水稻分蘖芽萌发与休眠相互转换的激素学机制
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摘要
分蘖是水稻的重要农艺性状之一,调控分蘖发生是塑造高质量水稻群体的一个重要手段。本研究采用水培、盆栽、大田等试验方式,通过氮素、外源激素和去穗等处理,构建了一系列水稻分蘖芽萌发与休眠相互转换的材料,研究了分蘖节分蘖芽和伸长节分蘖芽萌发与休眠转换过程中各种内源激素含量和相关基因表达的变化,探讨了外源激素对分蘖相关基因表达、内源激素变化和分蘖芽萌发与休眠相互转换的影响,分析了分蘖芽萌发与休眠转换过程中相关基因表达与激素的关系,从分蘖芽的萌发→休眠、休眠→萌发正反两个方面深入研究了水稻分蘖芽萌发与休眠相互转换的激素学机制。主要研究结果如下：
(1)内源CTK和IAA参与了对水稻分蘖芽萌发与休眠转换的调控,其中CTK是直接因子,IAA是间接因子。总体来讲,CTK促进分蘖芽萌发,IAA抑制分蘖芽萌发,但水稻分蘖芽萌发与休眠的转换不仅仅受到CTK和IAA绝对浓度的影响,而是受到IAA和CTK平衡的调控,当IAA/CTK值升高时,分蘖芽由萌发向休眠转换；当IAA/CTK值降低时,分蘖芽由休眠向萌发转换。氮素、外源激素、去穗等农艺措施对水稻分蘖芽萌发与休眠的影响主要通过调控CTK和IAA两种内源激素含量及其平衡实现。外源低浓度氮通过抑制CTK含量的提高进而提高IAA/CTK值抑制分蘖芽萌发,而高浓度氮和6-BA通过提高CTK含量进而降低IAA/CTK值促进分蘖芽萌发；去穗通过提高CTK含量、降低IAA含量进而降低IAA/CTK值促进分蘖芽萌发；外源GA3和IAA通过提高IAA含量、降低CTK含量进而提高IAA/CTK值抑制分蘖芽萌发；而外源NAA通过降低CTK含量进而提高IAA/CTK值抑制分蘖芽萌发。除CTK和IAA外,ABA也参与对水稻分蘖芽生长的调控,但在水稻分蘖芽萌发与休眠的转换过程中未起主导作用。
(2)IAA对水稻分蘖芽萌发与休眠的调控具有明显的位置效应,而且IAA的位置效应与其极性运输特性密切相关。分蘖期水稻基部节间不伸长,分蘖节各节密集在一处,地上部极性运输较弱,向各叶位叶涂抹的IAA均能通过极性运输运至节部通过提高IAA含量、降低CTK含量进而提高IAA/CTK值抑制分蘖节分蘖芽萌发,而根部施用的IAA则无法通过极性运输运输至节部位,所以根部施用IAA对分蘖节分蘖芽的萌发无抑制作用。齐穗后,水稻伸长节间伸长,伸长节彼此分开,地上部极性运输较强。向伸长节分蘖芽上部叶或着生叶涂抹的IAA能够通过极性运输运至节部通过提高IAA含量、降低CTK含量进而提高IAA/CTK值抑制伸长节分蘖芽萌发,而向伸长节分蘖芽下部叶涂抹的IAA则无法通过极性运输运至节部,所以向伸长节分蘖芽下部叶涂抹IAA对伸长节分蘖芽的萌发无抑制作用。与IAA不同,CTK对水稻分蘖芽萌发与休眠的调控没有明显的位置效应,向不同叶位叶和根部涂抹6-BA均能促进分蘖芽的萌发。
(3)调控水稻分蘖芽萌发与休眠的CTK在节部合成,然后运输至芽,通过调控芽中OsTB1等基因表达来调控水稻分蘖芽生长；外界因素对分蘖芽萌发与休眠的影响主要通过调控节部位CTK的合成进行；除CTK之外,外源IAA等调控措施还可能通过调控OsD3、OsD10和OsD27等基因的表达调控水稻分蘖芽生长。这表明,在水稻体内存在不止一条途径参与调节分蘖芽的萌发与休眠。
Tillering is an important agronomic trait of rice, and the tillering regulation is an effective means for the establishment of excellent population. In the present study, we conducted a series of experiments in the nutrient solution, pot and field, and external nitrogen, external hormones and panicle removal were used to construct a series of rice population which were difference for the tiller buds growth. The hormonal changes and the expression levels of the genes related to rice tiller buds growth were emphatically measured, and the objects of the present study were to investigate the relationship of hormones and genes during the transformation between dormancy and germination of rice tiller buds, and the mechanism of hormonal regulation of the transformation between dormancy and germination of the tiller buds. The main results of the study are listed below.
(1) The endogenous CTK and IAA play important roles in regulating the tiller buds growth. The CTK is the direct regulator for tiller buds growth, and the IAA is the indirect factor. In general, CTK promoted the growth of rice tiller buds, and IAA inhibited that. However, the transformation of dormancy and germination of tiller bus was not only regulated by the contents of IAA and CTK, but also regulated by the balance of them. The high IAA/CTK ratio promoted the transformation from germination to dormancy of the tiller bus, however, the low IAA/CTK ratio promoted the transformation from dormancy to germination of the tiller buds. The agronomic measures such as nitrogen, hormones and panicle removal all regulated the growth of tiller buds by regulated the contents and balance of the IAA and CTK. The low nitrogen inhibited the growth of tiller buds by inhibited the increasing of CTK and increased the ratio of IAA/CTK, and the promoting effect of high nitrogen and 6-BA on the tiller buds growth was significantly related to the increasing of the CTK contents and the decreasing of the ratio of IAA/CTK. The panicle removal increased the CTK contents and decreased the IAA contents, and then decreased the ratio of IAA/CTK and by this to promote the growth of tiller buds. External application of GA3 and IAA decreased the CTK contents and increased the IAA contents, and then increased the ratio of IAA/CTK and by this to inhibit the growth of tiller buds. The external NAA inhibited the growth of tiller buds by inhibited the increasing of CTK and increased the ratio of IAA/CTK. Besides IAA and CTK, ABA also regulated the growth of rice tiller buds, but ABA may not be the determining factor for the tiller buds growth.
(2) IAA has a notable positional effect on the regulation of rice tiller bud growth, and the positional effect of IAA was closely related to its polar transport. In tillering stage, the basal internodes of rice are not elongated, and this structural characteristic may lead to a deficiency in polar transport of IAA in the aboveground organs. IAA applied to different leaves all can transport to nodes through polar transport, and then decreased the CTK content and increased the ratio of IAA/CTK, and by this to inhibited the growth of tiller buds. However, the IAA applied to root cannot transport to nodes by polar transport, this lead to that applied IAA to roots has no significant effect on tiller buds growth. Contrary to the basal internodes, the upper internodes elongated after heading and that structural characteristic separate its nodes from one another and lead to an enhance in polar transport of IAA in the aboveground organs. IAA applied to the leaves above the buds or the leaves at which the buds locate can be transported to the node. IAA can then inhibit the increase in CTK levels and increased the ratio of IAA/CTK, thus inhibiting the growth of the bud. However, IAA applied to the leaf below the bud cannot be transported to the node by polar transport; this may be the main reason why applied IAA to the leaf below the bud could not inhibit the growth of the bud. In contrast to IAA, CTK had no positional effect on rice tiller bud growth. Application of 6-BA to different leaves and roots significantly promoted the growth of rice tiller buds at both unelongated and elongated nodes.
(3) The CTK which promotes the germination of rice tiller buds is biosynthesis mainly in the nodes and delivery to tiller buds, and then regulated the growth of tiller buds by regulated the expression of OsTBl. The agronomic measures such as external applied IAA regulated the growth of tiller buds by regulated the biosynthesis of CTK in nodes. Besides CTK, the external IAA also might have regulated the expression level of OsD3, OsD10 and OsD27 in tiller buds and by this to regulating the tiller bud growth. These results suggested that the rice have more than one ways to regulate the growth of tiller buds.
(1)内源CTK和IAA参与了对水稻分蘖芽萌发与休眠转换的调控,其中CTK是直接因子,IAA是间接因子。总体来讲,CTK促进分蘖芽萌发,IAA抑制分蘖芽萌发,但水稻分蘖芽萌发与休眠的转换不仅仅受到CTK和IAA绝对浓度的影响,而是受到IAA和CTK平衡的调控,当IAA/CTK值升高时,分蘖芽由萌发向休眠转换；当IAA/CTK值降低时,分蘖芽由休眠向萌发转换。氮素、外源激素、去穗等农艺措施对水稻分蘖芽萌发与休眠的影响主要通过调控CTK和IAA两种内源激素含量及其平衡实现。外源低浓度氮通过抑制CTK含量的提高进而提高IAA/CTK值抑制分蘖芽萌发,而高浓度氮和6-BA通过提高CTK含量进而降低IAA/CTK值促进分蘖芽萌发；去穗通过提高CTK含量、降低IAA含量进而降低IAA/CTK值促进分蘖芽萌发；外源GA3和IAA通过提高IAA含量、降低CTK含量进而提高IAA/CTK值抑制分蘖芽萌发；而外源NAA通过降低CTK含量进而提高IAA/CTK值抑制分蘖芽萌发。除CTK和IAA外,ABA也参与对水稻分蘖芽生长的调控,但在水稻分蘖芽萌发与休眠的转换过程中未起主导作用。
(2)IAA对水稻分蘖芽萌发与休眠的调控具有明显的位置效应,而且IAA的位置效应与其极性运输特性密切相关。分蘖期水稻基部节间不伸长,分蘖节各节密集在一处,地上部极性运输较弱,向各叶位叶涂抹的IAA均能通过极性运输运至节部通过提高IAA含量、降低CTK含量进而提高IAA/CTK值抑制分蘖节分蘖芽萌发,而根部施用的IAA则无法通过极性运输运输至节部位,所以根部施用IAA对分蘖节分蘖芽的萌发无抑制作用。齐穗后,水稻伸长节间伸长,伸长节彼此分开,地上部极性运输较强。向伸长节分蘖芽上部叶或着生叶涂抹的IAA能够通过极性运输运至节部通过提高IAA含量、降低CTK含量进而提高IAA/CTK值抑制伸长节分蘖芽萌发,而向伸长节分蘖芽下部叶涂抹的IAA则无法通过极性运输运至节部,所以向伸长节分蘖芽下部叶涂抹IAA对伸长节分蘖芽的萌发无抑制作用。与IAA不同,CTK对水稻分蘖芽萌发与休眠的调控没有明显的位置效应,向不同叶位叶和根部涂抹6-BA均能促进分蘖芽的萌发。
(3)调控水稻分蘖芽萌发与休眠的CTK在节部合成,然后运输至芽,通过调控芽中OsTB1等基因表达来调控水稻分蘖芽生长；外界因素对分蘖芽萌发与休眠的影响主要通过调控节部位CTK的合成进行；除CTK之外,外源IAA等调控措施还可能通过调控OsD3、OsD10和OsD27等基因的表达调控水稻分蘖芽生长。这表明,在水稻体内存在不止一条途径参与调节分蘖芽的萌发与休眠。
Tillering is an important agronomic trait of rice, and the tillering regulation is an effective means for the establishment of excellent population. In the present study, we conducted a series of experiments in the nutrient solution, pot and field, and external nitrogen, external hormones and panicle removal were used to construct a series of rice population which were difference for the tiller buds growth. The hormonal changes and the expression levels of the genes related to rice tiller buds growth were emphatically measured, and the objects of the present study were to investigate the relationship of hormones and genes during the transformation between dormancy and germination of rice tiller buds, and the mechanism of hormonal regulation of the transformation between dormancy and germination of the tiller buds. The main results of the study are listed below.
(1) The endogenous CTK and IAA play important roles in regulating the tiller buds growth. The CTK is the direct regulator for tiller buds growth, and the IAA is the indirect factor. In general, CTK promoted the growth of rice tiller buds, and IAA inhibited that. However, the transformation of dormancy and germination of tiller bus was not only regulated by the contents of IAA and CTK, but also regulated by the balance of them. The high IAA/CTK ratio promoted the transformation from germination to dormancy of the tiller bus, however, the low IAA/CTK ratio promoted the transformation from dormancy to germination of the tiller buds. The agronomic measures such as nitrogen, hormones and panicle removal all regulated the growth of tiller buds by regulated the contents and balance of the IAA and CTK. The low nitrogen inhibited the growth of tiller buds by inhibited the increasing of CTK and increased the ratio of IAA/CTK, and the promoting effect of high nitrogen and 6-BA on the tiller buds growth was significantly related to the increasing of the CTK contents and the decreasing of the ratio of IAA/CTK. The panicle removal increased the CTK contents and decreased the IAA contents, and then decreased the ratio of IAA/CTK and by this to promote the growth of tiller buds. External application of GA3 and IAA decreased the CTK contents and increased the IAA contents, and then increased the ratio of IAA/CTK and by this to inhibit the growth of tiller buds. The external NAA inhibited the growth of tiller buds by inhibited the increasing of CTK and increased the ratio of IAA/CTK. Besides IAA and CTK, ABA also regulated the growth of rice tiller buds, but ABA may not be the determining factor for the tiller buds growth.
(2) IAA has a notable positional effect on the regulation of rice tiller bud growth, and the positional effect of IAA was closely related to its polar transport. In tillering stage, the basal internodes of rice are not elongated, and this structural characteristic may lead to a deficiency in polar transport of IAA in the aboveground organs. IAA applied to different leaves all can transport to nodes through polar transport, and then decreased the CTK content and increased the ratio of IAA/CTK, and by this to inhibited the growth of tiller buds. However, the IAA applied to root cannot transport to nodes by polar transport, this lead to that applied IAA to roots has no significant effect on tiller buds growth. Contrary to the basal internodes, the upper internodes elongated after heading and that structural characteristic separate its nodes from one another and lead to an enhance in polar transport of IAA in the aboveground organs. IAA applied to the leaves above the buds or the leaves at which the buds locate can be transported to the node. IAA can then inhibit the increase in CTK levels and increased the ratio of IAA/CTK, thus inhibiting the growth of the bud. However, IAA applied to the leaf below the bud cannot be transported to the node by polar transport; this may be the main reason why applied IAA to the leaf below the bud could not inhibit the growth of the bud. In contrast to IAA, CTK had no positional effect on rice tiller bud growth. Application of 6-BA to different leaves and roots significantly promoted the growth of rice tiller buds at both unelongated and elongated nodes.
(3) The CTK which promotes the germination of rice tiller buds is biosynthesis mainly in the nodes and delivery to tiller buds, and then regulated the growth of tiller buds by regulated the expression of OsTBl. The agronomic measures such as external applied IAA regulated the growth of tiller buds by regulated the biosynthesis of CTK in nodes. Besides CTK, the external IAA also might have regulated the expression level of OsD3, OsD10 and OsD27 in tiller buds and by this to regulating the tiller bud growth. These results suggested that the rice have more than one ways to regulate the growth of tiller buds.
引文
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