套作大豆苗期倒伏与茎秆内源赤霉素代谢的关系

doi:10.3864/j.issn.0578-1752.2015.13.005

摘要/Abstract

摘要： 【目的】从内源赤霉素代谢角度，阐明玉米大豆套作环境下，大豆苗期茎秆发生藤蔓化倒伏的原因。【方法】在大豆单作和玉米大豆套作两种种植方式下，以耐荫抗倒型大豆南豆12和不耐荫抗倒型大豆南032-4为试验材料，采用石蜡切片、液相色谱质谱联用、SYBR^®GreenⅡ荧光定量PCR等方法，对茎秆形态、解剖结构、赤霉素含量、赤霉素代谢相关基因表达等进行测定和分析。【结果】与大豆单作相比，玉米大豆套作下，由于受玉米荫蔽的影响，南豆12和南032-4两个大豆材料苗期均出现典型的避荫性反应，即主茎节数减少，节间长和株高显著增加，苗期出现藤蔓化和倒伏，但两大豆材料间反应程度不同，差异极显著。与南032-4比，南豆12受套作荫蔽的影响较小，茎秆藤蔓化程度低，倒伏率仅29.93%，极显著低于南032-4（93.94%）（P<0.05）；南032-4株高增加32.64 cm，与南豆12（22.95 cm）呈显著差异（P<0.05）；两材料主茎节数减少，材料间无差异；南032-4中上部节间长度显著长于南豆12，说明套作下植株株高的增加来源于节间的伸长，而非节数的增多，且中部及上部节间过度伸长易导致植株茎秆藤蔓化和倒伏。从茎秆纵切面的解剖结构可知，玉米大豆套作条件下，大豆主茎细胞都有不同程度的伸长，南032-4茎秆髓部、木质部、韧皮部细胞伸长明显，而南豆12在两种种植模式下各部分细胞无明显变化，说明大豆株高的增加源于细胞的伸长，非细胞的分裂。内源赤霉素代谢分析结果表明，套作条件下，两材料茎秆内源赤霉素含量均显著降低，南豆12茎秆内赤霉素A₄（GA₄）的含量显著低于南032-4；对编码赤霉素代谢途径中关键酶基因表达量的分析表明，除赤霉素合成酶基因GmGA20ox5、GmGA3ox6和赤霉素降解酶基因GmGA2ox4在不同种植方式和不同材料间均保持较低的水平外，茎秆中其他内源赤霉素合成酶GA-20氧化酶基因、GA3-氧化酶基因和赤霉素降解酶GA2-氧化酶基因表达量均高于单作，且南032-4的表达量高于南豆12，说明植株内源赤霉素的含量对编码赤霉素代谢途径中关键酶基因具有前馈和反馈作用，南豆12茎尖维持较低活性赤霉素GA₄水平，抑制植株主茎过度伸长，最终表现出较强的耐荫抗倒性。【结论】大豆内源赤霉素GA₄的合成受基因型和光环境的共同影响，从而影响大豆茎秆生长发育，最终导致不同材料在耐荫抗倒表型上的典型差异。

关键词: 大豆, 套作, 苗期倒伏, 茎秆, 内源赤霉素

Abstract: 【Objective】 From the perspective of endogenous gibberellins(GAs) metabolism, the reasons of soybean seedling vine lodging under the maize-soybean relay strip intercropping system were studied. 【Method】Under maize-soybean relay strip intercropping system and soybean sole cropping system, shade-tolerant and lodging resistant Nandou12 and non-shade tolerant and non- lodging resistant Nan032-4 were taken as experimental materials. The stalk morphology, anatomical structure, the level of gibberellin A₄, and the genes that are concerned with gibberellins expression profiles were analyzed with the paraffin section, Gas chromatography-mass spectrometry analyses, real-time qPCR with SYBR^®Green Ⅱ and other. methods. 【Result】Compared with soybean monoculture, due to the effects of maize shading at the seeding stage, both Nandou12 and Nan032-4 appeared obvious shade avoidance responses in relay strip intercropping system, including reduction of node number, significant increase of plant height and internode length, vining and lodging of the seedlings. However, the shade avoidance phenotypes were quite different between these two materials. Compared with the shade-tolerant cultivar Nandou12, the shade-sensitive Nan032-4 was influenced more seriously, for which 93.94% of the plants showed a lodging phenotype, much higher than that of Nandou12 (29.93%) (P＜0.05). The height of Nan032-4 was increased by 32.64 cm, different from that of Nandou12 (22.95 cm). The node number of stem reduced, but there was no significance between the two varieties. The upper nodes of Nan032-4 were longer than Nanou12 remarkably, that is to say, in the relay intercropping system, the increase of plant height of these two varieties resulted from internodes elongation, not from the increase of the node number, and the excessive elongation of middle and upper nodes could lead stems to vine and lodge easily. Vertical section of soybean stem revealed that the cell length of stems showed different degrees in elongation between the two varieties, especially for Nan032-4, not Nandou12, showed an apparent increase in cell length of stem pith, xylem and phloem. Therefore, these results suggest that the increase of plant height of these two varieties was caused by cell elongation, but not cell division. Endogenous gibberellins metabolism analyses showed the GA₄ content in the stems of both varieties was decreased significantly in relay strip intercropping system, but was much lower in Nandou12 compared to Nan032-4. The transcript levels of genes encoding key enzymes in gibberellins metabolic pathways were also investigated in the present study. The results show that the expression of genes encoding GA20-oxidase, GA3-oxidase, and GA2-oxidase were not only higher in Nan032-4 than that in Nadou12, but also up-regulated by intercropping in both varieties. However, GA synthetic genes of GmGA20ox5 and GmGA3ox6, as well as the GA deactivating gene GmGA2ox4 displayed a lower expression pattern in different plant modes and varieties. These results suggest that the content of endogenous gibberellins negatively regulate the expression of key genes in GA related pathways with a feedforward and feedback manner. Low level of GA₄ in the stem tips of Nandou12 might repress the stem elongation excessively, and contribute to a shade avoidance syndrome.【Conclusion】Synthesis of soybean endogenous GA₄ is affected by the genotypes and the light environment, thus influencing the growth and development of soybean stem, which finally results in the typical differences in shade lodging phenotype among varieties.

Key words: soybean, relay intercropping, lodging at seedling stage, stem, endogenous gibberellins

罗玲，于晓波，万燕，蒋涛，杜俊波，邹俊林，杨文钰，刘卫国. 套作大豆苗期倒伏与茎秆内源赤霉素代谢的关系[J]. 中国农业科学, 2015, 48(13): 2528-2537.

LUO Ling, YU Xiao-bo, WAN Yan, JIANG Tao, DU Jun-bo, ZOU Jun-lin, YANG Wen-yu, LIU Wei-guo. The Relationship Between Lodging and Stem Endogenous Gibberellins Metabolism Pathway of Relay Intercropping Soybean at Seedling Stage[J]. Scientia Agricultura Sinica, 2015, 48(13): 2528-2537.

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