hào)的短距離傳遞和遠(yuǎn)距離傳輸(圖1)[17]。

圖1 茉莉酸信號(hào)的傳遞[17]

2.1 短距離信號(hào)傳遞

當(dāng)植物受到機(jī)械損傷或昆蟲(chóng)取食后，在損傷部位會(huì)造成迅速并且短暫的JA或JA-Ile的積累，由此激活傷口周圍組織防御基因的表達(dá)，從而產(chǎn)生局部防御反應(yīng)。在局部防御反應(yīng)中，茉莉酸信號(hào)的短距離傳遞可能有兩種方式。其一，由創(chuàng)傷反應(yīng)產(chǎn)生的系統(tǒng)素作為信號(hào)物質(zhì)，通過(guò)質(zhì)外體與韌皮部傳遞到鄰近部位，激活茉莉酸級(jí)聯(lián)反應(yīng)途徑。其二，由系統(tǒng)素誘導(dǎo)產(chǎn)生的JA與JA-Ile充當(dāng)了流動(dòng)信號(hào)，被轉(zhuǎn)運(yùn)到鄰近部位進(jìn)行防御反應(yīng)[18]。

2.2 遠(yuǎn)距離信號(hào)傳輸

目前為止知道，茉莉酸信號(hào)的遠(yuǎn)距離傳輸方式有維管束傳輸與空氣傳播兩種。

2.2.1 維管束傳輸

先前，許多研究者認(rèn)為系統(tǒng)素在遠(yuǎn)距離信號(hào)傳遞的過(guò)程中發(fā)揮直接作用，是一種可移動(dòng)的信號(hào)分子。但是，用擬南芥的茉莉酸不敏感突變體jai1、系統(tǒng)素不敏感突變體spr1、茉莉酸缺失突變體spr2與acx1A等開(kāi)展的一系列嫁接實(shí)驗(yàn)證明，系統(tǒng)素只是在局部起到信號(hào)放大與誘發(fā)茉莉酸類化合物合成的作用，之后，茉莉酸類化合物JA-Ile在植物體內(nèi)傳輸[19]。Thorpe等則通過(guò)同位素標(biāo)記實(shí)驗(yàn)證明，MeJA可以在維管束中的韌皮部和木質(zhì)部轉(zhuǎn)移[20]。

一些工作也表明，茉莉酸類化合物并不是簡(jiǎn)單地沿維管束運(yùn)輸，而是在運(yùn)輸過(guò)程中伴隨著茉莉酸類化合物的重新合成。在番茄維管束的伴胞和篩管復(fù)合體(companion cell-sieveelement complex, CC-SE)中也發(fā)現(xiàn)了各種JA合成酶(如LOX、AOS等)的定位[21]，且韌皮部中的篩分子有形成JA前體OPDA的能力[22]。

近些年Koo等[23]通過(guò)實(shí)驗(yàn)發(fā)現(xiàn)，傷誘導(dǎo)導(dǎo)致的全身性JA、JA-Ile的產(chǎn)生并非全部由受傷部位轉(zhuǎn)移而來(lái)，至少有一部分是重新合成的，并在未受傷部位級(jí)聯(lián)循環(huán)產(chǎn)生更多的JA-Ile。Wang等也證實(shí)了這一結(jié)果[24]。

2.2.2 空氣傳播

實(shí)驗(yàn)發(fā)現(xiàn)，番茄韌皮部信號(hào)的流動(dòng)速度為每小時(shí)1—5 cm[25]，但機(jī)械損傷后15 min內(nèi)就可在整株檢測(cè)到JA與JA-Ile的積累[26]。20世紀(jì)90年代Malone等的環(huán)割實(shí)驗(yàn)也證明，即使維管束傳輸被阻斷了，遠(yuǎn)端葉片中也存在迅速且強(qiáng)烈的防御基因表達(dá)[26]。大量的事實(shí)表明，除了維管束傳輸以外，茉莉酸信號(hào)還存在其他的遠(yuǎn)距離傳播途徑。JA可被甲基化形成MeJA。相對(duì)于沒(méi)有載體輔助就難以穿透細(xì)胞膜的JA，MeJA容易透過(guò)細(xì)胞膜且有很強(qiáng)的揮發(fā)性，可通過(guò)空氣傳播擴(kuò)散到遠(yuǎn)距離葉片和相鄰的植株而發(fā)揮作用[27]。在擬南芥[19]、煙草(Nicotianatabacum)[28]、利馬豆(Phaseoluslunatus)[29]、山艾樹(shù)(Artemisiakawakamii)[30]等植物中已經(jīng)證實(shí)，MeJA可以在同一植株的損傷與未損傷葉片之間或相鄰植物間通過(guò)空氣傳播。

3 茉莉酸信號(hào)的效應(yīng)

3.1 茉莉酸信號(hào)的受體

人們對(duì)茉莉酸信號(hào)受體的認(rèn)識(shí)經(jīng)歷了復(fù)雜的過(guò)程。1994年Feys最先確認(rèn)擬南芥的COI1蛋白是茉莉酸敏感蛋白[31]，進(jìn)一步發(fā)現(xiàn)擬南芥coi1突變體喪失了對(duì)茉莉酸的所有反應(yīng)[32]，證實(shí)了這一觀點(diǎn)。COI1是一種泛素連接酶E3的組成部分，SCF型泛素連接酶E3包括S、C、F 3個(gè)組分，其家族所有成員的S(SKP1蛋白)和C(Cullin蛋白)組分都相同，區(qū)分標(biāo)志是F組分(F-box)，F(xiàn)-box蛋白為COI1的SCF型泛素連接酶E3記為SCFCOI1[33]。COI1蛋白的發(fā)現(xiàn)對(duì)于茉莉酸信號(hào)途徑的研究有著重要意義。

人們?cè)欢纫詾镃OI1就是茉莉酸信號(hào)在細(xì)胞內(nèi)的受體，直到JAZ蛋白家族的發(fā)現(xiàn)使人們對(duì)茉莉酸信號(hào)轉(zhuǎn)導(dǎo)途徑有了新的認(rèn)識(shí)。2007年Thines在茉莉酸合成突變體opr3中發(fā)現(xiàn)了8個(gè)JAZ蛋白[34]。在擬南芥中有12個(gè)JAZ蛋白，它們都有Jas和ZIM兩個(gè)保守結(jié)構(gòu)域[35]。JAZ蛋白通過(guò)Jas結(jié)構(gòu)域與COI1相作用，通過(guò)ZIM結(jié)構(gòu)域與MYC2發(fā)生作用[36]。因此，許多研究者認(rèn)為，JAZ蛋白是COI1的靶蛋白，JAZ蛋白的降解是解除JA途徑抑制的關(guān)鍵步驟。然而，2010年Sheard等通過(guò)晶體結(jié)構(gòu)的分析又對(duì)茉莉酸受體提出了不同觀點(diǎn)，并證實(shí)COI1-JAZ復(fù)合物是茉莉酸的高親和受體，即COI1與JAZ是茉莉酸信號(hào)的共同受體[37]。目前認(rèn)為，當(dāng)植物受到外界環(huán)境刺激后，生成的JA-Ile與COI1-JAZ復(fù)合體直接結(jié)合，形成的復(fù)合物轉(zhuǎn)移到26s蛋白酶體后被降解，同時(shí)激活了下游基因的轉(zhuǎn)錄(圖2)。

圖2 茉莉酸信號(hào)的轉(zhuǎn)導(dǎo)通路

3.2 茉莉酸信號(hào)調(diào)控的轉(zhuǎn)錄因子

從圖2看，茉莉酸信號(hào)通過(guò)JAZ蛋白直接啟動(dòng)的是MYC類轉(zhuǎn)錄因子，不過(guò)最新的研究發(fā)現(xiàn)，MYB類轉(zhuǎn)錄因子也可以通過(guò)與JAZ蛋白的結(jié)合而被茉莉酸信號(hào)激活。此外，受茉莉酸信號(hào)調(diào)控的還有NAC、ERF、WRKY等幾類轉(zhuǎn)錄因子。茉莉酸信號(hào)通過(guò)這些轉(zhuǎn)錄因子進(jìn)而調(diào)節(jié)眾多基因的表達(dá)、影響植物的生長(zhǎng)發(fā)育過(guò)程特別是植物對(duì)環(huán)境的響應(yīng)與適應(yīng)。也有研究表明，茉莉酸信號(hào)還能誘發(fā)MAP級(jí)聯(lián)反應(yīng)途徑[38]、鈣離子通道[39]，以及與乙烯、水楊酸、脫落酸等信號(hào)分子互作調(diào)控植物生命活動(dòng)的眾多過(guò)程[40]。

3.2.1 MYC類轉(zhuǎn)錄因子

在MYC類轉(zhuǎn)錄因子中，MYC2蛋白是大家所熟知的調(diào)節(jié)蛋白。MYC2屬于bHLH(堿性螺旋-環(huán)-螺旋)類轉(zhuǎn)錄因子，由JIN1基因編碼。JAZ蛋白家族中的大多數(shù)成員都可與MYC2相互作用[41]。很長(zhǎng)時(shí)間以來(lái)，人們認(rèn)為只有MYC2蛋白可以與JAZ蛋白直接作用，2011年Fernandez-Calvo等鑒定出另外兩種bHLH類蛋白——MYC3與MYC4，發(fā)現(xiàn)它們也可以與JAZ蛋白相互作用，并與MYC2功能疊加[42]，不過(guò)這3種MYC蛋白在茉莉酸信號(hào)途徑中的分工以及它們相互之間的作用機(jī)制還不是很清楚。

3.2.2 MYB類轉(zhuǎn)錄因子

MYB類轉(zhuǎn)錄因子屬于R2R3-MYB轉(zhuǎn)錄因子家族，在植物應(yīng)答環(huán)境過(guò)程中起著重要的調(diào)節(jié)作用。Dombrecht等通過(guò)MYC2突變體jin1證明，在茉莉酸處理的情況下，MYC2可以通過(guò)調(diào)節(jié)MYB51和MYB34的表達(dá)量達(dá)到負(fù)調(diào)節(jié)色氨酸及吲哚族芥子油苷合成的作用，由此推斷MYB轉(zhuǎn)錄因子在MYC2的下游起作用[41]。但近期的研究發(fā)現(xiàn)，有一些MYB類轉(zhuǎn)錄因子可以與JAZ蛋白結(jié)合，如MYB75、Glabra1、MYB21和MYB24，從而影響植物花青素積累、毛狀體形成以及雄蕊成熟等發(fā)育過(guò)程。這些MYB類轉(zhuǎn)錄因子與JAZ作用的機(jī)制與MYC2相似，都是通過(guò)JAZ蛋白的降解來(lái)啟動(dòng)下游基因的轉(zhuǎn)錄[43- 44]。

3.2.3 NAC類轉(zhuǎn)錄因子

NAC類轉(zhuǎn)錄因子在生物與非生物脅迫時(shí)也被茉莉酸信號(hào)誘導(dǎo)，并且在植物的生長(zhǎng)發(fā)育過(guò)程中起到重要作用。擬南芥NAC家族中的ATAF1、ATAF2轉(zhuǎn)錄因子都受到茉莉酸信號(hào)的誘導(dǎo)，茉莉酸信號(hào)通過(guò)對(duì)ATAF1、ATAF2的調(diào)節(jié)提高了植物對(duì)干旱、鹽脅迫、灰霉病菌及其他一些病原體的抵抗能力，同時(shí)對(duì)植物的氧化應(yīng)激反應(yīng)、開(kāi)花、角果發(fā)育有著重要的調(diào)節(jié)作用。此外，ATAF1與ATAF2還調(diào)節(jié)著茉莉酸與水楊酸信號(hào)途徑之間的相互作用[45- 46]。擬南芥中NAC家族中的另2個(gè)轉(zhuǎn)錄因子ANAC019與ANAC055也存在于MYC2轉(zhuǎn)錄因子的下游，并與MYC2起平行作用，可促進(jìn)種子萌發(fā)、細(xì)胞分裂、細(xì)胞次生壁的合成以及影響脫落酸信號(hào)[47]。

3.2.4 ERF類轉(zhuǎn)錄因子

在擬南芥中，ERF轉(zhuǎn)錄因子家族有122個(gè)成員[48]。近幾年基因水平的微陣列實(shí)驗(yàn)證實(shí)茉莉酸信號(hào)可以誘導(dǎo)許多EFR轉(zhuǎn)錄因子的轉(zhuǎn)錄。在擬南芥中，EFR1、EFR2、EFR4轉(zhuǎn)錄因子位于MYC2轉(zhuǎn)錄因子的下游，同樣受到茉莉酸信號(hào)的誘導(dǎo)，但EFR4卻與EFR1、EFR2相互抑制[49- 50]。此外，茉莉酸信號(hào)通過(guò)誘導(dǎo)ORA59轉(zhuǎn)錄因子增加了對(duì)灰霉病的抵抗性[51]。在煙草中，OPBP1轉(zhuǎn)錄因子也受到茉莉酸信號(hào)的誘導(dǎo)[52]。由于ERF轉(zhuǎn)錄因子是一類乙烯應(yīng)答因子，因而也介導(dǎo)著茉莉酸信號(hào)途徑與乙烯信號(hào)途徑之間的相互作用[48]。

3.2.5 WRKY類轉(zhuǎn)錄因子

WRKY類轉(zhuǎn)錄因子在植物發(fā)育、衰老及應(yīng)對(duì)環(huán)境脅迫過(guò)程中具有重要調(diào)控作用。擬南芥中有74個(gè)WRKY家族成員[53]。近年來(lái)的研究表明，有一些WRKY類轉(zhuǎn)錄因子是受茉莉酸信號(hào)調(diào)控的，如擬南芥中的WRKY70[54]、WRKY18[55]、WRKY33[56]、WRKY53[57]、WRKY62[58]等，它們多與植物的防御功能相關(guān)。通過(guò)基因敲除、超表達(dá)等實(shí)驗(yàn)推測(cè)它們處于COI的下游，且受到COI蛋白的抑制作用。在煙草中，2個(gè)WRKY類轉(zhuǎn)錄因子——NaWRKY3和NaWRKY6可以調(diào)節(jié)茉莉酸生物合成相關(guān)基因(LOX、AOS、AOC、OPR)的表達(dá)從而提高JA和JA-Ile的水平，對(duì)茉莉酸信號(hào)起到反饋?zhàn)饔肹59]。此外，WRKY蛋白也是MAP激酶的作用目標(biāo)[53]。

4 茉莉酸信號(hào)參與的生物學(xué)過(guò)程

4.1 受茉莉酸信號(hào)影響的環(huán)境應(yīng)答

大量研究表明，茉莉酸類化合物是與抗性密切相關(guān)的植物生長(zhǎng)物質(zhì)，作為內(nèi)源信號(hào)分子參與植物在干旱、鹽脅迫、低溫等條件下的抗逆反應(yīng)。

4.1.1 光

植物受光的影響并觀察到茉莉酸信號(hào)變化的主要體現(xiàn)在兩個(gè)方面，即植物的光形態(tài)建成和UV-B對(duì)植物的傷害。在擬南芥和水稻中都觀察到紅光/遠(yuǎn)紅光介導(dǎo)的光形態(tài)建成伴隨著茉莉酸信號(hào)的變化，在藍(lán)光介導(dǎo)的擬南芥光形態(tài)建成中也發(fā)現(xiàn)茉莉酸信號(hào)途徑的參與[60- 61]。UV-B輻射的增強(qiáng)能夠誘導(dǎo)煙草屬和曼陀羅屬植物體內(nèi)茉莉酸的生物合成從而啟動(dòng)茉莉酸信號(hào)途徑，并且這些過(guò)程與植物的化學(xué)防御相關(guān)聯(lián)[62- 63]。

4.1.2 溫度

Zhao等[64]對(duì)香蕉(Musaacuminata)的研究發(fā)現(xiàn)，低溫處理后內(nèi)源茉莉酸含量會(huì)略有降低，但變化不顯著。不過(guò)，在寒冷中MYC基因會(huì)對(duì)外源MeJA快速反應(yīng)，并在體內(nèi)重新合成大量的茉莉酸，抵御寒害。此外，對(duì)番茄(Lycopersiconesculentum)[65]、石榴(Punicagranatum)[66]、枇杷(Eribotryajaponica)[67]、芒果(Mangiferaindica)[68]、番石榴(Psidiumguajava)[69]等植物的研究都表明，外源MeJA處理能夠誘導(dǎo)熱激蛋白家族轉(zhuǎn)錄、增加抗氧化劑合成、降低脂氧合酶活性從而增加植物抵御寒害(零上低溫)的能力，說(shuō)明茉莉酸信號(hào)途徑參與了植物對(duì)低溫的響應(yīng)與適應(yīng)過(guò)程。最近對(duì)香蕉的研究發(fā)現(xiàn)，冷儲(chǔ)存后MeJA可明顯誘導(dǎo)MYC家族轉(zhuǎn)錄因子以及許多冷敏感基因(MaCBF1,MaCBF2,MaCOR1,MaKIN2,MaRD2,MaRD5等)的表達(dá)，由此減緩寒害對(duì)植物的傷害[64]。

目前，有關(guān)茉莉酸信號(hào)參與植物抗熱性的報(bào)道相對(duì)較少，但在紅掌(Spathiphyllumfloribundum)幼苗和蝴蝶蘭(Phalaenopsisamabilis)中已發(fā)現(xiàn)外源MeJA處理能提高它們的耐熱性[70]。

4.1.3 水

已有不少研究發(fā)現(xiàn)干旱脅迫中有茉莉酸信號(hào)途徑的參與。在擬南芥[71]、柑橘(Citrusparadisi×Poncirustrifoliata)[72]中發(fā)現(xiàn)，干旱脅迫后內(nèi)源茉莉酸含量的增加是快速、瞬時(shí)的，而后隨著脅迫時(shí)間的延長(zhǎng)逐漸降低到基礎(chǔ)水平。另一方面，外源茉莉酸施加也可以有效地緩解干旱對(duì)植物造成的損傷。早期潘瑞熾等用MeJA處理提高了花生(Arachishypogaea)幼苗的抗旱性[73]，在水稻(Oryzasativa)[74]、大豆(Glycinemax)[75]、花椰菜(Brassicaoleracea)[76]中也觀察到MeJA處理可以通過(guò)調(diào)整新陳代謝來(lái)抵御干旱脅迫。對(duì)蠶豆(Viciafaba)和大麥(Hordeumvulgare)的研究表明，MeJA可能是通過(guò)K+通道調(diào)控氣孔運(yùn)動(dòng)、從而提高植株抗旱能力的[77- 78]。

4.1.4 鹽分

在擬南芥[79]、番茄[80]、大麥[81]等植物中都發(fā)現(xiàn)，鹽處理后內(nèi)源茉莉酸含量明顯增加。而且，在鹽敏感的植物中茉莉酸含量增加迅速而又持久，而耐鹽植物中茉莉酸含量的變化則不明顯[81]。人們也觀察到，外源茉莉酸可以提高大豆[82]、胡椒(Capsicumannuum)[83]和葡萄藤(Rupestrisriparia)[84]等植物對(duì)鹽脅迫的抵御能力。

4.1.5CO2濃度

Allhornl等發(fā)現(xiàn)，高濃度CO2(500、700、1000 μmol/mol)環(huán)境導(dǎo)致利馬豆(Phaseoluslunatus)向空氣中釋放茉莉酸類化合物(MeJA和cis-JA)的量增加[85]。不過(guò)，有關(guān)CO2作用下植物體內(nèi)茉莉酸信號(hào)途徑的研究還比較少。

4.1.6 臭氧

經(jīng)臭氧處理后野生型擬南芥的內(nèi)源茉莉酸含量明顯增加，而用臭氧敏感突變體rcd1(radical- induced cell death 1)[86]、oji1(ozone-sensitive and jasmonate-insensitive 1)[87]以及茉莉酸信號(hào)突變體jar1[88]開(kāi)展的實(shí)驗(yàn)表明，外源MeJA可以抑制由臭氧導(dǎo)致的細(xì)胞程序性死亡的傳播，而阻斷茉莉酸信號(hào)則會(huì)使植物對(duì)臭氧產(chǎn)生更強(qiáng)烈的過(guò)敏反應(yīng)。施加外源MeJA也導(dǎo)致雜交楊樹(shù)(Populusmaximowizii×P.trichocarpa)[89]、番茄[88]對(duì)臭氧的敏感性降低。但是，最近對(duì)棉花的研究顯示，只有在高濃度(685體積分?jǐn)?shù))臭氧下，MeJA才顯示出對(duì)臭氧損傷擴(kuò)散的抑制，并伴隨著與乙烯的拮抗作用[90]。

4.2 基因芯片和蛋白組學(xué)研究結(jié)果

近年發(fā)展起來(lái)的基因芯片和蛋白組學(xué)等系統(tǒng)生物學(xué)研究手段，可以從全局角度在基因和蛋白質(zhì)水平上檢測(cè)植物各種生理過(guò)程的反應(yīng)和變化，從而分析各代謝途徑間的相互關(guān)系。

Jung等采用基因芯片技術(shù)在MeJA處理的擬南芥中鑒定了137個(gè)表達(dá)水平發(fā)生變化的基因。其中有74個(gè)基因上調(diào)表達(dá)，包括茉莉酸生物合成相關(guān)基因、各種防御基因(如pdf1.2、黑芥子酶結(jié)合蛋白的編碼基因)、氧化應(yīng)激基因(氧化酶類、谷胱甘肽轉(zhuǎn)移酶、cyp450家族的編碼基因)、衰老相關(guān)基因、細(xì)胞壁修飾相關(guān)基因、激素代謝相關(guān)酶(如負(fù)責(zé)乙烯合成的ACC氧化酶)基因，以及涉及儲(chǔ)藏、信號(hào)轉(zhuǎn)導(dǎo)、初生和次生代謝相關(guān)的基因。63個(gè)基因下調(diào)表達(dá)，包括光合作用相關(guān)基因(Rubisco酶基因、葉綠素蛋白基因、早期光誘導(dǎo)蛋白基因)、寒冷調(diào)節(jié)基因、干旱調(diào)節(jié)基因、防御反應(yīng)相關(guān)基因、植物生長(zhǎng)發(fā)育相關(guān)基因、細(xì)胞壁修飾相關(guān)基因和一些其他的未知功能基因等[91]。

Chen等通過(guò)蛋白組學(xué)方法比較了擬南芥中MeJA處理前后各類蛋白含量變化，發(fā)現(xiàn)了186個(gè)差異表達(dá)的蛋白。這些蛋白涉及到植物的光合作用、碳水化合物代謝、激素代謝、次生代謝、產(chǎn)物運(yùn)輸、脅迫和防御以及基因轉(zhuǎn)錄等[92]。

基因芯片和蛋白組學(xué)的研究結(jié)果進(jìn)一步證實(shí)了茉莉酸信號(hào)途徑所涉及的植物生理代謝過(guò)程的廣泛性。

5 展望

近年來(lái)，關(guān)于植物環(huán)境信號(hào)分子——茉莉酸的研究取得了較大的進(jìn)展，但由于植物對(duì)環(huán)境的反應(yīng)是一個(gè)多種信號(hào)相互作用的復(fù)雜體系，茉莉酸信號(hào)在植物-環(huán)境關(guān)系中的作用機(jī)制還有待進(jìn)一步闡明。

有關(guān)茉莉酸信號(hào)的合成、傳遞等一系列轉(zhuǎn)導(dǎo)途徑已經(jīng)較為清晰，但是對(duì)于不同的環(huán)境信號(hào)如何被植物感知并啟動(dòng)茉莉酸合成的研究還不系統(tǒng)。由于細(xì)胞膜上受體以及激酶種類較多，且不同的生物以及非生物信號(hào)會(huì)刺激不同的酶活化，同時(shí)伴隨著鈣離子通道、鉀離子通道開(kāi)放等一系列復(fù)雜的反應(yīng)，所以環(huán)境信號(hào)的感知尚有很大的研究空間。

茉莉酸受體的研究已取得較大進(jìn)展，茉莉酸信號(hào)轉(zhuǎn)導(dǎo)模式也已經(jīng)建立，但這一調(diào)控過(guò)程中還有許多細(xì)節(jié)不甚清楚。如，茉莉酸信號(hào)如何誘導(dǎo)下游轉(zhuǎn)錄因子轉(zhuǎn)錄，以及這些下游轉(zhuǎn)錄因子間如何相互作用共同調(diào)控植物生長(zhǎng)發(fā)育等，都是一系列有待解決的問(wèn)題。

隨著對(duì)激素信號(hào)網(wǎng)絡(luò)作用機(jī)制研究的不斷深入，人們發(fā)現(xiàn)水楊酸、乙烯、生長(zhǎng)素等植物激素會(huì)與茉莉酸信號(hào)相互作用，以調(diào)節(jié)植物適應(yīng)環(huán)境。目前對(duì)植物接受環(huán)境信號(hào)后的復(fù)雜調(diào)控網(wǎng)絡(luò)及生物代謝過(guò)程的認(rèn)識(shí)還很有限。

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