Effect of Endogenous Hormones and Expressions of Relevant Genes on Tuber Growth of Bikeqi Yam
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摘要:
目的 分析毕克齐山药中内源激素含量与淀粉含量、可溶性总糖含量和还原糖含量的关系，以及与其相关合成基因表达量的相关性，探究毕克齐山药块茎膨大的机理。 方法 以毕克齐山药5个不同生长期块茎为材料，采用酶联免疫吸附法分别测定脱落酸（ABA）、赤霉素（GA3）、生长素（IAA）、茉莉酸（JA）、玉米素（ZR）、异戊烯基腺苷（IPA）等6种内源激素含量，采用高效液相色谱仪测定方法测定水杨酸（SA）含量。 结果 IAA、ZR、ABA、JA和SA含量与山药块茎形态指标正相关；GA3、IPA含量与形态指标负相关；IAA含量与山药块茎周长和块茎直径显著正相关；GA3含量与块茎长度显著负相关；与IAA相关的基因与内源激素IAA含量显著负相关。 结论 内源激素IAA、ZR、ABA、JA和SA促进山药块茎膨大；GA3、IPA抑制山药块茎生长；IAA促进山药增粗；GA3抑制其伸长生长；与IAA相关的基因的下调表达对IAA的合成有促进作用，即正调控IAA含量。 Abstract:Objective Effects of endogenous hormones and expressions of genes related to the hormone syntheses on the morphology and physiology of bikequ yam (Dioscorea Opposita) tubers during expanding stage were investigated. Correlation between the indicators was established to provide guidelines for upgrading the yield and quality of the yam farming. Method During expansion stage of the tubers, contents of starch, reducing sugar, and soluble total sugar were determined by chemical analysis, those of endogenous ABA, GA3, IAA, JA, ZR, and IPA by enzyme-linked immunosorbent assay, and that of salicylic acid (SA) by HPLC. Correlations between these indicators were tested by regression analysis. The gene expression levels of endogenous hormone related genes were obtained by transcriptome sequencing. Result The contents of IAA, ZR, ABA, JA, and SA correlated positively with the morphological indices on the tuber, while those of GA3 and IPA correlated negatively. IAA correlated with the tuber perimeter and diameter. GA3 correlated inversely with the length of tuber. The IAA-related gene was found to negatively correlate with IAA in the tubers. Conclusion Among the endogenous hormones, IAA, ZR, ABA, JA, and SA promoted the tuber enlargement; GA3 and IPA inhibited the growth; IAA enhanced the girth increase; and, GA3 retarded the elongation. The downregulation of IAA-related gene increased the hormone synthesis. In other words, the gene could regulate the IAA production in the yams, and in turn, affected the tuber growth. -
图 1 生长发育过程各个生理指标的变化
注：A-生长发育过程还原糖含量的变化；B-生长发育过程可溶性总糖含量的变化；C-生长发育过程淀粉含量的变化。
Figure 1. Physiological changes on Bikeqi yam during tuber development
Note: A-on reducing sugars; B-on soluble sugars; C-on starch.
表 1 毕克齐山药在块茎膨大过程内源激素含量变化
Table 1. Changes on endogenous hormones during tuber enlargement of Bikeqi yam
[单位: ng·g−1（FW）] 项目 Item 105 d 120 d 135 d 150 d 165 d 生长素 IAA 24.697±1.82 b 38.542±1.76 b 46.454±3.46 a 37.793±2.41 b 48.012±2.68 a 玉米素 ZR 5.295±0.23 b 6.97±0.39b 8.129±0.45 ab 9.39±0.72 a 6.751±0.36 b 脱落酸 ABA 33.274±1.93 bc 33.564±0.03 bc 28.402±1.70 c 40.707±2.11 b 81.939±1.88 a 茉莉酸 JA 11.976±0.43 b 12.678±0.39 b 15.806±1.02 ab 13.541±0.92 b 21.276±1.46 a 赤霉素 GA3 5.484±0.44 a 2.538±0.09 c 4.072±0.36 bc 3.209±0.20 bc 3.69±0.26 bc 异戊烯基腺苷 IPA 4.103±0.20 b 8.005±0.30 a 5.943±0.37 ab 4.202±0.21 b 4.419±0.27 b 水杨酸 SA 52.308±30.38 b 88.763±63.26 ab 19.226±10.83 b 81±37.48 ab 234.867±111.44 a 注：同行不同小写字母表示在0.05水平上差异显著。
Note: Data with different lowercase letters indicate significant differences at 0.05 level.
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表 2 毕克齐山药内源激素与形态、生理指标相关性
Table 2. Correlation between endogenous hormones and morphology and physiology of Bikeqi yam tubers
项目
Item块茎长度
Length块茎周长
Circumference块茎直径
Diameter块茎鲜重
Fresh
weight块茎干重
Dry
weight淀粉含量
Starch
content还原糖含量
Reducing sugar
content可溶性总糖含量
Total soluble
sugar contentIAA ZR ABA JA GA3 IPA SA 块茎长度
Length0.756 0.757 0.674 0.679 0.375 −0.78 −0.940* 0.841 0.562 0.431 0.531 −0.892* 0.435 0.494 块茎周长
Circumference1.000** 0.922* 0.981** 0.719 −0.823 −0.839 0.906* 0.589 0.64 0.833 −0.425 −0.161 0.553 块茎直径
Diameter0.924* 0.982** 0.715 −0.822 −0.838 0.904* 0.588 0.643 0.833 −0.425 −0.162 0.557 块茎鲜重
Fresh weight0.972** 0.489 −0.84 −0.75 0.692 0.657 0.665 0.694 −0.43 −0.362 0.591 块茎干重
Dry weight0.646 −0.817 −0.771 0.807 0.602 0.677 0.804 −0.36 −0.321 0.579 淀粉含量
Starch content−0.617 −0.611 0.767 0.542 0.072 0.524 −0.061 −0.04 −0.082 还原糖含量
Reducing sugar content0.927* −0.732 −0.932* −0.21 −0.375 0.673 −0.034 −0.170 可溶性总糖含量
Total soluble sugar content−0.883* −0.773 −0.289 −0.494 0.805 −0.315 −0.293 IAA 0.483 0.479 0.779 −0.533 0.227 0.433 ZR −0.116 0.043 −0.575 0.022 −0.159 ABA 0.871 −0.118 −0.378 0.975** JA −0.107 −0.242 0.787 GA3 −0.612 −0.252 IPA −0.222 注：*在0.05水平，相关性显著。**在0.01水平，相关性极显著。表5、6同。
Note:*Significant correlation （P＜0.05）; **Extremely significant correlation （P＜0.01）. Same for Tables 5 and 6.
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表 3 与内源激素合成相关基因的基因编号、基因代号和基因注释信息
Table 3. Number, code, and information on genes associated with endogenous hormone synthesis in Bikeqi yam tubers
项目
Item基因
ID Gene ID基因代号
Gene code基因注释信息
Gene annotation information与IAA合成相关基因 Genes related to IAA synthesis Unigene0016848 IAA 1 生长素响应蛋白基因 Auxin response protein gene Unigene0021936 IAA 2 生长素响应蛋白基因 Auxin response protein gene Unigene0023040 IAA 3 生长素响应蛋白基因 Auxin response protein gene Unigene0025388 IAA 4 IAA-氨基酸水解酶 ILR1基因 IAA-amino acid hydrolase ILR1 gene Unigene0027417 IAA 5 IAA-氨基酸水解酶 ILR1基因 IAA-amino acid hydrolase ILR1 gene Unigene0027794 IAA 6 生长素响应蛋白基因 Auxin response protein gene Unigene0028701 IAA 7 生长素响应蛋白基因 Auxin response protein gene Unigene0028916 IAA 8 生长素响应蛋白基因 Auxin response protein gene Unigene0030426 IAA 9 生长素响应蛋白基因 Auxin response protein gene 与 GA3合成相关基因 Genes related to GA3 synthesis Unigene0002430 GA3 1 赤霉素2-β-双加氧酶 Gibberellin 2-β-dioxygenase Unigene0019924 GA3 2 赤霉素2-β-双加氧酶 Gibberellin 2-β-dioxygenase Unigene0021502 GA3 3 赤霉素2-β-双加氧酶 Gibberellin 2-β-dioxygenase Unigene0021503 GA3 4 赤霉素2-β-双加氧酶 Gibberellin 2-β-dioxygenase Unigene0027812 GA3 5 赤霉素20氧化酶 Gibberellin 2-β-dioxygenase Unigene0023907 GA3 6 赤霉素调节蛋白基因 Gibberellin regulatory protein gene Unigene0024440 GA3 7 赤霉素调节蛋白基因 Gibberellin regulatory protein gene Unigene0026445 GA3 8 赤霉素调节蛋白基因 Gibberellin regulatory protein gene Unigene0020765 GA3 9 赤霉素调节蛋白基因 Gibberellin regulatory protein gene
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表 4 与内源激素合成相关基因的rpkm平均值
Table 4. Average rpkm of genes associated with endogenous hormone synthesis
项目
Item基因ID
Gene ID基因代号
Gene code105 d 120 d 135 d 150 d 165 d 与IAA合成相关基因
Genes related to IAA synthesisUnigene0016848 IAA 1 95.82±30.63 65.29±44.90 64.25±22.21 26.73±12.83 26.68±10.96 Unigene0021936 IAA 2 14.49±2.45 8.08±6.19 3.64±2.22 2.95±3.33 3.57±0.87 Unigene0023040 IAA 3 213.97±40.34 119.15±61.21 132.16±54.901 48.88±36.07 39.56±9.26 Unigene0025388 IAA 4 4.61±3.19 5.45±0.97 8.84±2.12 1.18±0.84 0.00 Unigene0027417 IAA 5 11.82±1.17 7.21±3.44 5.10±1.42 1.48±1.15 3.18±0.68 Unigene0027794 IAA 6 2.28±0.33 0.82±0.67 0.53±0.42 0.28±0.39 0.00 Unigene0028701 IAA 7 60.78±12.10 30.43±21.84 77.84±20.86 50.67±17.82 21.64±5.46 Unigene0028916 IAA 8 127.67±26.48 40.53±21.48 66.59±24.93 35.91±11.76 14.06±1.25 Unigene0030426 IAA 9 7.79±1.10 2.82±3.12 3.91±1.72 0.36±0.51 2.20±0.80 与GA3合成相关基因
Genes related to GA3 synthesisUnigene0002430 GA3 1 24.60±5.23 24.72±10.21 18.71±6.22 5.74±6.31 2.92±0.83 Unigene0019924 GA3 2 4.80±1.80 0.62±0.60 1.20±1.54 0.00 0.00 Unigene0021502 GA3 3 4.02±1.56 1.21±1.06 3.05±2.99 2.34±1.78 0.00 Unigene0021503 GA3 4 5.82±3.34 5.51±2.87 5.20±2.06 6.88±7.74 0.00 Unigene0027812 GA3 5 13.12±2.80 1.38±1.23 2.59±0.96 1.91±0.92 5.69±1.14 Unigene0023907 GA3 6 16.76±10.56 4.20±5.94 0.67±0.60 0.08±0.12 0.42±0.60 Unigene0024440 GA3 7 176.89±19.66 189.53±72.98 110.84±19.67 80.80±39.49 423.10±68.62 Unigene0026445 GA3 8 47.20±12.01 32.18±34.09 51.94±10.28 74.13±97.99 203.55±60.52 Unigene0020765 GA3 9 11.95±2.51 1.74±1.95 4.61±2.35 0.55±0.78 0.00
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表 5 内源激素IAA含量与其合成相关基因表达量的相关性分析
Table 5. Correlation between IAA and expression of IAA synthesis-related gene
项目
ItemIAA含量
IAA contentIAA 1 IAA 2 IAA 3 IAA 4 IAA 5 IAA 6 IAA 7 IAA 8 IAA 9 IAA含量 IAA content −0.681 −0.875 −0.719 −0.046 −0.744 −0.902* −0.230 −0.799 −0.644 IAA 1 0.868 0.992** 0.669 0.956* 0.912* 0.482 0.898* 0.916* IAA 2 0.869 0.218 0.961** 0.966** 0.128 0.833 0.869 IAA 3 0.638 0.944* 0.936* 0.547 0.946* 0.936* IAA 4 0.442 0.340 0.700 0.481 0.453 IAA 5 0.944* 0.245 0.855 0.937* IAA 6 0.373 0.944* 0.904* IAA 7 0.643 0.418 IAA 8 0.911* IAA 9
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表 6 内源激素GA3含量与其合成相关基因表达量的相关性分析
Table 6. Correlation between GA3 and expression of GA3 synthesis-related gene
项目
ItemGA3含量
GA3 contentGA3 1 GA3 2 GA3 3 GA3 4 GA3 5 GA3 6 GA3 7 GA3 8 GA3 9 GA3含量 GA3 content 0.256 0.863 0.680 0.020 0.900* 0.737 −0.007 −0.041 0.872 GA3 1 0.652 0.557 0.511 0.267 0.648 −0.400 −0.803 0.673 GA3 2 0.765 0.306 0.876 0.958* −0.185 −0.416 0.991** GA3 3 0.728 0.468 0.620 −0.730 −0.704 0.826 GA3 4 −0.096 0.277 −0.957* −0.910* 0.354 GA3 5 0.873 0.224 0.058 0.827 GA3 6 −0.085 −0.383 0.914* GA3 7 0.854 −0.265 GA3 8 −0.469 GA3 9
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