Article(id=1321852236892848641, tenantId=1146029695717560320, journalId=1316781310003478668, issueId=1321852178331979976, articleNumber=null, orderNo=null, doi=10.11983/CBB24187, pmid=null, cstr=32102.14.CBB24187, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1733068800000, receivedDateStr=2024-12-02, revisedDate=null, revisedDateStr=null, acceptedDate=1742227200000, acceptedDateStr=2025-03-18, onlineDate=1793083732102, onlineDateStr=2026-10-27, pubDate=1762704000000, pubDateStr=2025-11-10, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1793083732102, onlineIssueDateStr=2026-10-27, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1793083732102, creator=13701087609, updateTime=1793083732102, updator=13701087609, issue=Issue{id=1321852178331979976, tenantId=1146029695717560320, journalId=1316781310003478668, year='2025', volume='60', issue='6', pageStart='859', pageEnd='1035', issueExtLink='null', onlineDate='null', pubDate='1762704000000', pubDateStr='2025-11-10', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=0, createTime=1793083718140, creator='13701087609', updateTime=1793083787009, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1321852467248223111, tenantId=1146029695717560320, journalId=1316781310003478668, issueId=1321852178331979976, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1321852467252417416, tenantId=1146029695717560320, journalId=1316781310003478668, issueId=1321852178331979976, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=931, endPage=943, ext={EN=ArticleExt(id=1321852237119341058, articleId=1321852236892848641, tenantId=1146029695717560320, journalId=1316781310003478668, language=EN, title=Heterosis in Yield and Its Physiological Mechanism of Changzagu Series Millet Hybrids, columnId=1317485691135706105, journalTitle=Chinese Bulletin of Botany, columnName=RESEARCH ARTICLES, runingTitle=null, highlight=null, articleAbstract=

INTRODUCTION: In agricultural production, the utilization of heterosis has brought significant benefits to society and economic development by markedly increasing crop yield, stress resistance, and quality. Foxtail millet (Setaria italica var. germanica), as an important coarse grain crop in the arid and semi-arid regions of northern China, holds a significant position in dry land ecological agriculture. However, the slow increase in the yield of foxtail millet has limited the further realization of its production potential. Utilizing heterosis has thus become one of the effective ways to increase the yield of foxtail millet. Nevertheless, research on the physiological and molecular mechanisms of its heterosis is still relatively weak, and the mechanism remains unclear. Therefore, to understand the physiological mechanisms of heterosis in foxtail millet is of great importance for improving the yield of hybrid varieties.
RATIONALE: The Changzagu series of foxtail millet hybrids (Changzagu466, Changzagu2922, and Changzagu333) exhibit significant heterosis in yield. In order to elucidate its mechanism, we systematically analyzed the yield advantages of these hybrids and their influencing factors by measuring yield-related traits and key physiological indicators of the hybrids and their parental lines.
RESULTS: Throughout the entire growth period, the chlorophyll content of the three hybrid varieties was higher than that of their parents. Among them, Changzagu466 exhibited the highest chlorophyll content at the jointing stage, reaching 13.86 mg∙g-1 FW. During the seedling and jointing stages, the root activity of the three hybrids was significantly higher than that of their parents. Specifically, Changzagu466 showed the highest root activity at the seedling stage, measuring 1.76 mg∙g-1∙h-1, which was 7.8 times and 5.5 times higher than its female and male parents, respectively. The root activity values of Changzagu2922 at the seedling stage were 0.38 and 0.66 mg∙g-1∙h-1 higher than its female and male parents, respectively. Meanwhile, Changzagu333 displayed pronounced advantages at the jointing stage, with root activity values 0.31 and 0.62 mg∙g-1∙h-1 higher than its female and male parents, respectively. In terms of yield-related traits, compared to their parents, the hybrids showed significant improvements in both grain filling rate and spikelet number. Changzagu466 reached its maximum grain filling rate of 1.58 g∙d-1 per panicle at 19 days after flowering. Both Changzagu466 and Changzagu333 had significantly higher spikelet numbers than their parents, while Changzagu2922 also showed a significant increase in spikelet numbers. In addition, the hybrid varieties also demonstrate certain advantages in root nitrogen accumulation and nitrogen translocation efficiency. Among them, Changzagu2922 exhibits the strongest root nitrogen accumulation advantage and the highest nitrogen translocation efficiency (nearly 56%), both are significantly higher than that of its male parent line M22.
CONCLUSION: The Changzagu series of foxtail millet hybrids effectively enhances photosynthetic capacity, nutrient absorption and utilization efficiency by significantly increasing chlorophyll content, root activity during the early growth stages, and root nitrogen accumulation. Meanwhile, the significant increase in grain filling rate and spikelet number of the hybrids further enhances both grain weight and grain number per panicle, ultimately achieving high yield.

Foxtail millet and parental trait comparison chart

, authors=Yurong Guo1, 2, Hong Liu2, Zhenhua Wang2, Gang Tian2, Xin Liu1, 2, Jie Guo1, Chunyong Li3, Huixia Li1, 2, *, authorsList=Yurong Guo, Hong Liu, Zhenhua Wang, Gang Tian, Xin Liu, Jie Guo, Chunyong Li, Huixia Li, authorCompany=null, correspAuthors=Huixia Li, authorNote=null, correspAuthorsNote=
* E-mail:
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阐明谷子(Setaria italica var. germanica)杂种优势的生理机制是提高谷子杂交种产量的重要基础。2023-2024年, 以3个高产谷子杂交种长杂谷466、长杂谷2922和长杂谷333及其亲本为供试材料, 测定杂交种及其亲本的产量性状及生理指标, 对杂交种的产量优势及其影响因子进行系统分析, 初步解析了杂种优势的生理机制。结果表明, 在整个生育期内, 3个杂交种的叶绿素含量均高于双亲, 其中长杂谷466在拔节期的叶绿素含量最高, 达13.86 mg∙g-1 FW。在苗期和拔节期, 3个杂交种的根系活力均显著高于双亲, 其中长杂谷466在苗期的根系活力最高, 为1.76 mg∙g-1∙h-1, 分别为其父母本的7.8倍和5.5倍; 长杂谷2922在苗期的根系活力较父母本分别高0.38和0.66 mg∙g-1∙h-1; 而长杂谷333在拔节期优势更为显著, 较父母本分别高0.31和0.62 mg∙g-1∙h-1。在产量形成性状方面, 与双亲相比, 杂交种的灌浆速率和颖花数均显著提高, 长杂谷466在花后19天灌浆速率达最大值, 为1.58 g∙d-1 per panicle; 长杂谷466和长杂谷333颖花数极显著高于其亲本, 长杂谷2922颖花数显著增加。此外, 杂交种根系的氮素积累量和氮素转运效率也表现出一定的优势, 其中长杂谷2922根系氮素积累优势最大且氮素转运效率(近56%)最高, 显著高于其父本M22。综上表明, 长杂谷系列杂交种通过提高光合能力、养分吸收利用效率、籽粒灌浆速率和颖花数实现产量增加。

, authors=郭宇荣1, 2, 刘红2, 王振华2, 田岗2, 刘鑫1, 2, 郭杰1, 李春勇3, 李会霞1, 2, *, authorsList=郭宇荣, 刘红, 王振华, 田岗, 刘鑫, 郭杰, 李春勇, 李会霞, authorCompany=null, correspAuthors=李会霞, authorNote=null, correspAuthorsNote=
* 李会霞, 山西农业大学谷子研究所研究员, 谷子杂优育种科室负责人, 山西省谷子体系岗位专家, 硕士生导师。30多年来其研究团队致力于谷子杂种优势利用研究及新品种选育, 共选育谷子品种14个, 5个为谷子杂交种并全部转让, 转让费累计达120万元。其中长杂2号为我国较早自主培育的中晚熟谷子杂交种。2012年获山西省科技进步二等奖。近3年, 杂交种生产推广面积超过1.3×104 hm2, 平均产量居当地较高水平。E-mail:
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The materials Changzagu466, its paternal parent is K34 and its maternal parent is Gu3A; Changzagu2922, its paternal parent is M22 and its maternal parent is Jin29A; Changzagu333, its paternal parent is K410 and its maternal parent is Gao51A. Different lowercase letters indicate significant differences at the 0.05 level.

, figureFileSmall=4R1/oHpKniSb7FCbyvEc8w==, figureFileBig=TIVzx6DblJ4E6SxW1YTOYg==, tableContent=null), ArticleFig(id=1321852246317449813, tenantId=1146029695717560320, journalId=1316781310003478668, articleId=1321852236892848641, language=CN, label=图1, caption=不同生长时期谷子叶绿素含量

供试材料长杂谷466, 其父本为K34, 母本为谷3A; 长杂谷2922, 其父本为M22, 母本为晋29A; 长杂谷333, 其父本为K410, 母本为高51A。不同小写字母表示在0.05水平差异显著。

, figureFileSmall=4R1/oHpKniSb7FCbyvEc8w==, figureFileBig=TIVzx6DblJ4E6SxW1YTOYg==, tableContent=null), ArticleFig(id=1321852246531359318, tenantId=1146029695717560320, journalId=1316781310003478668, articleId=1321852236892848641, language=EN, label=Figure 2, caption=Root activity of millet at different growth stages

The materials are the same as shown in Figure 1. Different lowercase letters indicate significant differences at the 0.05 level.

, figureFileSmall=4F+XTTL9CluOH363FTGqNQ==, figureFileBig=188w44rx+QmoiUzdT+vDkg==, tableContent=null), ArticleFig(id=1321852248234246743, tenantId=1146029695717560320, journalId=1316781310003478668, articleId=1321852236892848641, language=CN, label=图2, caption=不同生长时期谷子根系活力

供试材料同图1。不同小写字母表示在0.05水平差异显著。

, figureFileSmall=4F+XTTL9CluOH363FTGqNQ==, figureFileBig=188w44rx+QmoiUzdT+vDkg==, tableContent=null), ArticleFig(id=1321852248355881560, tenantId=1146029695717560320, journalId=1316781310003478668, articleId=1321852236892848641, language=EN, label=Figure 3, caption=Heterosis in physiological index of millet

RA: Root activity; Chl: Chlorophyll content; S: Seedling stage; J: Jointing stage; FH: Full heading stage; G: Grouting stage; H: Over-parent advantage; L: Low-parent advantage; M: Medium-parent advantage. The scale color indicates the dominance range (-100-100); red represents the positive dominance level and blue represents the negative dominance level, and the dominance color beyond the range is indicated by the deepest color.

, figureFileSmall=ootBfdYz1tmU6PUDwk0nLw==, figureFileBig=zL9XHQpA+mIChg7Q1qjpkg==, tableContent=null), ArticleFig(id=1321852248418796121, tenantId=1146029695717560320, journalId=1316781310003478668, articleId=1321852236892848641, language=CN, label=图3, caption=谷子生理指标杂种优势

RA: 根系活力; Chl: 叶绿素含量; S: 苗期; J: 拔节期; FH: 齐穗期; G: 灌浆期; H: 超亲优势; L: 低亲优势; M: 中亲优势。标尺颜色指示优势度范围(-100-100); 红色代表正向优势水平, 蓝色代表负向优势水平, 超出范围的优势度颜色以最深颜色表示。

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(A) Nitrogen accumulation at seedling stage; (B) Nitrogen accumulation at jointing stage; (C) Nitrogen accumulation at full heading stage; (D) Nitrogen accumulation at grouting stage; (E) Nitrogen accumulation at maturity stage; (F) Nitrogen transport efficiency. Different lowercase letters indicate significant differences at the 0.05 level.

, figureFileSmall=b0WCGQ9fclY9F0N7fLuFHg==, figureFileBig=R2/mVSJS4N3H0FiVzXqALg==, tableContent=null), ArticleFig(id=1321852248779506269, tenantId=1146029695717560320, journalId=1316781310003478668, articleId=1321852236892848641, language=CN, label=图5, caption=不同生育期谷子各器官氮素积累量及氮素转运效率

(A) 苗期氮素积累量; (B) 拔节期氮素积累量; (C) 齐穗期氮素积累量; (D) 灌浆期氮素积累量; (E) 成熟期氮素积累量; (F) 氮素转运效率。不同小写字母表示在0.05水平差异显著。

, figureFileSmall=b0WCGQ9fclY9F0N7fLuFHg==, figureFileBig=R2/mVSJS4N3H0FiVzXqALg==, tableContent=null), ArticleFig(id=1321852248859198046, tenantId=1146029695717560320, journalId=1316781310003478668, articleId=1321852236892848641, language=EN, label=Figure 6, caption=Heterosis in agronomic traits of millet

The abbreviations of traits are the same as shown in Table 2. H: Over-parent advantage; L: Low-parent advantage; M: Medium-parent advantage. The scale color indicates the dominance range (-100-100); red represents the positive dominance level and blue represents the negative dominance level, and the dominance color beyond the range is indicated by the deepest color.

, figureFileSmall=hdld98zFku60Q9VYsOXtCg==, figureFileBig=sPwkzaO8mGUyAlPwUcrTrg==, tableContent=null), ArticleFig(id=1321852248951472735, tenantId=1146029695717560320, journalId=1316781310003478668, articleId=1321852236892848641, language=CN, label=图6, caption=谷子农艺性状杂种优势

性状缩写同表2。H: 超亲优势; L: 低亲优势; M: 中亲优势。标尺颜色指示优势度范围(-100-100); 红色代表正向优势水平, 蓝色代表负向优势水平, 超出范围的优势度颜色以最深颜色表示。

, figureFileSmall=hdld98zFku60Q9VYsOXtCg==, figureFileBig=sPwkzaO8mGUyAlPwUcrTrg==, tableContent=null), ArticleFig(id=1321852249018581600, tenantId=1146029695717560320, journalId=1316781310003478668, articleId=1321852236892848641, language=EN, label=Table 1, caption=

Over-parent advantage of dry matter accumulation of millet in different grouting stages

, figureFileSmall=null, figureFileBig=null, tableContent=
Variety Dry matter accumulation (g∙d-1 per panicle) Over-parent heterosis (%)
0-t1 t1-t2 t2-t3 0-t1 t1-t2 t2-t3
Changzagu466 0.53 1.39 0.39 63 28 29
K34 0.32 1.09 0.31
Changzagu2922 0.43 1.09 0.30 27 44 38
M22 0.34 0.76 0.21
Changzagu333 0.49 1.20 0.33 30 36 79
K410 0.38 0.88 0.25
), ArticleFig(id=1321852249115050593, tenantId=1146029695717560320, journalId=1316781310003478668, articleId=1321852236892848641, language=CN, label=表1, caption=

不同灌浆时期谷子干物质积累量的超亲优势

, figureFileSmall=null, figureFileBig=null, tableContent=
Variety Dry matter accumulation (g∙d-1 per panicle) Over-parent heterosis (%)
0-t1 t1-t2 t2-t3 0-t1 t1-t2 t2-t3
Changzagu466 0.53 1.39 0.39 63 28 29
K34 0.32 1.09 0.31
Changzagu2922 0.43 1.09 0.30 27 44 38
M22 0.34 0.76 0.21
Changzagu333 0.49 1.20 0.33 30 36 79
K410 0.38 0.88 0.25
), ArticleFig(id=1321852249198936674, tenantId=1146029695717560320, journalId=1316781310003478668, articleId=1321852236892848641, language=EN, label=Table 2, caption=

Analysis of yield-related agronomic traits of millet

, figureFileSmall=null, figureFileBig=null, tableContent=
), ArticleFig(id=1321852249278628451, tenantId=1146029695717560320, journalId=1316781310003478668, articleId=1321852236892848641, language=CN, label=表2, caption=

谷子产量相关农艺性状分析

, figureFileSmall=null, figureFileBig=null, tableContent=
), ArticleFig(id=1321852249349931620, tenantId=1146029695717560320, journalId=1316781310003478668, articleId=1321852236892848641, language=EN, label=Table 3, caption=

Yield and yieid components of dfent milet vanetes

, figureFileSmall=null, figureFileBig=null, tableContent=
), ArticleFig(id=1321852249442206309, tenantId=1146029695717560320, journalId=1316781310003478668, articleId=1321852236892848641, language=CN, label=表3, caption=

不同谷子品种产量及产量构成因素

, figureFileSmall=null, figureFileBig=null, tableContent=
), ArticleFig(id=1321852249555452518, tenantId=1146029695717560320, journalId=1316781310003478668, articleId=1321852236892848641, language=EN, label=Table 4, caption=

Yield and yield component factors for over-parent advantage in millet

, figureFileSmall=null, figureFileBig=null, tableContent=
Traits Changzagu- 466 Changzagu- 2922 Changzagu- 333
PW (%) 37 28 61
TGW (%) 1 6 2
GWPP (%) 28 39 65
GP (%) 27 31 61
SP (%) 35 30 44
SSR (%) -6 3 11
PY (%) 20 15 50
), ArticleFig(id=1321852249672893031, tenantId=1146029695717560320, journalId=1316781310003478668, articleId=1321852236892848641, language=CN, label=表4, caption=

谷子产量及产量构成因素超亲优势

, figureFileSmall=null, figureFileBig=null, tableContent=
Traits Changzagu- 466 Changzagu- 2922 Changzagu- 333
PW (%) 37 28 61
TGW (%) 1 6 2
GWPP (%) 28 39 65
GP (%) 27 31 61
SP (%) 35 30 44
SSR (%) -6 3 11
PY (%) 20 15 50
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长杂谷系列谷子杂交种产量优势及其生理机制
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郭宇荣 1, 2 , 刘红 2 , 王振华 2 , 田岗 2 , 刘鑫 1, 2 , 郭杰 1 , 李春勇 3 , 李会霞 1, 2, *
植物学报 | 研究论文 2025,60(6): 931-943
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植物学报 |研究论文 2025 , 60 (6) : 931 -943
长杂谷系列谷子杂交种产量优势及其生理机制
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郭宇荣1, 2, 刘红2, 王振华2, 田岗2, 刘鑫1, 2, 郭杰1, 李春勇3, 李会霞1, 2, *
作者信息
  • 1 山西农业大学农学院, 太谷 030801
  • 2 山西农业大学谷子研究所, 长治 046011
  • 3 山西农业大学山西功能农产品检验检测中心, 太原 030031
通讯作者:
* 李会霞, 山西农业大学谷子研究所研究员, 谷子杂优育种科室负责人, 山西省谷子体系岗位专家, 硕士生导师。30多年来其研究团队致力于谷子杂种优势利用研究及新品种选育, 共选育谷子品种14个, 5个为谷子杂交种并全部转让, 转让费累计达120万元。其中长杂2号为我国较早自主培育的中晚熟谷子杂交种。2012年获山西省科技进步二等奖。近3年, 杂交种生产推广面积超过1.3×104 hm2, 平均产量居当地较高水平。E-mail:
Heterosis in Yield and Its Physiological Mechanism of Changzagu Series Millet Hybrids
Yurong Guo1, 2, Hong Liu2, Zhenhua Wang2, Gang Tian2, Xin Liu1, 2, Jie Guo1, Chunyong Li3, Huixia Li1, 2, *
Affiliations
  • 1 College of Agronomy, Shanxi Agricultural University, Taigu 030801, China
  • 2 Millet Research Institute, Shanxi Agricultural University, Changzhi 046011, China
  • 3 Shanxi Center for Testing of Functional Agro-Products, Shanxi Agricultural University, Taiyuan 030031, China
  • Corresponding Author:
出版时间: 2025-11-10 doi: 10.11983/CBB24187
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阐明谷子(Setaria italica var. germanica)杂种优势的生理机制是提高谷子杂交种产量的重要基础。2023-2024年, 以3个高产谷子杂交种长杂谷466、长杂谷2922和长杂谷333及其亲本为供试材料, 测定杂交种及其亲本的产量性状及生理指标, 对杂交种的产量优势及其影响因子进行系统分析, 初步解析了杂种优势的生理机制。结果表明, 在整个生育期内, 3个杂交种的叶绿素含量均高于双亲, 其中长杂谷466在拔节期的叶绿素含量最高, 达13.86 mg∙g-1 FW。在苗期和拔节期, 3个杂交种的根系活力均显著高于双亲, 其中长杂谷466在苗期的根系活力最高, 为1.76 mg∙g-1∙h-1, 分别为其父母本的7.8倍和5.5倍; 长杂谷2922在苗期的根系活力较父母本分别高0.38和0.66 mg∙g-1∙h-1; 而长杂谷333在拔节期优势更为显著, 较父母本分别高0.31和0.62 mg∙g-1∙h-1。在产量形成性状方面, 与双亲相比, 杂交种的灌浆速率和颖花数均显著提高, 长杂谷466在花后19天灌浆速率达最大值, 为1.58 g∙d-1 per panicle; 长杂谷466和长杂谷333颖花数极显著高于其亲本, 长杂谷2922颖花数显著增加。此外, 杂交种根系的氮素积累量和氮素转运效率也表现出一定的优势, 其中长杂谷2922根系氮素积累优势最大且氮素转运效率(近56%)最高, 显著高于其父本M22。综上表明, 长杂谷系列杂交种通过提高光合能力、养分吸收利用效率、籽粒灌浆速率和颖花数实现产量增加。

杂交谷子  /  生理指标  /  籽粒灌浆速率  /  农艺性状  /  产量

INTRODUCTION: In agricultural production, the utilization of heterosis has brought significant benefits to society and economic development by markedly increasing crop yield, stress resistance, and quality. Foxtail millet (Setaria italica var. germanica), as an important coarse grain crop in the arid and semi-arid regions of northern China, holds a significant position in dry land ecological agriculture. However, the slow increase in the yield of foxtail millet has limited the further realization of its production potential. Utilizing heterosis has thus become one of the effective ways to increase the yield of foxtail millet. Nevertheless, research on the physiological and molecular mechanisms of its heterosis is still relatively weak, and the mechanism remains unclear. Therefore, to understand the physiological mechanisms of heterosis in foxtail millet is of great importance for improving the yield of hybrid varieties.
RATIONALE: The Changzagu series of foxtail millet hybrids (Changzagu466, Changzagu2922, and Changzagu333) exhibit significant heterosis in yield. In order to elucidate its mechanism, we systematically analyzed the yield advantages of these hybrids and their influencing factors by measuring yield-related traits and key physiological indicators of the hybrids and their parental lines.
RESULTS: Throughout the entire growth period, the chlorophyll content of the three hybrid varieties was higher than that of their parents. Among them, Changzagu466 exhibited the highest chlorophyll content at the jointing stage, reaching 13.86 mg∙g-1 FW. During the seedling and jointing stages, the root activity of the three hybrids was significantly higher than that of their parents. Specifically, Changzagu466 showed the highest root activity at the seedling stage, measuring 1.76 mg∙g-1∙h-1, which was 7.8 times and 5.5 times higher than its female and male parents, respectively. The root activity values of Changzagu2922 at the seedling stage were 0.38 and 0.66 mg∙g-1∙h-1 higher than its female and male parents, respectively. Meanwhile, Changzagu333 displayed pronounced advantages at the jointing stage, with root activity values 0.31 and 0.62 mg∙g-1∙h-1 higher than its female and male parents, respectively. In terms of yield-related traits, compared to their parents, the hybrids showed significant improvements in both grain filling rate and spikelet number. Changzagu466 reached its maximum grain filling rate of 1.58 g∙d-1 per panicle at 19 days after flowering. Both Changzagu466 and Changzagu333 had significantly higher spikelet numbers than their parents, while Changzagu2922 also showed a significant increase in spikelet numbers. In addition, the hybrid varieties also demonstrate certain advantages in root nitrogen accumulation and nitrogen translocation efficiency. Among them, Changzagu2922 exhibits the strongest root nitrogen accumulation advantage and the highest nitrogen translocation efficiency (nearly 56%), both are significantly higher than that of its male parent line M22.
CONCLUSION: The Changzagu series of foxtail millet hybrids effectively enhances photosynthetic capacity, nutrient absorption and utilization efficiency by significantly increasing chlorophyll content, root activity during the early growth stages, and root nitrogen accumulation. Meanwhile, the significant increase in grain filling rate and spikelet number of the hybrids further enhances both grain weight and grain number per panicle, ultimately achieving high yield.

Foxtail millet and parental trait comparison chart

hybrid millet  /  physiological index  /  grain filling rate  /  agronomic traits  /  yield
郭宇荣, 刘红, 王振华, 田岗, 刘鑫, 郭杰, 李春勇, 李会霞. 长杂谷系列谷子杂交种产量优势及其生理机制. 植物学报, 2025 , 60 (6) : 931 -943 . DOI: 10.11983/CBB24187
Yurong Guo, Hong Liu, Zhenhua Wang, Gang Tian, Xin Liu, Jie Guo, Chunyong Li, Huixia Li. Heterosis in Yield and Its Physiological Mechanism of Changzagu Series Millet Hybrids[J]. Chinese Bulletin of Botany, 2025 , 60 (6) : 931 -943 . DOI: 10.11983/CBB24187
杂种优势(heterosis)是杂交后代在长势、抗逆性、适应性、产量和品质等方面优于亲本的现象(肖层林等, 2015), 已广泛应用于作物育种。研究表明, 作物最终的产量由光合作用、同化产物运输和分配以及籽粒灌浆特性共同决定(宋慧等, 2022)。谷子(Setaria italica var. germanica)花后形成的干物质占谷物产量中干物质的60%-90% (黄玲等, 2013); 而小麦(Triticum aestivum)灌浆中后期持续时间的延长使籽粒重量增加(Zhang et al., 2024)。这表明谷物灌浆过程对作物高产至关重要。此外, 单穗重、穗粒重、株高、穗长、穗粗、出谷率以及千粒重也是影响谷子产量的关键因素(史根生等, 2016; 张霞和冯永忠, 2017; 魏玮等, 2019; 周花等, 2020)。因此, 作物产量杂种优势直接评价指标主要包括穗数、穗粒数和千粒重等。
解析杂种优势大多采用配合力、相关性和遗传距离等分析方法。吴立东等(2024)利用遗传距离和配合力方法预测了辣椒(Capsicum annuum)辣度的杂种优势。钏兴宽等(2024)对高原两系杂交水稻(Oryza sativa)主要经济性状杂种优势进行了相关性分析。在杂种优势机理研究方面, 对水稻和玉米(Zea mays)等作物的研究比较深入。苟亚军等(2023)从基因角度对籼粳杂交水稻杂种优势利用的关键性状进行遗传解析。侯美(2023)通过构建玉米骨干亲本自交系的泛基因组和相关性分析, 挖掘出玉米杂种优势以及杂种优势群形成相关基因。上述研究为深入理解和利用作物杂种优势提供了重要理论依据和实践指导。
多年来山西谷子单产提升较慢, 目前平均产量仅为3 000 kg∙hm-2。若要大幅提高谷子产量, 利用杂交种是有效途径之一。准确评估杂种优势的前提是对杂种优势进行度量, 常用的度量指标包括超亲优势、低亲优势、中亲优势及超标优势等。史关燕等(2021)对18个谷子杂交组合及其亲本产量等相关性状配合力进行分析, 发现产量性状具有较强的中亲优势, 部分组合还表现出较高的超亲优势。李素英等(2023)系统回顾了中国谷子杂种优势利用研究的发展历程, 探讨了杂种优势利用存在的问题, 强调基础研究与应用研究须紧密结合。然而, 谷子杂种优势生理与分子机制方面的研究较为薄弱, 目前对杂种优势的形成机理认识不清, 限制了突破性强优势组合的选育。因此, 有必要开展谷子杂种优势的机制分析, 研究与产量相关的外部形态特征和内部生理指标及其相关性, 从理论上指导鉴定筛选高配合力的亲本, 提高强优势杂交种的选育概率。
本研究以山西农业大学谷子研究所选育的3个高产杂交种长杂谷466、长杂谷2922和长杂谷333及其亲本为实验材料, 在各生育阶段对谷子的农艺性状和生理指标进行调查, 并对谷子杂种优势的生理机制进行分析, 旨在为选育强优势谷子杂交种提供理论指导。
供试材料为山西农业大学谷子研究所选育的3个谷子(Setaria italica var. germanica (Mill.) Schred.)高产杂交组合长杂谷466 (谷3A × K34)、长杂谷2922 (晋29A × M22)和长杂谷333 (高51A × K410)及6个亲本, 母本谷3A、晋29A和高51A均为高度雄性不育系。
2023-2024年在山西农业大学谷子研究所试验田进行试验。试验地海拔923 m, 土壤为石灰质褐土。采用随机区组设计, 设3次重复, 共27个小区, 每小区4行, 行距为0.45 m, 行长7 m, 小区面积为12.6 m2。试验地前茬作物为玉米。4月旋耕田地、施肥及备种。施肥量为氮肥105 kg∙hm-2, 磷肥63 kg∙hm-2, 钾肥42 kg∙hm-2。 肥料全部在播种前底施。5月中旬播种, 10月初收获。田间管理同丰产田管理。
在谷子4个生长时期(苗期、拔节期、齐穗期和灌浆期), 每小区各取3株长势一致的植株, 测定叶绿素含量及根系活力。采用乙醇浸提法(高俊凤, 2006)测定叶绿素含量。用TTC法(陈云风, 2015)测定根系活力。
谷子抽穗后, 选同一天抽穗的植株进行挂牌标记, 从抽穗后3天开始每隔10天取1次样, 摘取10个谷穗, 称鲜重后放入烘箱, 105°C杀青30分钟, 80°C烘干至恒重, 称量干重。用Logistic (Du et al., 2024)方程拟合谷子灌浆动态。
$W=\frac{K}{1+\boldsymbol{a} \times \mathbf{e}^{(-b t)}}$
式中, W代表花后籽粒干重, K为理论籽粒最大干重, e为常数, t为时间, a和b为参数。
$\text { 最大灌浆速率 } V_{\max }=K b / 4$
$\text { 达最大灌浆速率的时间 } t_{\text {max }}=\ln a / b$
灌浆速率曲线的2个拐点坐标值t1和t2,
$t_{1}=-\frac{\ln \left[\frac{2+\sqrt{3}}{a}\right]}{b}, t_{2}=-\ln [(2-\sqrt{3}) / a] / b{\tiny } $
谷子成熟后, 每个小区取长势基本一致的10株植株进行表型性状调查, 统计测量叶片数、株高、根长、根分布及顶三叶面积。植株风干后再测根干重、地上草重(除穗)、单株穗重、穗粒重、穗粒数、千粒重和颖花数等。小区实收测产。
在谷子各生育期(苗期、拔节期、齐穗期、灌浆期和成熟期), 每小区取3株, 分为根、茎+鞘、叶和穗4部分, 105°C杀青30分钟, 80°C烘干至恒重, 称量各器官干物质重。将各器官样品粉碎过筛后, 加浓硫酸消煮, 用凯氏定氮仪(K9840)测全氮含量。计算氮素积累量及氮素转运效率(朱灿灿等, 2023)。
各器官氮素积累量=各器官氮含量×各器官干物质积累量;
花前氮素积累量=花前干物质积累量×开花期植株氮含量;
成熟期氮素积累量=成熟期干物质积累量×成熟期植株氮含量;
氮素转运效率=((花前氮素积累量-成熟期营养器官氮素积累量)/花前氮素积累量)×100%。
使用Excel 2019和SPSS 25软件进行数据处理与分析。用Origin 2022软件作图。
研究结果显示, 随着生育进程的推进, 谷子叶绿素含量呈先升高后降低趋势(图1)。在整个生育期内, 杂交种的叶绿素含量均高于其父母本。长杂谷466的叶绿素含量在拔节期达到峰值(13.86 mg∙g-1 FW), 其余8个品种叶绿素含量的峰值则出现在齐穗期。长杂谷466在拔节期叶绿素含量显著高于其父母本(4.3和5.88 mg∙g-1 FW), 在齐穗期叶绿素含量也显著高于其父母本(0.55和0.98 mg∙g-1 FW)。在齐穗期和灌浆期长杂谷2922叶绿素含量显著高于双亲, 齐穗期分别比父母本高1.85和1.4 mg∙g-1 FW, 灌浆期分别比父母本高1.97和1.08 mg∙g-1 FW。在苗期和拔节期长杂谷333叶绿素含量显著高于双亲, 苗期分别比父母本高1.47和3.67 mg∙g-1 FW, 拔节期分别比父母本高2.59和1.78 mg∙g-1 FW。
研究结果表明, 随着生育进程的推进, 谷子根系活力呈先升高后降低趋势。长杂谷466根系活力在苗期达到峰值, 为1.76 mg·g-1 FW, 显著高于其它时期的所有品种(图2)。3个杂交种根系活力在苗期和拔节期均显著高于双亲。长杂谷466在苗期根系活力是父本K34的7.8倍, 是母本谷3A的5.5倍。长杂谷2922在苗期和拔节期根系活力差异不显著, 苗期较父母本分别高0.38和0.66 mg∙g-1∙h-1, 拔节期分别高0.38和0.26 mg∙g-1∙h-1。长杂谷333在苗期根系活力分别比父母本高0.31和0.62 mg∙g-1∙h-1, 拔节期分别高0.41和0.42 mg∙g-1∙h-1。
本研究显示, 3个杂交种的叶绿素含量在各生长时期普遍具有杂种优势(图3)。长杂谷466在拔节期的叶绿素含量表现出较强的杂种优势, 其超亲优势、低亲优势和中亲优势分别为45%、74%和58%。长杂谷333在苗期的叶绿素含量表现出56%的低亲优势。此外, 3个杂交种在苗期的根系活力均表现出极强的杂种优势。长杂谷466超亲优势、低亲优势和中亲优势分别达452%、677%和550%, 杂种优势非常明显。杂交种在拔节期根系活力仍表现出较强的杂种优势, 但在齐穗期和灌浆期根系活力低于双亲。
杂交种及其父本花后灌浆期籽粒干物质积累呈S型生长曲线(图4), 花后10-40天干物质迅速积累, 40天后干物质积累速率明显减缓。长杂谷466在花后19天达到最大灌浆速率(1.58 g∙d-1 per panicle), 其父本K34在花后20天达到最大灌浆速率(1.24 g∙d-1 per panicle)。长杂谷2922在花后17天达到最大灌浆速率(1.24 g∙d-1 per panicle), 其父本M22在花后14天达到最大灌浆速率(0.86 g∙d-1 per panicle)。长杂谷333在花后23天达到最大灌浆速率(1.36 g∙d-1 per panicle), 其父本K410在花后21天达到最大灌浆速率(1.00 g∙d-1 per panicle)。综上, 3个杂交种在不同灌浆时期的干物质积累量均高于其父本, 表现出超亲优势。
利用Logistic生长曲线方程拟合谷子灌浆曲线, 拟合相关系数r均大于0.98, 达极显著水平, 表明9个品种的灌浆特性均符合Logistic生长规律。对Logistic方程求导后, 可确定灌浆过程的2个拐点, 从而将灌浆期分为灌浆前期、灌浆中期和灌浆后期。长杂谷466、长杂谷2922和长杂谷333在灌浆期不同阶段均表现出显著的超亲优势(表1)。长杂谷466在灌浆前期的超亲优势达63%, 长杂谷2922在灌浆中期的超亲优势为44%, 长杂谷333在灌浆后期的超亲优势高达79%。综合分析表明, 长杂谷333在整个籽粒灌浆过程中超亲优势平均值显著高于其它2个杂交种。
从不同生育期各部位氮素积累量(图5A-E)来看, 杂交种在各生育期总氮素积累量高于其父本。各器官氮素积累量从苗期到齐穗期整体表现为叶>茎>根, 开始灌浆时表现为穗>叶≈茎>根。杂交种在各生育期根系氮素积累量均表现显著优势, 长杂谷466苗期至灌浆期根系氮素积累量显著高于其父本, 长杂谷2922在整个生育期根系氮素积累量均高于其父本, 长杂谷333在苗期和拔节期根系氮素积累量高于其父本。研究表明杂交种根系氮素吸收利用效率具有显著的杂种优势。
氮素转运效率表现出杂交种普遍高于其父本(图5F)。长杂谷466与K34之间氮素转运效率无显著差异。长杂谷2922氮素转运效率最高, 约为56%, 且显著高于其父本M22。长杂谷333的氮素转运效率约为35%, 其父本K410的氮素转运效率最低, 约为14%。 总体来看, 长杂谷2922在氮素转运效率上表现最佳, 显著高于其它品种。
F1代及双亲产量相关农艺性状方差分析显示, 长杂谷466最长根长、根集中分布深度、叶长、叶宽及根干重等农艺性状均优于双亲, 且最长根长显著高于双亲(表2)。长杂谷2922最长根长、根集中分布深度、顶三叶面积、叶宽、根干重、根冠比及根草比(根重/茎叶重)等农艺性状优于双亲, 其中根集中分布深度极显著高于双亲, 根干重显著高于双亲; 根草比极显著高于其父本, 显著高于其母本。长杂谷333最长根长、根集中分布深度、顶三叶面积、叶长及叶宽等农艺性状优于双亲, 其中叶长与双亲存在显著或极显著差异, 叶宽与其母本差异显著。
研究结果显示, 3个杂交种的农艺性状超亲优势不明显(图6)。长杂谷466的最长根长及根干重表现出明显的超亲优势, 分别为20%和33%; 长杂谷2922根集中分布深度与根干重超亲优势分别为35%和22%, 根草比超亲优势为47%; 长杂谷333叶长超亲优势为23%。3个杂交种普遍具有低亲优势。长杂谷466的根草比和单株草重低亲优势分别为60%和70%; 长杂谷2922顶三叶面积低亲优势为91%, 根干重低亲优势高达172%; 长杂谷333除根冠比和根草比外, 其它性状均表现较强的低亲优势。在中亲优势方面, 长杂谷466各农艺性状均表现不明显; 长杂谷2922除叶片数外, 其它农艺性状普遍具有中亲优势; 长杂谷333除根冠比和根草比外, 其它农艺性状也普遍表现中亲优势。
本研究显示, 3个杂交种的千粒重与亲本相比均无显著差异(表3)。而在单穗重、单穗粒重、单穗粒数、颖花数及小区产量等性状上, F1代与其父本之间均存在显著或极显著差异。此外, 长杂谷466的颖花数极显著高于双亲, 但其结实率却极显著低于双亲。长杂谷333的颖花数与亲本差异极显著, 结实率与其父本差异显著, 比父本高7.72%。
由于3个杂交组合的母本均为高度雄性不育系, 结实率仅有5%-10%, 因此超亲优势只与父本进行比较。长杂谷466在产量及产量构成因素上的超亲优势值为-6%-37%, 其中单穗重超亲优势为37%; 颖花数超亲优势达35%。长杂谷2922在产量及产量构成因素上的超亲优势值为3%-39%, 其中单穗粒重超亲优势达39%。长杂谷333在产量及产量构成因素上的超亲优势值为11%-65%, 其中单穗重、单穗粒重和单穗粒数超亲优势分别为61%、65%和61%。3个杂交种的小区产量超亲优势分别为20%、15%和50%, 产量优势明显(表4)。
产量是衡量谷子杂种优势利用效果的重要指标。杨艳君等(2012)研究表明, 张杂谷5号的产量与单穗粒数高度相关。李素英等(2018)发现谷子穗粒重和出谷率的杂种优势与产量杂种优势相关性最强, 其次是穗长、单穗重和穗粗。本研究表明, 与亲本相比, 3个杂交种的单穗重、单穗粒重及单穗粒数均表现显著或极显著差异, 表明超亲优势明显。杂交种的颖花数显著增加, 使其单穗粒数增加, 从而提高产量。Li等(2016)认为每穗小花数是决定两系杂交水稻产量优势的核心。而长杂谷466虽然结实率显著低于其父本, 但颖花数显著增加, 可能抵消了结实率降低对产量的影响, 从而确保杂交种的产量。
籽粒灌浆期是决定谷物粒重和品质的重要阶段(王蕾等, 2024)。刘童等(2024)对91个小麦杂交种灌浆特性的研究表明, 89%的杂交种灌浆时间更长, 导致籽粒增大。徐田军等(2021)的研究表明, 玉米杂交种的籽粒灌浆速率与产量呈极显著正相关。本研究表明, 3个杂交种在不同的灌浆期穗部干物质积累量均高于其父本, 灌浆速率提高可能使单穗粒重增加。该结果与前人研究结果(卢海博等, 2016)基本一致。
根系作为植物吸收养分的关键器官, 对谷子地上部生长发育产生重要影响(Wu et al., 2024)。Shalby等(2021)对甘蓝型油菜(Brassica napus‌)的研究表明, 杂交种早期生物量增加及根系活力增强在杂种优势形成中起关键作用。本研究表明, 长杂谷466和长杂谷2922在根长、根集中分布深度及根干重方面均表现出显著超亲优势, 且3个杂交种在生育前期的根系活力均表现极强的杂种优势, 这与皮凯等(2022)对烟草(Nicotiana tabacum)根系杂种优势的研究结果一致。同时, 杂交种在苗期和拔节期的根系氮素积累量显著高于其父本, 表明杂交谷子在生育前期的氮素吸收方面具有显著的杂种优势, 这种优势可能来源于杂交后代在遗传上的改良, 使其在氮素的吸收、运输和利用上更为高效。值得注意的是, 长杂谷2922的根草比(根重/茎叶重)表现出较强的杂种优势, 根草比越大, 根系越发达, 植株越健壮, 从而能更好地吸收和利用养分, 促进产量和品质的提升(彭翠兰, 2020)。
此外, 叶片作为植物光合作用的主要器官, 其叶绿素含量是衡量植株光合作用的重要指标, 并与植株产量密切相关(唐兴旺等, 2020; 任青等, 2021)。本研究表明, 3个杂交种在各生育期的叶绿素含量均高于其亲本, 杂种优势显著。长杂谷333在顶三叶面积、叶长和叶宽等性状上均表现出杂种优势。
长杂谷系列杂交种在多个生理和产量相关性状上表现出显著优势。在生理上, 杂交种具有更高的叶绿素含量, 长杂谷466在拔节期的叶绿素含量最高; 且杂交种生育前期根系活力强, 长杂谷466苗期根系活力尤为突出, 养分吸收能力强, 为生长发育提供了良好的物质基础。在产量形成性状上, 杂交种灌浆速率显著增高, 长杂谷333平均灌浆速率优于长杂谷466和长杂谷2922; 同时杂交种颖花数显著增加, 长杂谷466和长杂谷333的颖花数较双亲极显著增加。在氮素利用方面, 杂交种在根系氮素积累量和氮素转运效率方面表现出一定优势, 长杂谷2922在全生育期根系氮素积累量显著高于其父本M22, 且比长杂谷466和长杂谷333的氮素转运效率高。综上, 长杂谷系列杂交种通过增强光合能力、提高养分吸收效率、优化灌浆特性及增加颖花数等途径, 表现出显著的产量优势。研究结果为谷子高产育种提供了理论依据和实践指导。
郭宇荣: 实验数据调查与处理, 撰写论文初稿; 刘红, 王振华, 田岗, 刘鑫, 郭杰, 李春勇: 协助实验数据调查与处理; 李会霞: 提供理论和技术指导, 修改论文。
  • 山西农业大学科技创新提升工程(CXGC2023098)
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2025年第60卷第6期
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doi: 10.11983/CBB24187
  • 接收时间:2024-12-02
  • 首发时间:2026-10-27
  • 出版时间:2025-11-10
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  • 收稿日期:2024-12-02
  • 录用日期:2025-03-18
基金
山西农业大学科技创新提升工程(CXGC2023098)
作者信息
    1 山西农业大学农学院, 太谷 030801
    2 山西农业大学谷子研究所, 长治 046011
    3 山西农业大学山西功能农产品检验检测中心, 太原 030031

通讯作者:

* 李会霞, 山西农业大学谷子研究所研究员, 谷子杂优育种科室负责人, 山西省谷子体系岗位专家, 硕士生导师。30多年来其研究团队致力于谷子杂种优势利用研究及新品种选育, 共选育谷子品种14个, 5个为谷子杂交种并全部转让, 转让费累计达120万元。其中长杂2号为我国较早自主培育的中晚熟谷子杂交种。2012年获山西省科技进步二等奖。近3年, 杂交种生产推广面积超过1.3×104 hm2, 平均产量居当地较高水平。E-mail:
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2种不同金属材料的力学参数
科
Family
属数
Number of
genus
种数
Number of
species
占总种数比例
Percentage of
total species (%)
属
Genus
种数
Number of
species
占总种数比例
Percentage of total
species (%)
鹅膏菌科Amanitaceae 2 11 5.26 鹅膏菌属 Amanita 10 4.78
小菇科 Mycenaceae 2 12 5.74 丝盖伞属 Inocybe 5 2.39
多孔菌科 Polyporaceae 8 14 6.70 蜡蘑属 Laccaria 5 2.39
红菇科 Russulaceae 3 23 11.00 小皮伞属 Marasmius 6 2.87
小菇属 Mycena 11 5.26
光柄菇属 Pluteus 5 2.39
红菇属 Russula 17 8.13
栓菌属 Trametes 5 2.39
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