芸苔素内酯调控植物生长发育及抗逆性的研究进展

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周扬锟等

人[84]的研究则发现了BR的应用使得油菜子叶镉的积累降低了14%。Hayat等人[85]报道了BR的应用改善了镉胁迫下芸苔属植物的碳酸酐酶活性，叶绿素含量，净光合速率以及渗透调节。 3.4.2. 铜胁迫

高浓度的铜诱导通过哈伯–韦斯反应和芬顿反应产生大量自由基，从而造成氧化应激[86]，从而导致一系列氧化损伤。Fariduddin等人[87]报道了BR显著改善了铜胁迫下芸苔属植物的形态，增加了生物量的积累。Fariduddin等人[88]的另一项研究则发现黄瓜在盐胁迫和铜胁迫共同作用下，生物量的积累、叶绿素含量、碳酸酐酶活性、净光合速率以及PSII初级光化学的最大量子产量均显著下降，而在应用了BR后这些指标得到改善，除此以外植物多种抗氧化酶活性和脯氨酸含量同时上升，增强了植物的抗逆性。 3.4.3. 铝胁迫

Pereira等人[89]的实验发现了铝对植物生长，以及一些特定的酶，如δ-氨基乙酰丙酸脱水酶(ALA-D)活性的抑制作用。除此之外，铝还会干扰根尖和侧根的细胞分裂、降低根系呼吸和酶活性、干扰植物营养平衡、且能通过增强DNA双链的刚性来抑制DNA复制[90]。Ali等[91]的研究表明BR的应用降低了铝胁迫对绿豆幼苗的毒性，改善了幼苗生长、光合以及其它生理过程。Madhan等人[92]的研究发现BR的应用促进了铝胁迫下木豆种子的萌发和幼苗的生长。幼苗体内过氧化氢酶、过氧化物酶、超氧化物歧化酶和抗坏血酸过氧化物酶等抗氧化酶的活性也随着BR的应用而提高，脯氨酸等渗透调节物质水平也同时提高，维持了幼苗渗透平衡。 3.4.4. 镍胁迫

过量的镍对植物的光合作用、蒸散作用等生理过程以及抗氧化酶活性以及渗透调节物质水平产生影响，还会产生活性氧并造成脂质过氧化[93] [94]。Kanwar等人[95]报道了外源BR抑制了芥菜根茎对镍的吸收，恢复了受镍胁迫的芥菜生长。他们的研究结果还表明外源BR可提高植物体内的SOD、CAT、POD、APOX等氧化酶活性，改善植物形态和生理过程。Soares等人[96]则报道BR降低了镍在植物根中的积累，并且改善了渗透调节物质水平，提高了植物光合色素含量以及Rubisco活性。 3.4.5. 铬胁迫

Parr和Fred [97]的研究表明镉离子抑制了矮菜豆的萌芽。与其它重金属一样，镉毒性也会产生大量的自由基和活性氧，从而抑制植物生长[98]。Anket等人[99]研究发现，BR的应用显著减少了水稻中铬的积累，改善了水稻的生长发育。Arora等人[100]发现BR调节了抗氧化剂和蛋白质的生物合成，从而减轻铬胁迫。 3.4.6. 锌胁迫

锌浓度过高会导致光合色素分解，使叶片黄化，抑制根生长和发生，同时产生活性氧造成氧化损伤并打破营养平衡，抑制植物生长[101] [102]。Ramakrishna和Rao [103]报道了应用BR可以提高锌胁迫下萝卜幼苗的抗氧化酶活性，从而减轻活性氧对植物的损伤，同时幼苗体内的脯氨酸水平上升，幼苗抗逆性得到提高。它们的研究还指出应用了BR后MAD含量下降，脂氧合酶活性和电解液泄漏得到改善。

4. 总结与展望

BR作为一种活性的信号化合物在不同的代谢和生理过程中，改善植物胁迫下的生长发育，在植物生长和抗逆性方面有重要的作用。BR通过激活多种抗氧化酶，提高抗氧化物含量来减少活性氧对植物的损伤。不仅如此，BR也能保护光合色素的超微结构不被降解，从而增加光合作用和其他叶片的气交换。应用BR还可以增加各类渗透调节物质的积累，特别是脯氨酸，以调节胁迫下植物的渗透势。在不同的应

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用方法中，通过种子处理和叶面喷雾应用BR比根应用更方便有效。此外，也有研究认为BR具有低浓度促进生长，高浓度抑制生长的双重性。BR对于不同基因表达水平上具有调节作用，但是相关研究较少，还有研究表明外源芸苔素内酯影响植物体内多种激素水平，表明芸苔素内酯与其它植物激素间存在相互作用关系，但其机制尚不明确，也还需要深入研究。

基金项目

浙江省重点研发计划项目“药食同源植物黄精品质提升关键技术研究及功能食品开发”(2020C02039)。

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