二倍体与四倍体苦荞的充实差异研究

张余; 何佩云; 罗庆华; 吴兴慧; 孔德章; 黄小燕; 黄凯丰

doi:10.19303/j.issn.1008-0384.2020.10.001

二倍体与四倍体苦荞的充实差异研究

doi: 10.19303/j.issn.1008-0384.2020.10.001

贵州师范大学荞麦产业技术研究中心，贵州　贵阳　550001

基金项目: 国家自然科学基金项目（31560358）；贵州省科技支撑计划项目（黔科合支撑[2019]2297、[2020]1Y048、[2017] 2505）；贵州省教育厅创新群体重大研究项目（黔教合KY字[2018]015）；贵阳市科技计划项目（筑科合同[2019]11-6）；贵州省科技计划项目（黔科合成果[2019] 4334）

详细信息

作者简介:
张余（1999−），女，硕士研究生，主要从事荞麦栽培生理研究（E-mail：zhangyu19990000@163.com）

通讯作者:
黄凯丰（1979−），男，博士，教授，主要从事荞麦栽培生理研究（E-mail：hkf1979@163.com）

中图分类号: S 517
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出版历程
- 收稿日期: 2020-02-13
- 修回日期: 2020-04-23
- 刊出日期: 2020-10-28

Grain-filling of Diploid and Tetraploid Tartary Buckwheat

Buckwheat Industry Technology Research Center, Guizhou Normal University, Guiyang, Guizhou　550001, China

摘要

摘要: 目的比较分析二倍体和四倍体苦荞材料的根系、灌浆特性等相关生理性状，明确四倍体苦荞充实差的生理原因。方法以二倍体苦荞品种小黄荞和六苦2081及其四倍体苦荞材料TB193、TB198为试验材料，比较分析了四倍体和二倍体苦荞间灌浆特性、淀粉合成酶活性、根系形态生理、农艺性状、充实度及产量的差异。结果 2个苦荞品种及其四倍体苦荞材料在花后25 d百粒重几乎不再增加，且二倍体苦荞的百粒重高于四倍体苦荞；四倍体苦荞的灌浆起始势、最大灌浆速率和平均灌浆速率低于二倍体苦荞，达最大灌浆速率的天数高于二倍体；灌浆前期二倍体苦荞籽粒的AGPase和SSS活性强于四倍体苦荞，中后期则低于四倍体苦荞；四倍体苦荞的根系长度、根系表面积、根系体积、根系活力、根系伤流液总体显著高于二倍体苦荞；二倍体苦荞的株高、主茎分枝数、主茎节数、1～2节节间粗度、1～2节节间粗度总体均低于四倍体苦荞，充实度和产量则分别是四倍体苦荞的2.97和1.23倍（两品种的平均值）。结论较低的灌浆起始势、灌浆速率，以及灌浆前期较低的淀粉合成酶活性是四倍体苦荞充实较差的重要生理原因。
- 苦荞 /
- 四倍体 /
- 灌浆特性 /
- 淀粉合成酶活性 /
- 根系形态生理 /
- 充实度
Abstract: Objective Grain-filling of Tartary buckwheat (Fagopyrum tataricum) was studied to determine the cause of inferior crop yield of tetraploid variety. Method Diploid Tartary buckwheat varieties, Xiaohuang and Liuku 2081, were compared with the tetraploid TB193 and TB198 on grain-filling, starch biosynthesis, root morphology and physiology, agronomic characteristics, and grain yield for the analysis. Result Although both Tartary buckwheat varieties stopped the 100-grain weight increase 25 d after anthesis, the diploid was higher than the tetraploid in that respect. In addition, compared to the tetraploid varieties, the diploid Tartary buckwheat started earlier on the filling initiation; achieved higher peak and average filling rates; reached the maximum grain-filling in a shorter time; showed greater AGPase and SSS activities in grains at early grain-filling stage but lower in the middle and late filling stages; had significantly lower length, surface area, volume, activity, and fluid on the roots, height on the plant, number of branches on the main stem, node count on the main stem, girth between the 1^st and 2^nd nodes, and 1-2 internode thickness. On average, the diploid Tartary buckwheat was 2.97 times higher on plumpness and 1.23 times higher on yield than the tetraploid counterpart. Conclusion The lower grain filling initiation, filling rate, and starch synthase activity at early filling stage appeared to be the key physiological indicators of the inferior performance of the tetraploid Tartary buckwheat on grain production.
- Tetraploid Tartary buckwheat /
- tetraploid /
- grain-filling /
- starch synthase activity /
- root morphology and physiology /
- plumpness

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表 1 籽粒的百粒重

Table 1. Hundred-grain weight of Tartary buckwheat （单位：g）

品种 Variety	5 d	10 d	15 d	20 d	25 d	30 d
E1	0.07±0.01 a	0.76±0.07 a	1.37±0.06 a	1.57±0.07 a	1.79±0.03 a	1.80±0.03 a
S1	0.04±0.02 b	0.41±0.02 b	1.12±0.07 b	1.54±0.06 a	1.71±0.02 a	1.77±0.05 a
E2	0.13±0.02 a	0.99±0.03 a	1.63±0.04 a	1.76±0.07 a	1.92±0.05 a	1.96±0.03 a
S2	0.11±0.03 b	0.82±0.05 b	1.40±0.03 b	1.74±0.04 a	1.79±0.03 b	1.83±0.02 a
注：同列数据后不同小写字母表示同一品种不同材料之间差异显著（P＜0.05），表2～6同。 Note: Different letter means significant difference of each level (P＜0.05). The same as table 2-6.

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表 2 籽粒灌浆的Richards方程参数

Table 2. Parameters in Richards equation for evaluating grain-filling of Tartary buckwheat

品种 Variety	生长终值量 A	初值参数 B	生长速率参数 K	形状参数 N	决定系数 R²	灌浆起始势 R₀	灌浆速率最大时间 T_max.G/d	最大灌浆速率 G_max/ （g·d⁻¹）	平均灌浆速率 G_mean/ （g·d⁻¹）	灌浆速率最大生长量/ 籽粒生长终值量 I/%
E1	1.80±0.11	0.51±0.02	0.24±0.03	0.05±0.01	0.995±0.052	4.41±0.42 a	9.48±0.87 b	0.15±0.01 a	0.10±0.02 a	37.76±3.12 a
S1	1.79±0.13	1.73±0.09	0.24±0.02	0.10±0.02	1.000±0.021	2.28±0.21 b	11.92±1.12 a	0.15±0.01 a	0.10±0.01 a	38.61±2.97 a
E2	1.92±0.14	1.09±0.10	0.29±0.01	0.08±0.01	0.996±0.034	3.45±0.27 a	8.80±0.80 b	0.20±0.01 a	0.13±0.01 a	38.29±2.11 a
S2	1.85±0.09	1.13±0.98	0.24±0.01	0.12±0.01	0.998±0.042	1.99±0.28 b	9.64±1.02 a	0.15±0.02 b	0.10±0.01 b	38.86±2.07 a
注：A：生长终值量；B：初值参数；K：生长速率参数；N：形状参数；t：开花后的时间；R²：判断系数；R₀：灌浆起始势；T_max.G：达最大灌浆速率的时间；G_max：最大灌浆速率；G_mean：平均灌浆速率；I为灌浆速率为最大时的生长量与籽粒最终质量比值。 A: The final grain weight at harvest; B: The initial value of parameter; K: The constant growth rate; N: The shape parameter; R² : The compatibility; R₀: The initial growth power; T_max.G : The time with maximum grain-filling rate; G_max: The maximum grain-filling rate; G_mean: The mean grain-filling rate; I: the ratio of the growth at the maximum grain filling rate to the final grain dry weight.

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表 3 籽粒灌浆阶段的划分

Table 3. Grain-filling stages of Tartary buckwheat

品种 Variety	灌浆前期 Early filling stage			灌浆中期 Middle filling stage			灌浆后期 Later filling stage
品种 Variety	天数 Days/d	平均速率 Average rate/（g·d⁻¹）	贡献率 Contribution/%	天数 Days/d	平均速率 Average rate/（g·d⁻¹）	贡献率 Contribution/%	天数 Days/d	平均速率 Average rate/（g·d⁻¹）	贡献率 Contribution/%
E1	5.33±0.47 b	0.03±0.01 a	8.17±0.48 b	13.63±1.21 b	0.13±0.02 a	60.95±5.21 a	28.85±1.27 b	0.04±0.01 a	29.88±1.38 a
S1	7.66±0.49 a	0.02±0.01 b	8.97±0.69 a	16.19±1.09 a	0.13±0.01 a	60.90±4.23 a	31.41±2.34 a	0.03±0.02 b	29.12±2.03 b
E2	5.36±0.34 a	0.03±0.02 a	8.67±0.57 b	12.24±1.57 b	0.17±0.03 a	60.92±5.28 a	24.62±2.10 b	0.05±0.03 a	29.41±2.14 a
S2	5.32±0.36 a	0.03±0.02 a	9.21±0.61 a	13.96±1.02 a	0.13±0.01 b	60.88±1.20 a	29.25±3.02 a	0.04±0.02 b	28.91±2.07 b

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表 4 苦荞籽粒的淀粉合成酶活性

Table 4. Starch synthase activity of Tartary buckwheat

指标 Item	品种 Variety	时期 Period
指标 Item	品种 Variety	5 d	10 d	15 d	20 d	25 d	30 d
ADPG 焦磷酸化酶AGPase/（U ·g⁻¹·min⁻¹）	E1	0.31±0.02 a	0.55±0.03 a	0.46±0.04 b	0.36±0.03 b	0.27±0.02 b	0.26±0.02 b
	S1	0.20±0.01 b	0.36±0.02 b	0.49±0.05 a	0.43±0.05 a	0.33±0.04 a	0.30±0.02 a
	E2	0.26±0.03 a	0.46±0.03 a	0.36±0.04 b	0.27±0.03 b	0.20±0.01 b	0.20±0.03 a
	S2	0.15±0.0.2 b	0.32±0.02 b	0.41±0.02 a	0.32±0.01 a	0.23±0.02 a	0.20±0.02 a
可溶性淀粉合酶SSS/（U ·mg⁻¹ ·min⁻¹）	E1	6.56±0.28 a	10.03±1.07 a	4.97±0.36 b	2.47±0.17 b	2.01±0.11 b	1.03±0.09 b
	S1	4.92±0.39 b	8.55±0.38 b	5.98±0.32 a	3.38±0.25 a	2.98±0.23 a	1.53±0.11 a
	E2	3.72±0.34 a	9.28±0.59 a	4.24±0.51 b	2.16±0.19 b	1.68±0.10 b	0.97±0.01 b
	S2	3.41±0.16 b	7.54±0.46 b	4.74±0.40 a	2.38±0.20 a	2.00±0.25 a	1.33±0.20 a
淀粉分支酶SBE/（U ·g⁻¹·min⁻¹）	E1	2.37±0.18 a	4.55±0.32 a	4.39±0.29 a	3.97±0.45 a	3.45±0.27 a	3.17±0.28 a
	S1	1.33±0.09 b	3.31±0.17 b	4.07±0.24 b	3.71±0.31 b	3.22±0.18 b	2.95±0.32 b
	E2	2.29±0.17 a	4.45±0.35 a	4.32±0.38 a	3.87±0.28 a	3.25±0.41 a	2.53±0.24 a
	S2	1.12±0.11 b	3.13±0.21 b	3.96±0.21 b	3.49±0.24 b	2.54±0.20 b	1.93±0.17 b

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表 5 二倍体和四倍体苦荞的根系形态及生理指标比较

Table 5. Root morphology and physiology of diploid and tetraploid Tartary buckwheat

时期 Stage	品种 Variety	根系长度 Length/cm	根系表面积 Surface area/cm²	根系体积 Volume/cm³	根系平均直径 Average diameter/mm	根系伤流液 Root system fluid/（g·h⁻¹）	根系活力 Root activity/ （μg·g⁻¹·h⁻¹）
苗期 Seedling stage	E1	44.91±1.49 b	5.03±0.16 b	0.10±0.01 b	0.42±0.02 a	0.044±0.003 a	21.71±1.21 a
	S1	55.50±1.34 a	6.46±0.21 a	0.12±0.01 a	0.43±0.0.3 a	0.041±0.002 a	21.23±1.06 a
	E2	43.57±1.27 b	4.31±0.20 b	0.07±0.03 b	0.37±0.02 b	0.025±0.001 b	13.91±0.98 b
	S2	55.08±1.12 a	6.36±0.18 a	0.13±0.04 a	0.44±0.01 a	0.033±0.003 a	17.16±1.13 a
开花期 Anthesis stage	E1	168.49±9.45 b	31.06±1.34 b	1.28±0.03 b	0.53±0.06 a	0.026±0.001 b	3.31±0.11 b
	S1	200.85±8.69 a	35.95±1.26 a	1.50±0.04 a	0.54±0.05 a	0.035±0.002 a	6.40±0.28 a
	E2	156.79±9.12 b	25.48±1.33 b	0.78±0.02 b	0.51±0.02 a	0.025±0.001 b	3.92±0.32 a
	S2	300.07±9.34 a	55.29±1.06 a	2.53±0.16 a	0.54±0.04 a	0.052±0.002 a	3.94±0.40 a
灌浆期 Grain filling stage	E1	264.61±8.37 a	31.28±1.25 b	0.96±0.01 b	0.42±0.02 a	0.004±0.002 b	4.79±0.29 b
	S1	220.99±8.67 b	36.72±2.26 a	1.23±0.17 a	0.44±0.03 a	0.006±0.004 a	7.90±0.47 a
	E2	254.45±9.61 a	23.09±1.63 b	0.58±0.03 b	0.40±0.02 b	0.010±0.001 b	7.49±0.51 b
	S2	186.76±6.47 b	36.50±1.71 a	1.21±0.11 a	0.44±0.04 a	0.016±0.002 a	5.35±0.20 a
成熟期 Mature stage	E1	229.37±10.55 b	29.49±0.99 b	0.85±0.04 b	0.44±0.03 a	0.005±0.001 b	2.98±0.37 b
	S1	245.02±9.91 a	36.04±1.01 a	1.31±0.03 a	0.46±0.02 a	0.006±0.002 a	5.37±0.31 a
	E2	171.82±7.34 b	27.21±1.13 b	0.94±0.05 b	0.44±0.03 a	0.009±0.002 b	3.55±0.26 a
	S2	230.18±10.25 a	34.38±1.08 a	1.38±0.02 a	0.46±0.04 a	0.016±0.004 a	3.57±0.29 a

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表 6 二倍体和四倍体苦荞的农艺性状及产量比较

Table 6. Agronomic traits and yield of Tartary buckwheat

品种 Variety	株高 Plant height/cm	主茎节数 Number of main stem nodes/个	主茎分枝数 Main stem branch number/个	子叶节高度 Cotyledon node height/cm	1～2节节间长度 1-2 internode length/cm	1～2节节间粗度 1-2 internode thickness/mm	充实度 Plumpness/%	产量 Yield/（kg·hm⁻²）
E1	112.3±1.3 b	12.2±1.1 b	7.1±1.2 b	1.6±0.1 a	1.7±0.1 b	4.7±0.4 b	74.2±1.4 a	2452.1±102.2 a
S1	125.0±1.1 a	17.6±1.1 a	9.6±1.1 a	1.8±0.2 a	3.3±0.2 a	6.3±0.5 a	27.7±0.8 b	2058.1±112.3 b
E2	119.4±1.35 b	15.3±1.7 b	5.0±1.0 b	1.7±0.2 a	1.7±0.1 b	4.7±0.4 b	85.2±1.2 a	2305.8±106.3 a
S2	127.0±1.0 a	18.2±1.3 a	7.7±1.3 a	1.6±0.2 a	3.0±0.2 a	6.2±0.5 a	26.2±0.7 b	1807.3±99.4 b

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