Effects of exogenous 14-hydroxybrassinolide on floret development into grain and physiological characteristics of winter wheat
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摘要: 为探索14-羟基芸苔素甾醇(14-HBR)对小麦穗花发育成粒的影响, 以多穗型品种‘豫麦49-198’(V1)和大穗型品种‘周麦16’(V2)为材料, 在小麦拔节后20 d叶面喷施0.05 μmol∙L−1的14-HBR, 通过观察穗花发育进程, 测定旗叶碳氮代谢酶和不同器官碳氮含量及产量构成因素, 分析外源14-HBR对穗花发育的调控效应及促粒增产的生理机制。结果表明, 外源14-HBR可以有效增加小麦可孕小花数量, 提高可孕小花和小穗结实率, 对大穗型品种V2促进效果更明显。喷施14-HBR可以提高小麦旗叶Rubisco酶、蔗糖磷酸合成酶(SPS)和蔗糖合成酶(SS)以及谷氨酰胺合成酶(GS)和硝酸还原酶(NR)的活性, 且总体上对品种V2的促进效应更高。外源14-HBR可以提高穗和非穗器官的可溶性糖和氮含量及干物质积累量, 促进可溶性糖向穗部转运, 提高穗器官C/N比, 为穗花发育提供充足的碳营养, 且总体对品种V2的促进效果更好。考察V1和V2两品种产量及其构成因素发现, 喷施14-HBR对两品种穗数和千粒重影响较小, 而对穗粒数和产量促进效果显著, 其中产量增幅分别为9.31%和12.03%, 穗粒数增幅分别为9.85%和11.40%。喷施14-HBR主要通过提高小麦碳同化能力, 增强氮素吸收与积累, 促进碳营养物质向穗部转运和分配, 从而为穗花发育提供良好的物质基础以减少可孕小花的退化和败育, 进而提高小花结实数和结实率, 且对大穗型品种的影响更明显。
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关键词:
- 冬小麦 /
- 14-羟基芸苔素甾醇 /
- 小花发育 /
- 生理特性 /
- 穗粒数
Abstract: A high grain number per spike is the basis for achieving a high yield. Promoting spike and floret development, and reducing floret degeneration and abortion can effectively increase the development of fertile florets into more grains. Exploring the effect of spraying exogenous 14-hydroxybrassinosterol (14-HBR) on the development of wheat florets into grains contributes to increasing grains number per spike and improving yield. A field experiment of spraying 0.05 μmol∙L−1 14-HBR on wheat leaf surface at 20 days after jointing was conducted with multi-spike wheat cultivar ‘Yumai 49-198’ and large-spike wheat cultivar ‘Zhoumai 16’. The development of young spikes and florets, carbon and nitrogen metabolism enzymes in flag leaves, dry matter weight, soluble sugar and nitrogen contents of different organs, and yield components were measured to analyze the effect of exogenous 14-HBR on the optimization of spike and floret development, as well as the physiological mechanism of 14-HBR promoting grains number. The results showed that spraying exogenous 14-HBR increased the number of fertile florets and the setting rate of fertile florets and spikelets. The promoting effect was more obvious for the large-spike wheat cultivar ‘Zhoumai 16’. Spraying exogenous 14-HBR increased the activities of RuBisCo enzyme, sucrose phosphorylase, and sucrose synthase for carbon metabolism, and activities of glutamine synthetase and nitrate reductase for nitrogen metabolism in wheat flag leaf, which promoted the synthesis of photosynthetic products and the metabolism and accumulation of carbon and nitrogen. Overall, the promoting effect was higher for large-spike wheat cultivar ‘Zhoumai 16’ than multi-spike wheat cultivar ‘Yumai 49-198’. Spraying exogenous 14-HBR increased the contents of soluble sugar and nitrogen and dry matter accumulation of spike organ and non-spike organ, promoted the transport and distribution of soluble sugar to spike, and improved the C/N ratio of spike organs, which provided sufficient carbon nutrients for spike and floret development, and overall, the promoting effect was more effective for large-spike wheat cultivar ‘Zhoumai 16’. To analyze the effect of exogenous 14-HBR on the yield and yield components, spraying 14-HBR 20 days after elongation had little effect on spike number and thousand-grain weight, but a significant impact on grains number per spike and yield was observed. Spraying exogenous 14-HBR significantly improved the grains number per spike, which was the main way to increase the yield. Compared with CK treatment (no spraying 14-HBR but spraying water), for the spraying 14-HBR treatment of multi-spike wheat cultivar ‘Yumai 49-198’ and large-spike wheat cultivar ‘Zhoumai 16’, the increase in grains number per spike was 9.85% and 11.40%, respectively, and the increase in yield was 9.31% and 12.03%, respectively. Spraying exogenous 14-HBR mainly improved the carbon assimilation ability of wheat, enhanced nitrogen absorption and accumulation, and promoted the transport and distribution of carbon nutrients to spike organs, thereby providing a good material basis for the two-stage differentiation of florets, reducing the degradation and abortion of florets, and increasing the number and rate of floret settings. -
图 1 外源喷施14-羟基芸苔素甾醇(14-HBR)对小麦多穗型品种‘豫麦49-198’(V1)和大穗型品种‘周麦16’(V2)穗花发育及结实的影响
CK: 喷施清水对照; BR: 喷施14-HBR处理。不同小写字母表示不同处理间差异显著(P<0.05)。
Figure 1. Effect of exogenous spraying 14-hydroxybrassinolide (14-HBR) on spike and floret development and setting of multi-spike wheat cultivar ‘Yumai 49-198’ (V1) and large-spike wheat cultivar ‘Zhoumai 16’ (V2)
CK: control with spraying water; BR: spraying 14-HBR. Different lowercase letters show significant differences among treatments (P<0.05).
图 2 外源喷施14-羟基芸苔素甾醇(14-HBR)对小麦多穗型品种‘豫麦49-198’(V1)和大穗型品种‘周麦16’(V2)穗和非穗器官干物质积累的影响
CK: 喷施清水对照; BR: 喷施14-HBR处理。
Figure 2. Effect of exogenous spraying 14-hydroxybrassinolide (14-HBR) on dry matter accumulation in spike and non-spike organs of multi-spike wheat cultivar ‘Yumai 49-198’ (V1) and large-spike wheat cultivar ‘Zhoumai 16’ (V2)
CK: control with spraying water; BR: spraying 14-HBR.
图 3 外源喷施14-羟基芸苔素甾醇(14-HBR)对小麦多穗型品种‘豫麦49-198’(V1)和大穗型品种‘周麦16’(V2)穗与非穗器官氮含量和氮积累量的影响
CK: 喷施清水对照; BR: 喷施14-HBR处理。
Figure 3. Effects of exogenous spraying 14-hydroxybrassinolide (14-HBR) on nitrogen content and nitrogen accumulation in spike and non-spike organs of multi-spike wheat cultivar ‘Yumai 49-198’ (V1) and large-spike wheat cultivar ‘Zhoumai 16’ (V2)
CK: control with spraying water; BR: spraying 14-HBR.
图 4 外源喷施14-羟基芸苔素甾醇(14-HBR)对小麦多穗型品种‘豫麦49-198’(V1)和大穗型品种‘周麦16’(V2)穗和非穗器官可溶性糖含量的影响
CK: 喷施清水对照; BR: 喷施14-HBR处理。CK: control with spraying water; BR: spraying 14-HBR.
Figure 4. Effect of exogenous spraying 14-hydroxybrassinolide (14-HBR) on soluble sugar content in spike and non-spike organs of multi-spike wheat cultivar ‘Yumai 49-198’ (V1) and large-spike wheat cultivar ‘Zhoumai 16’ (V2)
图 5 外源喷施14-羟基芸苔素甾醇(14-HBR)对小麦多穗型品种‘豫麦49-198’(V1)和大穗型品种‘周麦16’(V2)穗和非穗器官C/N比的影响
CK: 喷施清水对照; BR: 喷施14-HBR处理。CK: control with spraying water; BR: spraying 14-HBR.
Figure 5. Effect of exogenous spraying 14-hydroxybrassinolide (14-HBR) on C / N ratio in spike and non-spike organs of multi-spike wheat cultivar ‘Yumai 49-198’ (V1) and large-spike wheat cultivar ‘Zhoumai 16’ (V2)
图 6 外源喷施14-羟基芸苔素甾醇(14-HBR)对小麦多穗型品种‘豫麦49-198’(V1)和大穗型品种‘周麦16’(V2)旗叶Rubisco、蔗糖磷酸合成酶(SPS)和蔗糖合成酶(SS)活性的影响
CK: 喷施清水对照; BR: 喷施14-HBR处理。CK: control with spraying water; BR: spraying 14-HBR.
Figure 6. Effects of exogenous spraying 14-hydroxybrassinolide (14-HBR) on activities of Rubisco, sucrose phosphate synthase (SPS) and sucrose synthase (SS) enzymes in flag leaf of multi-spike wheat cultivar ‘Yumai 49-198’ (V1) and large-spike wheat cultivar ‘Zhoumai 16’ (V2)
图 7 外源喷施14-羟基芸苔素甾醇(14-HBR)对小麦多穗型品种‘豫麦49-198’(V1)和大穗型品种‘周麦16’(V2)旗叶谷氨酰胺合成酶(GS)和硝酸还原酶(NR)活性的影响
CK: 喷施清水对照; BR: 喷施14-HBR处理。CK: control with spraying water; BR: spraying 14-HBR.
Figure 7. Effect of exogenous spraying 14-hydroxybrassinolide (14-HBR) on activities of glutamine synthetase (GS) and nitrate reductase (NR) enzymes in flag leaf of multi-spike wheat cultivar ‘Yumai 49-198’ (V1) and large-spike wheat cultivar ‘Zhoumai 16’ (V2)
表 1 外源喷施14- 羟基芸苔素甾醇(14-HBR)对不同小麦品种产量及其构成因素的影响
Table 1. Effect of exogenous spraying 14-hydroxybrassinolide (14-HBR) on yield and yield components of different wheat cultivars
年度
Year品种
Cultivar处理
Treatment穗数
Spike number
(×104spikes∙hm−2)穗粒数
Grains number per spike千粒重
1000-grain weight (g)产量
Grain yield
(kg∙hm−2)2018—2019 豫麦49-198 Yumai 49-198 CK 612.87±2.29a 33.66±0.76c 40.41±0.68b 7229.12±138.86d BR 608.44±0.67a 36.70±0.80b 41.23±0.63b 7968.66±37.90bc 周麦16 Zhoumai 16 CK 520.86±2.19b 37.01±1.04b 44.26±2.13a 8204.11±81.49b BR 524.42±0.41b 41.62±0.52a 44.55±0.13a 9143.48±87.81a 2019—2020 豫麦49-198 Yumai 49-198 CK 645.04±0.89a 36.17±0.69c 37.70±0.74bc 7637.31±42.14c BR 643.15±1.10a 40.03±0.94b 39.19±1.25b 8278.03±31.27b 周麦16 Zhoumai 16 CK 540.03±0.98b 40.89±0.63b 46.57±0.75a 8308.42±100.77b BR 542.46±1.24b 45.12±0.72a 47.85±0.21a 9356.10±98.08a 变异来源
Source of variation品种 Cultivar ** ** ** ** 处理 Treatment NS ** NS ** 品种×处理
Cultivar × treatmentNS NS NS * ‘豫麦49-198’为多穗型品种, ‘周麦16’为大穗型品种。CK: 喷施清水; BR: 喷施14-HBR。同列数据后不同小写字母表示不同品种不同处理间差异显著(P<0.05)。NS: 差异不显著; *: P<0.05; **: P<0.01。‘Yumai49-198’ is a multi-spike wheat cultivar, and ‘Zhoumai16’ is a large-spike wheat cultivar. CK: control with spraying water; BR: spraying 14-HBR treatment. Different lowercase letters in the same column show significant differences among treatments of different cultivars (P<0.05). NS: no significant difference. * and ** indicate significant differences at P<0.05 and P<0.01, respectively. 
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