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韓貝 孫思敏 孫偉男 楊細(xì)燕* 張獻(xiàn)龍 華中農(nóng)業(yè)大學(xué)作物遺傳改良國(guó)家重點(diǎn)實(shí)驗(yàn)室，湖北武漢 430070 摘要 Abstract 在體細(xì)胞胚胎發(fā)生過程中，通過不同激素的配合使用可以有效調(diào)控體細(xì)胞胚胎發(fā)生的各個(gè)發(fā)育階段。比如，通過調(diào)節(jié)生長(zhǎng)素(auxin)與細(xì)胞分裂素(cytokinin，CTK)比率促進(jìn)脫分化和愈傷增殖、通過添加低濃度的乙烯(ethylene)促進(jìn)體細(xì)胞胚胎發(fā)生、通過赤霉素(gibberellins，GAs)調(diào)控胚性培養(yǎng)物到球形胚的轉(zhuǎn)化、通過添加脫落酸(abscisic acid，ABA)來提高體細(xì)胞胚的質(zhì)量等。外源激素對(duì)體細(xì)胞胚胎的作用主要是通過胞外或胞內(nèi)的激素受體將外界刺激信號(hào)轉(zhuǎn)到核內(nèi)，從而調(diào)控基因的表達(dá)，啟動(dòng)發(fā)育程序。 1.1 生長(zhǎng)素信號(hào)傳導(dǎo)及體細(xì)胞胚胎發(fā)生
圖1 生長(zhǎng)素信號(hào)傳導(dǎo)及體細(xì)胞胚胎發(fā)生 Fig. 1 Auxin signaling and somatic embryogenesis 1.2 細(xì)胞分裂素信號(hào)與愈傷組織的增殖 1.3 乙烯信號(hào)與體細(xì)胞胚胎發(fā)生 理論上所有離體植物細(xì)胞在合適的外界環(huán)境下均可表現(xiàn)全能性。外界環(huán)境因素(特別是逆境因素)是體細(xì)胞胚胎發(fā)生的重要影響因素。逆境因子(包括機(jī)械損傷)在體細(xì)胞胚胎發(fā)生的幾個(gè)主要階段都起著重要的作用，很多研究者都將逆境因子的調(diào)控作為優(yōu)化體細(xì)胞胚胎發(fā)生體系的重要手段。在現(xiàn)有的誘導(dǎo)愈傷組織體系中，培養(yǎng)基多采用MS培養(yǎng)基，相對(duì)于維持植株正常生長(zhǎng)或萌發(fā)胚生根成苗的低糖、低鹽、低滲的SH培養(yǎng)基及1/2MS培養(yǎng)基來說，MS培養(yǎng)基無機(jī)鹽含量較高，微量元素種類較全，濃度也較高。另外，很多物種，包括胡蘿卜、苜蓿、煙草等，均有通過利用逆境處理來促進(jìn)體細(xì)胞胚胎形成及發(fā)育的研究，涉及到的逆境因素也多種多樣，主要有ABA處理、饑餓處理、滲透脅迫、高溫處理等[14-15]。
圖2 機(jī)械損傷通過調(diào)節(jié)因子WINs調(diào)控愈傷組織形成 Fig. 2 Mechanical damage regulates callus formation through the regulatory factor WINs 雖然體細(xì)胞胚胎發(fā)生過程受諸多外界環(huán)境因素的影響，但歸根結(jié)底是在各種因素的作用下，體細(xì)胞中某些特異的基因啟動(dòng)表達(dá)，從而使體細(xì)胞脫分化并再分化轉(zhuǎn)變?yōu)榕咝约?xì)胞。許多研究人員致力于解析體細(xì)胞胚胎發(fā)生過程的相關(guān)基因，目前已鑒定和克隆了大量與體細(xì)胞胚胎發(fā)生相關(guān)的轉(zhuǎn)錄因子基因，其中許多參與調(diào)節(jié)合子胚發(fā)生、分生組織分化和維持的轉(zhuǎn)錄因子都在體細(xì)胞胚胎發(fā)生過程中起著重要用。 3.1 核因子Y (nuclear factor Y, NF-Y) 3.2 B3-結(jié)構(gòu)域轉(zhuǎn)錄因子 3.3 AP2/ERF結(jié)構(gòu)域蛋白 3.4 同源異形域轉(zhuǎn)錄因子 4.1 類受體激酶與體細(xì)胞胚胎發(fā)生 4.2 鈣信號(hào)與體細(xì)胞胚胎發(fā)生 5.1 阿拉伯半乳糖蛋白 5.2 脂質(zhì)轉(zhuǎn)運(yùn)蛋白 6.1 DNA甲基化 6.2 組蛋白甲基化 組蛋白甲基化由組蛋白甲基轉(zhuǎn)移酶完成的。擬南芥中，組蛋白甲基轉(zhuǎn)移酶PRC2通過H3K27me3來抑制相關(guān)基因的表達(dá)，從而促進(jìn)細(xì)胞分化，反之則引起細(xì)胞脫分化，誘導(dǎo)體細(xì)胞胚胎發(fā)生[66]。在擬南芥中PRC2 (polycomb repressive complex 2)基因(CURLY LEAF，CLF和SWINGER，SWN)或(VERNALIZATION 2，VRN2和EMBRYONIC FLOWER 2，EMF2)雙突變體在莖尖脫分化形成愈傷組織，間接導(dǎo)致體細(xì)胞胚胎發(fā)生并形成異位根[67]，并且與野生型相比，PRC2的突變體在營(yíng)養(yǎng)組織中顯示出更高的體細(xì)胞胚胎誘導(dǎo)能力[68]。大部分胚性相關(guān)基因LEC1、LEC2、AGL15和BBM以及分生組織調(diào)節(jié)因子STM、WUS和WOX5等基因的染色質(zhì)區(qū)域都含有H3K27me3等甲基化位點(diǎn)[69]。而PRC1和PRC2與胚胎發(fā)生轉(zhuǎn)錄抑制因子VAL1和VAL2等互作，并通過表觀修飾抑制胚胎發(fā)生相關(guān)靶標(biāo)基因的表達(dá)從而抑制愈傷組織的形成和體細(xì)胞胚胎發(fā)生。除了H3K27me3在體細(xì)胞胚胎發(fā)生發(fā)揮功能外，在擬南芥還發(fā)現(xiàn)賴氨酸特異性去甲基酶LDL3可以在愈傷組織形成過程中特異性消除H3K4me2，進(jìn)一步使愈傷組織具有芽分化的能力[70]。 6.3 組蛋白去乙?；?/span> 6.4 miRNA的調(diào)控作用 本研究由國(guó)家重點(diǎn)研發(fā)計(jì)劃(2018YFD1000907)項(xiàng)目資助。 *通信作者: 楊細(xì)燕, E-mail: yxy@mail.hzau.edu.cn 第一作者聯(lián)系方式: E-mail: bhan_z@163.com 參考文獻(xiàn) [1] Yang X Y, Zhang X L, Yuan D J, Jin F Y, Zhang Y C, Xu J. 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Cham: Springer International Publishing, 2016. pp 11–22. 本文已在中國(guó)知網(wǎng)網(wǎng)絡(luò)首發(fā)，網(wǎng)址： https://kns.cnki.net/kcms/detail/11.1809.s.20220808.1609.002.html 期刊簡(jiǎn)介 |
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